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Интересные факты по эволюции гоминид, установленные в последние годы (1997-2004).

дополнение к обзору "Происхождение и эволюция человека"


Охотники или падальщики?

Кем были наши предки - охотниками или падальщиками? Этот спор продолжается уже много десятилетий. В целом общее мнение сейчас склоняется к тому, что австралопитеки и хабилисы были преимущественно падальщиками, гейдельбержцы и неандертальцы - охотниками, по поводу же питекантропов (H.ergaster, H.erectus, H.georgicus) мнения сильно расходятся.

Неандертальцы и сапиенсы

Эволюция двуногости

Систематика, филогения




1.  Br J Nutr. 1998 Jan;79(1):3-21.

Rift Valley lake fish and shellfish provided brain-specific nutrition for early Homo.

Broadhurst CL, Cunnane SC, Crawford MA.

An abundant, balanced dietary intake of long-chain polyunsaturated fatty acids is an absolute requirement for sustaining the very rapid expansion of the hominid cerebral cortex during the last one to two million years. The brain contains 600 g lipid/kg, with a long-chain polyunsaturated fatty acid profile containing approximately equal proportions of arachidonic acid and docosahexaenoic acid. Long-chain polyunsaturated fatty acid deficiency at any stage of fetal and/or infant development can result in irreversible failure to accomplish specific components of brain growth. For the past fifteen million years, the East African Rift Valley has been a unique geological environment which contains many enormous freshwater lakes. Paleoanthropological evidence clearly indicates that hominids evolved in East Africa, and that early Homo inhabited the Rift Valley lake shores. Although earlier hominid species migrated to Eurasia, modern Homo sapiens is believed to have originated in Africa between 100 and 200 thousand years ago, and subsequently migrated throughout the world. A shift in the hominid resource base towards more high-quality foods occurred approximately two million years ago; this was accompanied by an increase in relative brain size and a shift towards modern patterns of fetal and infant development. There is evidence for both meat and fish scavenging, although sophisticated tool industries and organized hunting had not yet developed. The earliest occurrences of modern H. sapiens and sophisticated tool technology are associated with aquatic resource bases. Tropical freshwater fish and shellfish have long-chain polyunsaturated lipid ratios more similar to that of the human brain than any other food source known. Consistent consumption of lacustrine foods could have provided a means of initiating and sustaining cerebral cortex growth without an attendant increase in body mass. A modest intake of fish and shellfish (6-12% total dietary energy intake) can provide more arachidonic acid and especially more docosahexaenoic acid than most diets contain today. Hence, 'brain-specific' nutrition had and still has significant potential to affect hominid brain evolution.


2. J Hum Evol. 2000 Apr;38(4):497-521.

Adults only. Reindeer hunting at the middle palaeolithic site salzgitter lebenstedt, northern Germany.

Gaudzinski S, Roebroeks W.

The Middle Palaeolithic site Salzgitter Lebenstedt (northern Germany), excavated in 1952, is well known because of its well-preserved faunal remains, dominated by adult reindeer (Rangifer tarandus). The archaeological assemblage accumulated in an arctic setting in an earlier part of the last (Weichsel) glacial (OIS5-3). The site is remarkable because of the presence of unique Middle Palaeolithic bone tools and the occurrence of the northernmost Neanderthal remains, but this paper focuses on an analysis of its reindeer assemblage. The results indicate autumn hunting of reindeer by Middle Palaeolithic hominids. After the hunt, carcasses were butchered and in subsequent marrow processing of the bones a selection against young and sub-adult animals occurred. Adults were clearly preferred, and from their bones, again, poorer marrow bones were neglected. This focus on primeness of resources has been documented in other domains of Neanderthal behaviour, but Salzgitter Lebenstedt is the best example yet known in terms of systematic and routinized processing of game. The Salzgitter Lebenstedt assemblage displays some remarkable similarities to the Late Glacial reindeer assemblages from the Ahrensburg tunnel valley sites. The subsequent review of the evidence on subsistence strategies from earlier periods of the European Palaeolithic shows that hunting of large mammals may have been a part of the behavioural repertoire of the Middle Pleistocene occupants of Europe from the earliest occupation onwards. At the same time, it is suggested that these early hunting strategies were incorporated in ways of moving through landscapes ("settlement systems") which were different from what we know from the middle parts of the Upper Palaeolithic onwards. Copyright 2000 Academic Press.


