Human handedness: an ethological perspective

Human handedness is thought to be unique and universal. Critical examination of primary sources challenges both of these conclusions. Data on laterality of function in apes, humankind’s closest living relations, indicates hand preference and task specialization, but not yet handedness, as crucial studies remain to be done. Ethnographic data across cultures reveal more variance than is usually acknowledged, but existing findings are based on minimal and limited data. Ethological studies of spontaneous behavior in every day life are needed for both human and non-human primates.     

Right handedness of Homo heidelbergensis from Sima de los Huesos (Atapuerca, Spain) 500,000 years ago

Handedness is a product of brain specialization, which in turn seems to be responsible for the higher cognitive capabilities of humans, such as language and technology. Handedness in living humans is well established and shows the highest degree of manual specialization. Studies on hand laterality in nonhuman primates, particularly in chimpanzees, remain a matter of controversy as results tend to vary depending on factors such as the tasks performed and the environment in which the individuals live. Studies in several disciplines have attempted to determine where in the course of human evolution handedness established itself, with evidence collected from sources such as paleoneurological analyses, stone tool flaking, zooarchaeological studies and dental wear analyses, the last one of which have proven the most reliable source of information. Here we report an experimental and paleoanthropological study on hand laterality of a sample of 28 hominids from Sima de los Huesos (Atapuerca, Spain), dated at about 500,000 years ago, and compare our results with dental microwear analysis in other fossil samples such as that from Krapina (Croatia), as well as modern traditional societies. Our results indicate that European Middle Pleistocene Homo heidelbergensis was already as right-handed as modern populations.     

Body size and body shape in early hominins - implications of the Gona Pelvis

Discovery of the first complete Early Pleistocene hominin pelvis, Gona BSN49/P27, attributed to Homo erectus, raises a number of issues regarding early hominin body size and shape variation. Here, acetabular breadth, femoral head breadth, and body mass calculated from femoral head breadth are compared in 37 early hominin (6.0-0.26 Ma) specimens, including BSN49/P27. Acetabular and estimated femoral head sizes in the Gona specimen fall close to the means for non-Homo specimens (Orrorin tugenesis, Australopithecus africanus, Paranthropus robustus), and well below the ranges of all previously described Early and Middle Pleistocene Homo specimens. The Gona specimen has an estimated body mass of 33.2 kg, close to the mean for the non-Homo sample (34.1 kg, range 24-51.5 kg, n = 19) and far outside the range for any previously known Homo specimen (mean = 70.5 kg; range 52-82 kg, n = 17). Inclusion of the Gona specimen within H. erectus increases inferred sexual dimorphism in body mass in this taxon to a level greater than that observed here for any other hominin taxon, and increases variation in body mass within H. erectus females to a level much greater than that observed for any living primate species. This raises questions regarding the taxonomic attribution of the Gona specimen. When considered within the context of overall variation in body breadth among early hominins, the mediolaterally very wide Gona pelvis fits within the distribution of other lower latitude Early and Middle Pleistocene specimens, and below that of higher latitude specimens. Thus, ecogeographic variation in body breadth was present among earlier hominins as it is in living humans. The increased M-L pelvic breadth in all earlier hominins relative to modern humans is related to an increase in ellipticity of the birth canal, possibly as a result of a non-rotational birth mechanism that was common to both australopithecines and archaic Homo.     

Liang Bua Homo floresiensis mandibles and mandibular teeth: a contribution to the comparative morphology of a new hominin species

In 2004, a new hominin species, Homo floresiensis, was described from Late Pleistocene cave deposits at Liang Bua, Flores. H. floresiensis was remarkable for its small body-size, endocranial volume in the chimpanzee range, limb proportions and skeletal robusticity similar to Pliocene Australopithecus, and a skeletal morphology with a distinctive combination of symplesiomorphic, derived, and unique traits. Critics of H. floresiensis as a novel species have argued that the Pleistocene skeletons from Liang Bua either fall within the range of living Australomelanesians, exhibit the attributes of growth disorders found in modern humans, or a combination of both. Here we describe the morphology of the LB1, LB2, and LB6 mandibles and mandibular teeth from Liang Bua. Morphological and metrical comparisons of the mandibles demonstrate that they share a distinctive suite of traits that place them outside both the H. sapiens and H. erectus ranges of variation. While having the derived molar size of later Homo, the symphyseal, corpus, ramus, and premolar morphologies share similarities with both Australopithecus and early Homo. When the mandibles are considered with the existing evidence for cranial and postcranial anatomy, limb proportions, and the functional anatomy of the wrist and shoulder, they are in many respects closer to African early Homo or Australopithecus than to later Homo. Taken together, this evidence suggests that the ancestors of H. floresiensis left Africa before the evolution of H. erectus, as defined by the Dmanisi and East African evidence.     

