Hominid cranial capacity versus time: A regression approach

The application of a linear regression approach to hominid data shows that there is more regularity in hominid cranial capacity over time than has been supposed. Two outcomes of this analysis are: (1) the possibility that the South African Australopithecus africanus sites are older than the presently accepted estimates of perhaps around 2·5 million years and (2) the suggestion that the Ngandong (Solo) cranial material is also considerably older than generally assumed; probably, in excess of approximately 250,000 years B.P.     

Paleoclimatic Variation and Brain Expansion during Human Evolution

One of the major adaptations during the evolution of Homo sapiens was an increase in brain size. Here we present evidence that a significant and substantial proportion of variation in brain size may be related to changes in temperature. Based on a sample of 109 fossilized hominid skulls, we found that cranial capacities were highly correlated with paleoclimatic changes in temperature, as indexed by oxygen isotope data and sea-surface temperature. Indeed, as much as 52% of the variance in the cranial capacity of these skulls could be accounted for by temperature variation at 100 ka intervals. As an index of more short-term seasonal fluctuations in temperature, we examined the latitude of the sites from which the crania originated. More than 22% of the variance in cranial capacity of these skulls could be accounted for by variation in equatorial distance.     

Cranial capacity in hominid evolution

We present an analysis of cranial capacity of 118 hominid crania available from the literature. The crania belong to both the genusAustralopithecus andHomo and provide a clear outline of hominid cranial evolution starting at more than 3 million years ago. Beginning withA. afarensis there is a clear increase in both absolute and relative brain size with every successive time period.H.s. neandertal has an absolutely and relatively smaller brain size (1412cc, E.Q.=5.6) than fossil modernH.s. sapiens (1487cc, E.Q.=5.9). Three evolutionary models of hominid brain evolution were tested: gradualism, punctuated equilibrium, and a mixed model using both gradualism and punctuated equilibrium. Both parametric and non-parametric analyses show a clear trend toward increasing brain size withH. erectus and a possible relationship within archaicH. sapiens. An evolutionary stasis in cranial capacity could not be refuted for all other taxa. Consequently, the mixed model appears to more fully explain hominid cranial capacity evolution. However, taxonomic decisions could directly compromise the possibility of testing the evolutionary mechanisms hypothesized to be operating in hominid brain expansion.     

Hominid brain size versus time: Revised regression estimates

This paper represents a revision of the regression estimates proposed in an earlier version (Journal of Human Evolution 1973 2, 405–411). An attempt has been made to incorporate new fossil material and the increasing number of absolute dates. The results of this new analysis suggest that the trend in the relationship of hominid cranial capacity with time remains consistent but the regression parameters have changed.     

A multivariate study of evolutionary change in the hominid cranial vault and some evolution rates

Multivariate analysis is used to describe the total morphological pattern of the hominid cranial vault, and to obtain distances between samples of Plio-Pleistocene Hominidae. Such techniques provide a means of quantifying phyletic change within a lineage, and therefore usefully complement the traditional Linnean nomenclature. When divided by elapsed time, the multivariate distances between groups provide a measure of the rate of evolution of a character complex and such data are given for the hominid cranial vault over the Quaternary as a whole, and for more detailed changes within Upper Pleistocene H. sapiens. The evolutionary significance of the observed rates, and their implications for the construction of phyletic schemes, are discussed.     

化石人类脑演化研究概况

脑演化是人类演化的一个重要组成部分,其研究可以为人类起源、演化、人群关系及语言、智力等方面提供重要的信息。脑演化的主要证据是通过研究颅内模（endocast）及颅骨的形态得到的。颅内模是从颅骨内表面得到的脑的外部形态。有时颅骨的内腔充满泥沙,并且为钙质所结固,可以自然形成颅内模。也可以人工制作颅内模。颅内模和颅骨的内表面能够提供人类脑髓及神经进化方面的直接证据。对化石人类脑演化的研究主要包括以下几个方面的内容：测量或估计脑量的大小及其和身体大小之间的关系;研究脑量随时间的变化过程;通过对早期人类颅内模表面沟回形态特征的研究,探索脑功能区在早期人类和猿类的区别及在演化上的变化;左右大脑不对称性与一侧优势关系,探讨语言的起源和惯用手的脑功能基础等;脑膜中动脉系统、静脉窦系统及与血液循环相关的排泄孔的变化,探讨大脑各部分比例的变化和功能的日益复杂对供血需求的影响;通过对人类脑演化的研究,探讨人类进化的原因。本文通过对以上几个方面及其中国化石脑演化研究的介绍,对化石人类脑演化的研究概况作综合论述和简单回顾。     