3. J Hum Evol. 1998 Aug;35(2):111-36. 

A critique of the evidence for scavenging by Neanderthals and early modern humans: new data from Kobeh Cave (Zagros Mountains, Iran) and Die Kelders Cave 1 layer 10 (South Africa).

Marean CW.

The primary mode of faunal exploitation by Neandertals and early modern humans remains a debated topic. Binford (1981, 1984, 1985, 1988) has argued for an obligate scavenging mode, Stiner (1991a, 1991b, 1991c, 1993, 1994) for a more opportunistic scavenging mode, while other researchers (Chase, 1986, 1988, 1989; Klein, 1989, 1994, 1995; Klein & Cruz-Uribe, 1996) deny the importance of scavenging as a faunal exploitation tactic. The scavenging interpretations rely primarily on several patterns in the faunal remains: the presence of a skeletal element pattern dominated by heads or head and foot parts, the presence of carnivore tooth marks on bone fragments, and infrequent cut marks that typically are not located on shaft regions of long bones or on fleshy bones. Five sites have been used to argue for scavenging: Klasies River Mouth, Combe Grenal, Grotta Guattari, Grotta dei Moscerini, and Grotte Vaufrey. The former four of the five sites are biased samples in that long bone shafts and other difficult to identify fragments were discarded at excavation. The analysis of Grotte Vaufrey included only those shafts identifiable to species or genus, thus excluding the vast majority of shaft specimens. This bias systematically shapes the skeletal element and surface modification patterning in ways that make the assemblages appear to fit a model of scavenging, when in fact the main determinant of the pattern is the bias in the flawed samples. This problem is illustrated with two unbiased faunal assemblages (Kobeh Cave and Die Kelders Layer 10). Skeletal element abundance is calculated in a way that mimics the bias in the sites listed above by excluding the shafts. Using this procedure, both Kobeh and Die Kelders have a head and foot skeletal element pattern and thus appear scavenged. Both assemblages are then analyzed in their entirety and a new pattern, consistent with hunting, is revealed. Taphonomic data on bone survival and destruction provide an explanation for this result. Excluding shaft fragments from the analysis also biases the surface modification patterning in such a way as to produce a pattern more consistent with scavenging. The conclusion is that there is no reliable evidence for scavenging by Neandertals or early modern humans.

4.   Published online before print June 13, 2000, 10.1073/pnas.120178997
Proc Natl Acad Sci U S A. 2000 June 20; 97 (13): 7663–7666

Neanderthal diet at Vindija and Neanderthal predation: The evidence from stable isotopes

Michael P. Richards,*† Paul B. Pettitt,*‡ Erik Trinkaus,§¶|| Fred H. Smith,** Maja Paunović ,‡‡ and Ivor Karavanić ††


* Research Laboratory for Archaeology and the History of Art, University of Oxford, 6 Keble Road, Oxford OX1 3QJ, United Kingdom; Department of Archaeology, Simon Fraser University, Burnaby, BC V5A 1S6, Canada; Keble College, Oxford OX1 3PG, United Kingdom; § Department of Anthropology, Campus Box 11, Washington University, St. Louis MO 63130; Unité Mixte de Recherche 5809 du Centre National de la Recherche Scientifique, Laboratoire d'Anthropologie, Université de Bordeaux I, 33405 Talence, France; ** Department of Anthropology, Northern Illinois University, DeKalb, IL 60115; ‡‡ Zavod za paleontologiju i geologiju kvartara, Hrvatske akademije znanosti i umjetnosti, Ulica A. Kovačića 5/II, HR-10000 Zagreb, Croatia; and †† Arheološki zavod Filozofskog fakulteta Sveučilišta u Zagrebu, I. Lučića 3, HR-10000 Zagreb, Croatia


Contributed by Erik Trinkaus, April 19, 2000


|| To whom reprint requests should be addressed. E-mail:


This article has been cited by other articles in PMC.