Side steps: the erratic pattern of hominin postcranial change through time

The hominin fossil record reveals brain-size expansion, canine reduction, premolar metaconid development, and numerous other craniodental features that become more human-like through time. In general, the postcranial skeleton also gets more human-like through time, but in some respects it does not. This is particularly apparent in the overall morphology of one of the most frequently preserved elements, the distal humerus. Some of the earliest hominins display quite human-like morphologies, whereas later specimens are quite unusual among extant species of Hominoidea: when described metrically and subjected to multivariate discriminant analyses in the context of large samples of extant hominoid humeri, the shapes of the earliest hominin fossils are more human-like than many of the later specimens. The Mahalanobis distances between many of the 1.5-2Ma hominin humeri and Homo sapiens are remarkably large. Many of the less well-represented postcranial specimens do not follow a linear path through time of increasing hominization either. This is particularly noticeable in the fore-to-hind limb joint-size proportions, ulnar morphology, and pelvic architecture. The hominin postcranial fossil record reveals many side-steps: there appears to be no simple march toward our human bodies, but a pattern better explained as adaptations to proximate conditions and constrained by ontogeny and history.     

Pierolapithecus and the functional morphology of Miocene ape hand phalanges: paleobiological and evolutionary implications

The partial skeleton of Pierolapithecus, which provides the oldest unequivocal evidence of orthogrady, together with the recently described phalanges from Pa?alar most likely attributable to Griphopithecus, provide a unique opportunity for understanding the changes in hand anatomy during the pronogrady/orthogrady transition in hominoid evolution. In this paper, we describe the Pierolapithecus hand phalanges and compare their morphology and proportions with those of other Miocene apes in order to make paleobiological inferences about locomotor evolution. In particular, we investigate the orthograde/pronograde evolutionary transition in order to test whether the acquisition of vertical climbing and suspension were decoupled during evolution. Our results indicate that the manual phalanges of Miocene apes are much more similar to one another than to living apes. In particular, Miocene apes retain primitive features related to powerful-grasping palmigrady on the basal portion, the shaft, and the trochlea of the proximal phalanges. These features suggest that above-branch quadrupedalism, inherited from stem hominoids, constituted a significant component of the locomotor repertories of different hominoid lineages at least until the late Miocene. Nonetheless, despite their striking morphological similarities, several Miocene apes do significantly differ in phalangeal curvature and/or elongation. Hispanopithecus most clearly departs by displaying markedly-curved and elongated phalanges, similar to those in the most suspensory of the extant apes (hylobatids and orangutans). This feature agrees with several others that indicate orang-like suspensory capabilities. The remaining Miocene apes, on the contrary, display low to moderate phalangeal curvature, and short to moderately-elongated phalanges, which are indicative of the lack of suspensory adaptations. As such, the transition from a pronograde towards an orthograde body plan, as far as this particular anatomical region is concerned, is reflected only in somewhat more elongated phalanges, which may be functionally related to enhanced vertical-climbing capabilities. Our results therefore agree with the view that hominoid locomotor evolution largely took place in a mosaic fashion: just as taillessness antedated the acquisition of an orthograde body plan, the emergence of the latter—being apparently related only to vertical climbing—also preceded the acquisition of suspensory adaptations, as well as the loss of primitively-retained, palmigrady-related features.     