Hominid Brain Evolution

Hypotheses regarding the selective pressures driving the threefold increase in the size of the hominid brain since Homo habilis include climatic conditions, ecological demands, and social competition. We provide a multivariate analysis that enables the simultaneous assessment of variables representing each of these potential selective forces. Data were collated for latitude, prevalence of harmful parasites, mean annual temperature, and variation in annual temperature for the location of 175 hominid crania dating from 1.9 million to 10 thousand years ago. We also included a proxy for population density and two indexes of paleoclimatic variability for the time at which each cranium was discovered. Results revealed independent contributions of population density, variation in paleoclimate, and temperature variation to the prediction of change in hominid cranial capacity (CC). Although the effects of paleoclimatic variability and temperature variation provide support for climatic hypotheses, the proxy for population density predicted more unique variance in CC than all other variables. The pattern suggests multiple pressures drove hominid brain evolution and that the core selective force was social competition. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.

Predicting the Evolution of Sex on Complex Fitness Landscapes

Most population genetic theories on the evolution of sex or recombination are based on fairly restrictive assumptions about the nature of the underlying fitness landscapes. Here we use computer simulations to study the evolution of sex on fitness landscapes with different degrees of complexity and epistasis. We evaluate predictors of the evolution of sex, which are derived from the conditions established in the population genetic literature for the evolution of sex on simpler fitness landscapes. These predictors are based on quantities such as the variance of Hamming distance, mean fitness, additive genetic variance, and epistasis. We show that for complex fitness landscapes all the predictors generally perform poorly. Interestingly, while the simplest predictor, ΔVar_HD, also suffers from a lack of accuracy, it turns out to be the most robust across different types of fitness landscapes. ΔVar_HD is based on the change in Hamming distance variance induced by recombination and thus does not require individual fitness measurements. The presence of loci that are not under selection can, however, severely diminish predictor accuracy. Our study thus highlights the difficulty of establishing reliable criteria for the evolution of sex on complex fitness landscapes and illustrates the challenge for both theoretical and experimental research on the origin and maintenance of sexual reproduction.     

History of American paleoanthropological research on early Hominidae, 1925–1980

Understanding of the early stages of hominid evolution prior to 1925 was based primarily on comparative morphological evidence derived from extant primates. With the publication of Australopithecus by Dart in 1925 and subsequent research in South Africa, new possibilities for empirical assessment of early hominid evolutionary history were opened. It was Gregory's work, with Hellman, reported at the first meeting of the AAPA in 1930, that convinced many workers of the hominid status of Australopithecus. The debunking of Eoanthropus as a Pliocene hominid, far from having a totally negative effect, showed that cranial expansion had occurred after bipedalism in hominid evolution, demonstrated that chemical dating had come of age, and in a broader sense, had underlined that phylogenetic hypotheses are falsifiable by recourse to the evidence. The input of biological sciences into early hominid studies, as exemplified by Washburn's “new physical anthropology,” reduced taxonomic diversity and focused attention on paleoecology and behavior. The development of the multidisciplinary approach to field research, pioneered by L. Leakey and brought to fruition by Howell, was of fundamental importance in accurately dating and understanding the context of early hominids. Archaeology, primatology, comparative and functional morphology, and morphometrics have contributed substantially in recent years to a fuller understanding of early hominid evolution. American granting agencies have heavily supported early hominid research but patterns of funding have not kept pace with the change from research based largely on individualistic enterprise to multidisciplinary research projects. Future early hominid research, if funding is available, will likely be directed toward investigating temporal and geographic gaps now known in the fossil record and in more rigorous and multidisciplinary investigations of early hominid behavior.     