Archeological analysis of faunal remains and of lithic and bone tools has suggested that hunting of medium to large mammals was a major element of Neanderthal subsistence. Plant foods are almost invisible in the archeological record, and it is impossible to estimate accurately their dietary importance. However, stable isotope (δ13C and δ15N) analysis of mammal bone collagen provides a direct measure of diet and has been applied to two Neanderthals and various faunal species from Vindija Cave, Croatia. The isotope evidence overwhelmingly points to the Neanderthals behaving as top-level carnivores, obtaining almost all of their dietary protein from animal sources. Earlier Neanderthals in France and Belgium have yielded similar results, and a pattern of European Neanderthal adaptation as carnivores is emerging. These data reinforce current taphonomic assessments of associated faunal elements and make it unlikely that the Neanderthals were acquiring animal protein principally through scavenging. Instead, these findings portray them as effective predators.

paleodiet | Croatia | Europe | δ13C | δ15N



Reconstructions of European Neanderthal subsistence strategies have overwhelmingly focused on the specialized hunting and scavenging of herbivores as the predominant method of obtaining food (16). These reconstructions are based principally on the analysis of the abundantly preserved faunal remains, supplemented by artifactual evidence of lithic and wood hunting apparatuses, as well as on the relative importance of the faunal biomass in the environments that European Neanderthals occupied during later oxygen isotope stage 5 and especially oxygen isotope stages 4 and 3 of the Late Pleistocene. Understanding Neanderthal diet has implications for understanding Neanderthal land use, social organization, and behavioral complexity. Yet despite the abundant evidence for successful hunting techniques across Neanderthal Eurasia, faunal remains can indicate only hunting or scavenging episodes; they cannot tell us about the predominant foods in the diet over the long term.

By contrast, the measurement of the ratios of the stable isotopes of carbon and nitrogen in mammal bone collagen provides an indication of aspects of diet over the last few years of life (79). This stable isotope evidence can therefore provide us with direct information on Neanderthal diet. This method has been applied to Neanderthal remains from the sites of Marillac, France (10), and Scladina Cave, Belgium (11). These studies, focusing particularly on their high δ15N values, indicated that the Neanderthals measured occupied the top trophic level, obtaining nearly all of their dietary protein from animal sources. In the context of this finding, we undertook stable isotope analyses of the two late Neanderthal specimens from Vindija Cave, in the Hrvatsko Zagorje of northern Croatia [Vi-207 and Vi-208 (12)], and of the fauna with which they were stratigraphically associated.

Vindija Neanderthal and Faunal Specimens. Recently, the Vi-207 and Vi-208 Neanderthal specimens, as well as various other archeological materials from level G1 of Vindija Cave, Croatia, were submitted for accelerator mass spectrometer radiocarbon dating at the Oxford Radiocarbon Accelerator Unit, University of Oxford (13). The two Neanderthal specimens were dated to 29,080 ± 400 years before present (B.P.) (OxA-8296, Vi-207) and 28,020 ± 360 years B.P. (OxA-8295, Vi-208), making them the youngest directly dated Neanderthal specimens in Europe (13). Because the radiocarbon sample preparation process includes assessments of stable isotopes, in part to control for potential contamination, this analysis also yielded stable isotope profiles for these late archaic humans. Combined with similar data obtained from faunal remains from level G1 and the older level G3 of Vindija Cave, this provides a means of assessing the dietary profiles of these Neanderthals.