The comparative amino acid sequences, substrate specificities and gene or cDNA nucleotide sequences of some prokaryote and eukaryote amidinotransferases: implications for evolution

The amino acid sequences of the amidinotransferases and the nucleotide sequences of their genes or cDNA from four Streptomyces species (seven genes) and from the kidneys of rat, pig, human and human pancreas were compared. The overall amino acid and nucleotide sequences of the prokaryotes and eukaryotes were very similar and further, three regions were identified that were highly identical. Evidence is presented that there is virtually zero chance that the overall and high identity regions of the amino acid sequence similarities and the overall nucleotide sequence similarities between Streptomyces and mammals represent random match. Both rat and lamprey amidinotransferases were able to use inosamine phosphate, the amidine group acceptor of Streptomyces. We have concluded that the structure and function of the amidinotransferases and their genes has been highly conserved through evolution from prokaryotes to eukaryotes. The evolution has occurred with: (1) a high degree of retention of nucleotide and amino acid sequences; (2) a high degree of retention of the primitive Streptomyces guanine+cytosine (G+C) third codon position composition in certain high identity regions of the eukaryote cDNA; (3) a decrease in the specificities for the amidine group acceptors; and (4) most of the mutations silent in the regions suggested to code for active sites in the enzymes.     

The rise of the hominids as an adaptive shift in fallback foods: plant underground storage organs (USOs) and australopith origins

We propose that a key change in the evolution of hominids from the last common ancestor shared with chimpanzees was the substitution of plant underground storage organs (USOs) for herbaceous vegetation as fallback foods. Four kinds of evidence support this hypothesis: (1) dental and masticatory adaptations of hominids in comparison with the African apes; (2) changes in australopith dentition in the fossil record; (3) paleoecological evidence for the expansion of USO-rich habitats in the late Miocene; and (4) the co-occurrence of hominid fossils with root-eating rodents. We suggest that some of the patterning in the early hominid fossil record, such as the existence of gracile and robust australopiths, may be understood in reference to this adaptive shift in the use of fallback foods. Our hypothesis implicates fallback foods as a critical limiting factor with far-reaching evolutionary effects. This complements the more common focus on adaptations to preferred foods, such as fruit and meat, in hominid evolution.     

Dental microwear and diets of African early Homo

Conventional wisdom ties the origin and early evolution of the genus Homo to environmental changes that occurred near the end of the Pliocene. The basic idea is that changing habitats led to new diets emphasizing savanna resources, such as herd mammals or underground storage organs. Fossil teeth provide the most direct evidence available for evaluating this theory. In this paper, we present a comprehensive study of dental microwear in Plio-Pleistocene Homo from Africa. We examined all available cheek teeth from Ethiopia, Kenya, Tanzania, Malawi, and South Africa and found 18 that preserved antemortem microwear. Microwear features were measured and compared for these specimens and a baseline series of five extant primate species (Cebus apella, Gorilla gorilla, Lophocebus albigena, Pan troglodytes, and Papio ursinus) and two protohistoric human foraging groups (Aleut and Arikara) with documented differences in diet and subsistence strategies. Results confirmed that dental microwear reflects diet, such that hard-object specialists tend to have more large microwear pits, whereas tough food eaters usually have more striations and smaller microwear features. Early Homo specimens clustered with baseline groups that do not prefer fracture resistant foods. Still, Homo erectus and individuals from Swartkrans Member 1 had more small pits than Homo habilis and specimens from Sterkfontein Member 5C. These results suggest that none of the early Homo groups specialized on very hard or tough foods, but that H. erectus and Swartkrans Member 1 individuals ate, at least occasionally, more brittle or tough items than other fossil hominins studied.     

Carrying and hand-use patterns in Panamanian white-faced capuchins (Cebus capucinus)

The aim of this project was to examine the potential influence of postural regulation on capuchin hand-use patterns by focusing on tasks that involved the carrying of objects. Two months were spent on Barro Colorado Island, Panama, collecting data on ten white-faced capuchins (Cebus capucicus) (four adult males/six adult females). Over 215 contact hours were spent with the monkeys, and a total of 213 carries were recorded. No hand preference bias at the population level was found for the carry task; and no detectable hand-use patterns for carry were correlated with the weight of the object carried (except a right-hand bias for carrying medium-sized objects), the plane of the monkey's movement, its locomotor pattern, or its height above the ground. Therefore, the results of this study do not support the hypothesis that the need for postural regulation while carrying an object influences hand-use patterns in free-ranging capuchins. Our results are especially important because they are contrary to the results of the only other free-ranging capuchin laterality study conducted to date (Panger, 1998).     