Variation in hominid brain size: how much is due to method?

Brain size represented by cranial capacity (CC) is one of the most frequently analysed characters of hominids. Accuracy of individual CC estimates depends on completeness of specimens and methods used for reconstruction and measurement. A file of published estimates of CC of hominids dated from 3.2 Ma (million years) to 10 Ka (thousand years) including 606 estimates for 243 specimens was compiled. In the file, 75 specimens are available with multiple values (3 to 15) obtained by various methods and/or by various authors. Using individuals as classes in ANOVA, intraclass variation, which represents "error" of estimates, was calculated. For the total sample of multiple estimates (N = 382) the error variance is 5315 ml2. The error standard deviation is 73 ml (coefficient of variation (CV = 7.3%), quite large in comparison to the actual variation in CC in modern humans, SD = 157 ml (CV = 11.6%). This large error makes us wonder whether any fossil can be reliably placed with respect to a particular "cerebral Rubicon" between palaeospecies. Recent discussions concerning cranial capacity of Stw505 are a case in point regarding errors in CC estimation. In actual repeated 30 time measurements on a research quality cast we obtained with various methods (water, seeds, plasticine) CC estimates ranging from 484 to 586 ml. The range of estimates in the literature is from 515 to 625 ml. When hominid CC by taxon with date as a covariate is subjected to ANOVA, taxon is responsible for 5% of the variance while date is responsible for the main portion, (89%). The relationship between CC and date is best characterised as a gradual time trend. It is proven by the ANOVA test for linearity, by gamma test for trend and by ASReml fitting of a linear function. The line of best fit to this time trend is a double exponential curve which explains 90% of the total variance in CC: CC = 306.63 (4.83(0.9995)DATE) Essentially the same curve fits subsamples of CC dated at less than 1 Ma and at 3.2-1.0 Ma. This has several implications for the nature of the Darwinian process to be reconstructed.     

Variability selection in hominid evolution

Variability selection (abbreviated as VS) is a process considered to link adaptive change to large degrees of environment variability. Its application to hominid evolution is based, in part, on the pronounced rise in environmental remodeling that took place over the past several million years. The VS hypothesis differs from prior views of hominid evolution, which stress the consistent selective effects associated with specific habitats or directional trends (e.g., woodland, savanna expansion, cooling). According to the VS hypothesis, wide fluctuations over time created a growing disparity in adaptive conditions. Inconsistency in selection eventually caused habitat-specific adaptations to be replaced by structures and behaviors responsive to complex environmental change. Key hominid adaptations, in fact, emerged during times of heightened variability. Early bipedality, encephalized brains, and complex human sociality appear to signify a sequence of VS adaptations—i.e., a ratcheting up of versatility and responsiveness to novel environments experienced over the past 6 million years. The adaptive results of VS cannot be extrapolated from selection within a single environmental shift or relatively stable habitat. If some complex traits indeed require disparities in adaptive setting (and relative fitness) in order to evolve, the VS idea counters the prevailing view that adaptive change necessitates long-term, directional consistency in selection. © 1998 Wiley-Liss, Inc.     

The enigma of the Sangiran 4 palate revisited

The Sangiran 4 palate has been controversial since its discovery in the 1930s because it retains a number of more primitive morphologies such as projecting canines and precanine diastemata. These characters have led some workers to question the hominid status of the palate, suggesting that it is both too large and too primitive to belong to the same individual as the Sangiran 4 cranial fragments. The palate has instead been diagnosed as a new species of Pongo. The present study re-evaluates this controversy through the analysis of new metric data and comparisons with more recently published fossil discoveries. This analysis shows that the Sangiran 4 palate is not unique, and shares several of these putative pongid traits with other Javan hominid fossils as well as recently described hominid specimens from Dmanisi, Georgia. These results suggest that the evolution of the earliest Asians was more complex than has previously been appreciated.     