Stable Isotope Analyses. Mammal bone collagen δ13C and δ15N values reflect the δ13C and δ15N values of dietary protein (14). They furnish a long-term record of diet, giving the average δ13C and δ15N values of all of the protein consumed over the last years of the measured individual's life. δ13C values can be used to discriminate between terrestrial and marine dietary protein in humans and other mammals (15, 16). In addition, because of the canopy effect, species that live in forest environments can have δ13C values that are more negative than species that live in open environments (17). δ15N values are, on average, 2–4‰ higher than the average δ15N value of the protein consumed (18). Therefore, δ15N values can be used to determine the trophic level of the protein consumed. By measuring the δ13C and δ15N values of various fauna in a paleo-ecosystem, it is possible to reconstruct the trophic level relationships within that ecosystem. Therefore, by comparing the δ13C and δ15N values of omnivores such as hominids with the values of herbivores and carnivores from the same ecosystem, it is possible to determine whether those omnivores were obtaining dietary protein from plant or animal sources.

Vindija Neanderthal and Faunal Isotope Values. Collagen was extracted from the two Neanderthal specimens from level G1 of Vindija Cave and from various faunal remains from level G1 and the older level G3 according to standard collagen extraction procedures; the Neanderthal specimens were extracted according to the methods outlined in Law and Hedges (19), and the faunal specimens were extracted according to the procedure outlined in Richards and Hedges (16). The collagen extracts varied in quality, and only those samples that had acceptable collagen attributes were used. These attributes are based on values determined by DeNiro (20) and Ambrose (21) and used by the majority of stable isotope researchers and radiocarbon dating labs. The acceptable values are a C:N ratio between 2.9 and 3.6, “percent collagen” >1%, and %C and %N in the extracted collagen of >13% for carbon and >5% for nitrogen. These collagen attributes allow us to identify and exclude collagen that is heavily degraded or contaminated. This is in contrast to stable isotope measurements of bioapatite in bone mineral and enamel, where no such criteria exist. The stable isotope values and various collagen attributes are given in Table 1; based on these, we are confident that the collagen δ13C and δ15N values reported here are robust and reflect the organisms' original collagen δ13C and δ15N values. The Neanderthal samples were measured at the Oxford Radiocarbon Accelerator Unit, and the faunal samples were measured at the Stable Isotope Laboratory, Research Laboratory for Archaeology and the History of Art, University of Oxford.

We used the ecosystem approach and compared the omnivores of interest, in this case the Neanderthals, with the isotope values of temporally and geographically associated fauna. Unfortunately, it was possible to extract collagen from only a few of the faunal samples taken from Vindija. A particular problem was our inability to extract collagen from our carnivore samples. For this reason, we have supplemented the Vindija faunal sample with data from the slightly later (?23,000–26,000 B.P.) sites of Dolní Vstonice II and Milovice in the Czech Republic (22). In addition, we have contributed a single herbivore sample from the site of Brno-Francouzská, which dates within this time range (23).

There are fluctuations in faunal δ15N values through time that are correlated with climate changes (24, 25). For example, Richards et al. (26) observed faunal δ15N values dated to ?12,000 years B.P. from Gough's Cave, U.K. that were ?2‰ lower than the δ15N values of similar species from the Holocene. Therefore, comparing isotope values between sites, especially sites of different ages, could be problematic. However, by employing fauna that are as geographically and temporally as close to our samples as possible, we should be providing an appropriate comparative framework for the Vindija Neanderthal samples. Moreover, the relative distribution, especially of δ15N values, for the species included in this pooled sample is similar to the distributions derived for various faunal species from single sites (10, 11).

Fauna. The Bos/Bison and cervid samples from Vindija (Table 1) have herbivore δ13C and δ15N values that are within the ranges observed for European Holocene specimens (25, 27). The δ13C values are more indicative of open-ranging species (?20Equation), rather than forest-dwelling species (?22Equation), but ranges of variation in Late Pleistocene Bos/Bison δ13C values (24) as well as the hilly terrain in the vicinity of Vindija Cave make it difficult to assess which of these bovine genera is most likely represented. The cave bear samples are interesting from a paleobiological, rather than an anthropological, perspective as they have very low δ15N values. Similarly low Ursus spelaeus δ15N values have been observed for samples from Slovenia (28), France (29), and Belgium (30). The low U. spelaeus values probably reflect a high degree of herbivory (31); they may also be a result of their unusual metabolism related to hibernation (32), although the hibernation model has been disputed (30).