The Belohdelie frontal: new evidence of early hominid cranial morphology from the Afar of Ethiopia

Study of the Belohdelie frontal has demonstrated that this four-million-year-old specimen belongs to a very generalized hominid that may be close to the divergence point of the hominid and African ape clades. Features associated with the temporalis muscle in the Belohdelie frontal and other new hominids from Hadar (AL 333-125) and West Turkana (KNM-ER 17000) suggest that the earliest hominids shared a large anterior component of this muscle relative to the extinct and extant apes. Results of this study support the phylogenetic hypothesis put forward by many workers that A. afarensis gave rise to the “robust” Australopithecus and A. africanus clades.     

Inferring hominoid and early hominid phylogeny using craniodental characters: the role of fossil taxa

Recent discoveries of new fossil hominid species have been accompanied by several phylogenetic hypotheses. All of these hypotheses are based on a consideration of hominid craniodental morphology. However, Collard and Wood (2000) suggested that cladograms derived from craniodental data are inconsistent with the prevailing hypothesis of ape phylogeny based on molecular data. The implication of their study is that craniodental characters are unreliable indicators of phylogeny in hominoids and fossil hominids but, notably, their analysis did not include extinct species. We report here on a cladistic analysis designed to test whether the inclusion of fossil taxa affects the ability of morphological characters to recover the molecular ape phylogeny. In the process of doing so, the study tests both Collard and Wood's (2000) hypothesis of character reliability, and the several recently proposed hypotheses of early hominid phylogeny. One hundred and ninety-eight craniodental characters were examined, including 109 traits that traditionally have been of interest in prior studies of hominoid and early hominid phylogeny, and 89 craniometric traits that represent size-corrected linear dimensions measured between standard cranial landmarks. The characters were partitioned into two data sets. One set contained all of the characters, and the other omitted the craniometric characters. Six parsimony analyses were performed; each data set was analyzed three times, once using an ingroup that consisted only of extant hominoids, a second time using an ingroup of extant hominoids and extinct early hominids, and a third time excluding Kenyanthropus platyops. Results suggest that the inclusion of fossil taxa can play a significant role in phylogenetic analysis. Analyses that examined only extant taxa produced most parsimonious cladograms that were inconsistent with the ape molecular tree. In contrast, analyses that included fossil hominids were consistent with that tree. This consistency refutes the basis for the hypothesis that craniodental characters are unreliable for reconstructing phylogenetic relationships. Regarding early hominids, the relationships of Sahelanthropus tchadensis and Ardipithecus ramidus were relatively unstable. However, there is tentative support for the hypotheses that S. tchadensis is the sister taxon of all other hominids. There is support for the hypothesis that A. anamensis is the sister taxon of all hominids except S. tchadensis and Ar. ramidus. There is no compelling support for the hypothesis that Kenyanthropus platyops shares especially close affinities with Homo rudolfensis. Rather, K. platyops is nested within the Homo + Paranthropus + Australopithecus africanus clade. If K. platyops is a valid species, these relationships suggest that Homo and Paranthropus are likely to have diverged from other hominids much earlier than previously supposed. There is no support for the hypothesis that A. garhi is either the sister taxon or direct ancestor of the genus Homo. Phylogenetic relationships indicate that Australopithecus is paraphyletic. Thus, A. anamensis and A. garhi should be allocated to new genera.     

Size- and locomotion-related aspects of hominid and anthropoid pelves: An osteometrical multivariate analysis

Two multivariate methods — the logarithmic principal component analysis (LPCA), and the logarithmic factorial analysis (LFA) — have been used tocompare the hip bone proportions of hominoids biometrically. The results have shown that size effects among apes and hominids interact to a centain extent with locomotor specializations, which are related to the attainment of more or less terrestrial behaviors. The pelvic morphology of great apes (Pongo, Pan, Gorilla) has retained numerous morphological traits — such as a gracile and elongated hip bone —, which were inherited from common adaptations to arboreal locomotion. In spite of these common traits, the African pongids (Pan, Gorilla) present two very different pelvic morphologies corresponding to two adaptative modes of terrestrial quadrupedalism. The hip bone of humans is proportionnally short and robust, most particularly at the level of its axial part. These characteristics, as well as the whole pelvic proportions, clearly indicate that gravitational forces exert a strong pressure on the pelvic walls during bipedalism. Among hominids, the transition from an australopithecine-like pelvic pattern to a human-like one corresponds to an increase of loading constraints on the hip jiont. This seems to indicate an evident change in locomotor behavior. Progression apparently became exclusively terrestrial with the genusHomo.     

