The evolution of early Homo: a reply to Scott.

Scott presents a welcome reply to our article, "A single lineage in early Pleistocene Homo" (Van Arsdale and Wolpoff ). However, Scott's reply mischaracterizes and fails to directly address the hypothesis of a single lineage that we test. Additionally, the approach taken by Scott fails to replicate the methods used in our analysis. As Scott himself suggests, our null hypothesis of a single evolving lineage in early Homo remains without refutation. Although many evolutionary scenarios might explain the complex pattern of variation present in the early Homo fossil record, the most parsimonious remains that of a single lineage displaying evolutionary change over time.

The accretion model of Neandertal evolution.

The Accretion model of Neandertal evolution specifies that this group of Late Pleistocene hominids evolved in partial or complete genetic isolation from the rest of humanity through the gradual accumulation of distinctive morphological traits in European populations. As they became more common, these traits also became less variable, according to those workers who developed the model. Its supporters propose that genetic drift caused this evolution, resulting from an initial small European population size and either complete isolation or drastic reduction in gene flow between this deme and contemporary human populations elsewhere. Here, we test an evolutionary model of gene flow between regions against fossil data from the European population of the Middle and Late Pleistocene. The results of the analysis clearly show that the European population was not significantly divergent from its contemporaries, even in a subset of traits chosen to show the maximum differences between Europeans and other populations. The pattern of changes, over time within Europe of the traits in this subset, does not support the Accretion model, either because the characters did not change in the manner specified by the model or because the characters did not change at all. From these data, we can conclude that special phenomena such as near-complete isolation of the European population during the Pleistocene are not required to explain the pattern of evolution in this region.

Brief communication: paleoanthropology and the population genetics of ancient genes.

The Mezmaiskaya cave mtDNA is similar in many ways to the Feldhofer cave Neandertal sequence and the more recently obtained Vindija cave sequence. If we accept the contention that the Mezmaiskaya cave specimen is a Neandertal infant, its mtDNA provides no new information about the fate of the European Neandertals. However, there is reason to believe that the Mezmaiskaya cave infant is not a Neandertal, and this places its importance in another light, because it delimits the possible hypotheses of Neandertal and recent human genetic relationships. One possibility is a that the pattern found in ancient mtDNA results from the replacement of an isolated gene pool (Neandertals) by one of its contemporaries (modern humans). A second possibility is natural selection expressed as the substitution of an advantageous mtDNA variant within a single large species, including both Neandertals and modern humans. The geologic, archaeological, and dating evidence shows the Mezmaiskaya cave infant to be a burial from a level even more recent than the Upper Paleolithic preserved at the site, and its anatomy does not contradict the assessment that the Mezmaiskaya cave infant is not a Neandertal. Therefore, the second pattern can be favored over the first.

Modern human ancestry at the peripheries: a test of the replacement theory.

The replacement theory of modern human origins stipulates that populations outside of Africa were replaced by a new African species of modern humans. Here we test the replacement theory in two peripheral areas far from Africa by examining the ancestry of early modern Australians and Central Europeans. Analysis of pairwise differences was used to determine if dual ancestry in local archaic populations and earlier modern populations from the Levant and/or Africa could be rejected. The data imply that both have a dual ancestry. The diversity of recent humans cannot result exclusively from a single Late Pleistocene dispersal.

An Australasian test of the recent African origin theory using the WLH-50 calvarium.

This analysis investigates the ancestry of a single modern human specimen from Australia, WLH-50 (Thorne et al., in preparation; Webb, 1989). Evaluating its ancestry is important to our understanding of modern human origins in Australasia because the prevailing models of human origins make different predictions for the ancestry of this specimen, and others like it. Some authors believe in the validity of a complete replacement theory and propose that modern humans in Australasia descended solely from earlier modern human populations found in Late Pleistocene Africa and the Levant. These ancestral modern populations are believed to have completely replaced other archaic human populations, including the Ngandong hominids of Indonesia. According to this recent African origin theory, the archaic humans from Indonesia are classified as Homo erectus, a different evolutionary species that could not have contributed to the ancestry of modern Australasians. Therefore this theory of complete replacement makes clear predictions concerning the ancestry of the specimen WLH-50. We tested these predictions using two methods: a discriminant analysis of metric data for three samples that are potential ancestors of WLH-50 (Ngandong, Late Pleistocene Africans, Levant hominids from Skhul and Qafzeh) and a pairwise difference analysis of nonmetric data for individuals within these samples. The results of these procedures provide an unambiguous refutation of a model of complete replacement within this region, and indicate that the Ngandong hominids or a population like them may have contributed significantly to the ancestry of WLH-50. We therefore contend that Ngandong hominids should be classified within the evolutionary species, Homo sapiens. The Multiregional model of human evolution has the expectation that Australasian ancestry is in all three of the potentially ancestral groups and best explains modern Australasian origins.