Cranial thickening in an Australian hominid as a possible palaeoepidemiological indicator

This paper describes the cranial thickening of a late Pleistocene hominid (Willandra Lakes Hominid 50) from Australia. The unusual development of the vault structures in this individual has few, if any, equals among other hominids or more recent populations from around the world. The vault morphology is, therefore, described in terms of a pathologically related condition associated with the modern haemolytic blood dyscrasias, typical of sickle cell anamia and thalassemia. A possible palaeoepidemiology for these genetic adaptations among early Australasian populations is proposed together with a discussion of similar changes observed in the vault of the Singa calvarium from the Sudan. It is tentatively suggested that the cranial thickening of the Australian hominid has its origins in some form of genetic blood disease and that if this diagnosis is correct, this individual provides a rare glimpse of human biological adaptation in the late Upper Pleistocene.     

The dating of Lantian man and his significance for analyzing trends in human evolution

The Lantian fossil hominid cranium from Southern Shensi Province, China, provides the earliest record of Homo erectus in northern east Asia, and is morphologically the most primitive specimen in the entire world. Importantly, the Kungwangling Lantian cranium (calvarium plus face), with associated stone tools in good geologic and paleontological context, is demonstrably both earlier and more primitive than the Choukoutien I remains. Faunal and palynological evidence support a mid-Mosbachium equivalent age (some 700,000 years). These facts are not recognized in the original Chinese reports. The Chenchiawo Lantian mandible, like the Choukoutien I remains, is attributable to the Holstein-equivalent in China (some 300,000 years ago), and therefore should no longer be temporally associated with the Kungwangling Lantian cranium. However, that the mandible may be morphologically associated with either calls attention to the relative independence of the mandible in human evolution. A comparative study of some modern Mongoloid populations in which very large mandibles may or may not be associated with a scaphoid keel or sagittal elevation depending upon the size and shape of the cranium demonstrates the relative autonomy of the mandible. Continuing selection pressure for a masticatory complex with large jaws provides another point of continuity between East Asian fossil and modern Mongoloid hunting populations such as Eskimos and Aleuts. A number of morphological features of the cranium, especially vault thickness, cranial capacity and reinforcement system, conform to expectation and confirm a general trend of reduction in vault thickness and reinforcement system with increase in cranial capacity over time within the single human species.     

Cranial vault shape in fossil hominids: Fourier descriptors in norma lateralis

Two major views of human evolution have elicited considerable controversy. These are: [1] the “out of Africa” hypothesis and [2] the “multiregional” hypothesis. This paper is an attempt to try to reconcile these two scenarios using hominid cranial vault data. Elliptical Fourier functions (EFFs) were used to describe, in visual and numerical terms, the shape of the human cranial vault in norma lateralis.Using jpeg images, contours of the cranial vault of a large sample of hominid specimens were pre-processed in Photoshop CS and rotated in 2D space (positional-orientation) so that a line drawn from nasion to porion was horizontal. The cranial vault image was then digitized with 72 closely-spaced points and submitted to a specially written routine that computed EFFs normalized by scaling (size-standardization). This ensured that the representation was invariant with respect to starting point, size and orientation.Statistically significant differences were found between the H. sapiens sample and both the H. erectus and H. neanderthalensis samples. In contrast, there were no statistically significant differences between the H. erectus and H. neanderthalensis groups, leading to three conclusions: [1] the similarity in cranial vault shape between H. erectus and H. neanderthalensis suggests a single gradually evolving lineage; [2] The taxon H. heidelbergensis can be embedded into the H. erectus → H. neanderthalensis line; and [3] H. sapiens seems to be a separate evolutionary development and is considered here either as a separate species or as a possible example of an allopatric semispecies (Grant, 1977). The results here suggest that human evolution over the last 2 Ma may turn out to be neither totally multiregional or simply out of Africa but rather represents a considerably more complicated picture.     