Body size and its consequences: Allometry and the lower limb length of Liang Bua 1 (Homo floresiensis)

Bivariate femoral length allometry in recent humans, Pan, and Gorilla is investigated with special reference to the diminutive Liang Bua (LB) 1 specimen (the holotype of Homo floresiensis) and six early Pleistocene femora referred to the genus Homo. Relative to predicted body mass, Pan and Gorilla femora show strong negative length allometry while recent human femora evince isometry to positive allometry, depending on sample composition and line-fitting technique employed. The allometric trajectories of Pan and Homo show convergence near the small body size range of LB 1, such that LB 1 manifests a low percentage deviation (d_yx of Smith [1980]) from the Pan allometric trajectory and falls well within the 95% confidence limits around the Pan individuals (but also outside the 95% confidence limits for recent Homo). In contrast, the six early Pleistocene Homo femora, belonging to larger individuals, show much greater d_yx values from both Pan and Gorilla and fall well above the 95% confidence limits for these taxa. All but one of these Pleistocene Homo specimens falls within the 95% confidence limits of the recent human sample. Similar results are obtained when femoral length is regressed on femoral head diameter in unlogged bivariate space. Regardless of the ultimate taxonomic status of LB 1, these findings are consistent with a prediction made by us (Franciscus and Holliday, 1992) that hominins in the small body size range of A.L. 288-1 (“Lucy”), including members of the genus Homo, will tend to possess short, ape-like lower limbs as a function of body size scaling.     

Neanderthal Skeletal Structure and the Place of Homo neanderthalensis in European Hominid Phylogeny

Although the debate rages on over whether the Neanderthals merit their own species status or should be viewed as an odd variant of Homo sapiens, recent evidence has accumulated that overwhelmingly supports the former interpretation. Among this evidence is a recent full-body skeletal reconstruction that not only highlights the extreme differences between the highly apomorphic H. sapiens and H. neanderthalensis in the construction of the thorax and pelvic girdle, but strongly suggests significant gait differences between the two species that add to the probability that the two kinds of hominid would not have recognized each other as breeding partners. This is hardly surprising since the two species possessed a relatively remote common ancestry, and it is indeed suggested here that Homo neanderthalensis was merely one species embedded within a diverse and endemic middle Pleistocene European hominid radiation. Clearly more than one lineage of hominids simultaneously occupied Europe during the middle Pleistocene.     

The Laetoli footprints and early hominin locomotor kinematics

A critical question in human evolution is whether the earliest bipeds walked with a bent-hip, bent-knee gait or on more extended hindlimbs. The differences between these gaits are not trivial, because the adoption of either has important implications for the evolution of bipedalism. In this study, we re-examined the Laetoli footprints to determine whether they can provide information on the locomotor posture of early hominins. Previous researchers have suggested that the stride lengths of Laetoli hominins fall within the range of modern human stride lengths and therefore, Laetoli hominins walked with modern-human-like kinematics. Using a dynamic-similarity analysis, we compared Laetoli hominin stride lengths with those of both modern humans and chimpanzees. Our results indicate that Laetoli hominins could have used either a bent-hip, bent-knee gait, similar to a chimpanzee, or an extended-hindlimb gait, similar to a human. In fact, our data suggest that the Laetoli hominins could have walked near their preferred speeds using either limb posture. This result contrasts with most previous studies, which suggest relatively slow walking speeds for these early bipeds. Despite the many attempts to discern limb-joint kinematics from Laetoli stride lengths, our study concludes that stride lengths alone do not resolve the debate over early hominin locomotor postures.