Multiregional, not multiple origins.

Multiregional evolution is a model to account for the pattern of human evolution in the Pleistocene. The underlying hypothesis is that a worldwide network of genic exchanges, between evolving human populations that continually divide and reticulate, provides a frame of population interconnections that allows both species-wide evolutionary change and local distinctions and differentiation. "Multiregional" does not mean independent multiple origins, ancient divergence of modern populations, simultaneous appearance of adaptive characters in different regions, or parallel evolution. A valid understanding of multiregional evolution would go a long way toward reducing the modern human origins controversy.

Zuttiyeh face: a view from the East.

We analyze the phylogenetic position of the frontofacial fragment from Zuttiyeh, Israel. This specimen is dated to the Middle Pleistocene (the latest estimate is between 250 and 350 kyr) and is associated with the Acheulo-Yabrudian, which makes it the oldest cranium from the region. It has been previously regarded as a Neandertal, and early "anatomically modern Homo sapiens," and a generalized specimen ancestral to both. These different phylogenetic interpretations of its features have a historic basis but in our view also result from a confusion of grade and intraspecies clade as valid sources of variation. We show here that generally the differences that distinguish Zuttiyeh from Neandertals are similarities it shares with the Zhoukoudian remains. These similarities involve a unique combination of features, and suggest the possibility of an ancestral relationship. It is less likely that Middle Pleistocene remains from Europe or sub-Saharan Africa are uniquely or significantly ancestral to Zuttiyeh. An accurate understanding of the relationship between populations of eastern and western Asia is important for resolving the more general questions surrounding the position of the Upper Pleistocene Levant populations in human evolution, including (1) whether there are significantly different contemporary Mousterian populations in the Upper Pleistocene, (2) whether Neandertals are clearly intrusive in the region, and (3) whether there is an early appearance of (what many workers call) "anatomically modern Homo sapiens." The hypothesis of a recent unique African ancestry for all modern humans is disproved by our study, which shows Asia as a significant source area for at least some living populations.

Lucy's lower limbs: long enough for Lucy to be fully bipedal?

The recent attempt to show that the Hadar australopithecine female 'Lucy' (AL 288-1) had hindlimbs too short to allow a modern pattern of striding bipedal gait has important implications for understanding the origin of bipedalism, if not for the more general problem of hominid origins. Combined with previous claims that Lucy had a forelimb unusually long in proportion and ape-like in morphology, the additional contention of a relatively short hindlimb would suggest a very different pattern of gait from the norm of today because the effectiveness of the pendulum action of the lower limb during stride is a function of the amount of mass in the limb, and because a short hindlimb would necessitate a short stride length. Yet, these contentions seem contradicted by the analyses of Lucy's pelvis (and the innominates of other australopithecines) that indicate a similar pattern of muscle use and imply a lack of significant gait differences. Are Lucy's legs too short to allow an effective stride, or is there a different solution to this contradiction? I propose here that there is.

Regional continuity in Australasian Pleistocene hominid evolution.

A reconstruction of the Indonesian male Homo erectus specimen Sangiran 17 reveals a suite of features that suggests regional morphological continuity in Australasia during the middle and late Pleistocene. Characteristics reflecting a morphological clade are established through a comparison with the males from Kow Swamp, the late Pleistocene Australian site with the largest number of hominid specimens. It is argued that the "center and edge" hypothesis could account for the initial appearance of regional variation, and that a dynamic model of geneflow and opposing selection provides the mechanism that maintained a long-standing dynamic clinal equilibrium in this area.

Anterior dental cutting at Laetolil.

No evidence supports the interpretation of malocclusion in the LH-14 right P3. The pattern of anterior cutting shown in the younger laetolil specimens has its analogue in the deciduous dentitions of Pongo.