Evolution of longevity in primates

Evolution of maximum lifespan potential (MLP) and its possible relation to the evolution of extra-brain functions are investigated in the primates. MLP is related to a species' characteristic aging rate and is considered a basic biological property of an organism. MLP may be of more importance than realized in the past in determining the evolutionary success of a species. It is related to the postnatal development rate, the length of time general vigor is maintained, the length of reproductive period, generation time and the time available for learning and teaching behavior. Three other parameters are considered to be importantly related to MLP: MLP calorie consumption (MCC), encephalization quotient (EQ) and extra number of cortical neurons (N_c). MCC is calculated as the product of MLP and specific metabolic rate (SMR) and is considered to represent total “life-capacity” of an organism. It is of potential value in studying the biological mechanisms involved in the evolution of MLP. Brain function is estimated by the Jerison EQ and N_c parameters. These parameters estimate the “extra” brain capacity involved in functions beyond normal body requirements. The rate of change in MLP and N_c per unit time occurring during an ancestral-descendant sequence is used to estimate the biological complexity of the genetic processes which have evolved in governing the rate of expression of the general aging process and increasing brain function. The average rate of change of MLP during the emergence of the primates was analyzed by the difference between MLP in closely related living primate species and the evolutionary time of appearance of a common ancestor. MLP, SMR, MCC, EQ and N_c were estimated from fossil cranial capacity and body weight measurements. The rate of change in these values was calculated according to the time of appearance of the fossil species. MLP and N_c were found to increase together and reached their highest rate of increase approximately 200,000 years ago along the hominid ancestral-descendant sequence leading to modern man. The high rate of increase of these parameters suggests that few genetic changes were responsible. The general increase in MLP during the evolution of the primate species indicates that a corresponding general decrease in mutation rate may have occurred. The high levels of MLP, MCC, EQ and N_c represented in the living primates, as compared to other mammals, are considered to represent a major characteristic determining their evolutionary success.     

Variance matters: The shape of a datum

In the quantitative analysis of behaviour, choice data are most often plotted and analyzed as logarithmic transforms of ratios of responses and of ratios of reinforcers according to the generalized-matching relation, or its derivatives such as conditional-discrimination models. The relation between log choice ratios and log reinforcer ratios has normally been found using ordinary linear regression, which minimizes the sums of the squares of the y deviations from the fitted line. However, linear regression of this type requires that the log choice data be normally distributed, of equal variance for each log reinforcer ratio, and that the x (log reinforcer ratio) measures be fixed with no variance. We argue that, while log transformed choice data may be normally distributed, log reinforcer ratios do have variance, and because these measures derive from a binomial process, log reinforcer ratio distributions will be non-normal and skewed to more extreme values. These effects result in ordinary linear regression systematically underestimating generalized-matching sensitivity values, and in faulty parameter estimates from non-linear regression to assume hyperbolic and exponential decay processes. They also lead to model comparisons, which assume equal normally distributed error around every data point, being incorrect. We describe an alternative approach that can be used if the variance in choice is measured.     

Paranthropus boisei: An example of evolutionary stasis?

Of the presently recognised early hominid species, Paranthropus boisei is one of the better known from the fossil record and arguably the most distinctive; the latter interpretation rests on the numbers of apparently derived characters it incorporates. The species as traditionally diagnosed is distributed across approximately one million years and is presently confined to samples from East African sites. The hypodigm has been examined for evidence of intraspecific phyletic evolution, with particular emphasis on the areas best represented in the fossil record, namely the teeth and mandible. The results of this examination of 55 mandibular and dental variables, which uses the Γ test statistic for the detection of trend, and nonparametric spline regression (Loess regression) for investigating pattern and rate of temporal change, show that within Paranthropus boisei sensu stricto most evidence of temporally related morphological trends relates to the morphology of the P₄ crown. There is little or no evidence of any tendency to increase in overall size through time. Fossils from the Omo Shungura Formation and West Turkana which have been recovered from a time period earlier than the P. boisei sensu stricto hypodigm resemble the latter taxon in some features, but differ from it in aspects of cranial morphology. There is insufficient fossil evidence of the earlier taxon to tell whether it changes through time, but when trends of 47 mandibular and dental variables are assessed across the combined East African “robust” australopithecine sample, there is evidence for a relatively abrupt change around 2.2–2.3 Myr in approximately 25% of the dental and mandibular remains. Some of these changes correspond with the temporal trends within P. boisei sensu stricto, but others, such as mandible height, do not. If the earlier material is ancestral to P. boisei sensu stricto, evidence from the teeth and jaws is consistent with a punctuated origin for the latter taxon. © 1994 Wiley-Liss, Inc.