Neanderthals. The Neanderthal samples from Vindija have high δ15N values, which indicate that the overwhelming majority of their dietary protein was from animal, rather than plant, sources (Table 1, Fig. 1). The associated δ13C values indicate the exploitation of more open-ranging herbivores, despite the hilly terrain of the Hrvatsko Zagorje. The Neanderthal values are close to the later carnivore isotope values from Dolní Vstonice II and Milovice (22), as well as those of earlier carnivores from Marillac and Scladina (10, 11), indicating that these Neanderthals had diets similar to nonhuman carnivores.

The insufficient associated faunal samples make it impossible to identify which herbivore species were preferentially being consumed by the Neanderthals. The mammoth δ15N values from Milovice are intriguing, as they are higher than the other herbivores. This pattern of higher mammoth values has been observed previously (30, 33, 34) and may relate to mammoths targeting specific plant species, whereas other herbivores consume a wider range of species. The higher mammoth δ15N values may be of relevance here, as the Neanderthal δ15N values could make sense if their main dietary protein source was mammoths rather than the other faunal species. However, archeological evidence for Neanderthal exploitation of proboscideans is extremely rare, and a broader series of fauna needs to be analyzed before the spectrum of predated herbivores can be evaluated through stable isotope analysis.

Our findings concerning the diet of the Vindija Neanderthals are remarkably similar to those observed by Bocherens and colleagues for other European Neanderthals (10, 11). They obtained similar δ13C and δ15N values for two Neanderthals from the site of Marillac dated to ?40,000–45,000 years B.P. and for a Neanderthal specimen from Scladina Cave, Belgium, which is earlier, dated to between 80,000 and 130,000 years B.P. (Table 2). Moreover, the high δ15N for the Marillac Neanderthal remains are most closely approached by the values for Canis lupus and Crocuta crocuta from that site (10), whereas the earlier Neanderthal δ15N value from Scladina is most closely approached in that site's faunal assemblage by Panthera spelaea and secondarily by slightly lower values for Crocuta crocuta and Canis lupus (11). For these five Neanderthal specimens, therefore, we have stable isotope data indicating that geographically and chronologically dispersed Neanderthals consistently behaved as top-level carnivores.

Neanderthals as Predators. Neanderthal subsistence strategies were varied in space and time, with carcass utilization patterns varying on intersite and interspecies levels (4, 35). The role of hunting versus scavenging in meat acquisition by Middle Paleolithic humans has been debated particularly over the last two decades (3, 36, 37), and from this discussion it has become clear that the Neanderthals were capable of, and frequently engaged in, predation on mammals.

In particular, taphonomic analyses of a number of Middle Paleolithic, Neanderthal-associated mammalian faunal assemblages in recent years have concluded that focused and selective hunting strategies resulting in high meat utility acquisition were carried out by these late archaic humans in areas of Europe and the Near East as dispersed as France (Bau de l'Aubesier, La Borde, Canalettes, Coudoulous, Mauran, Le Portel), Germany (Salzgitter Lebenstedt, Wallertheim), Italy (Grotta Breuil), Croatia (Krapina), Iran (Kobeh), Israel (Kebara), and Russia (Il'skaja) (13, 6, 35, 3844). These interpretations are based principally on mortality profiles and/or distributions of skeletal part frequencies of the prey species being processed, combined with direct evidence of human carcass processing with lithic tools. In the former, prime age-dominated assemblages are usually taken to indicate selective and active predation by these hominids. In the latter, a proximal limb element-dominated assemblage or a preserved skeletal distribution representative of anatomical frequencies, as opposed to a head and foot-dominated assemblage, are generally taken to indicate primary carcass access and hence active predation.

However, not only do a significant number of these assemblages not meet both criteria for active predation on the part of the Neanderthals, it is increasingly apparent that a variety of factors can contribute to the mortality and skeletal element distributions documented in archeological faunal assemblages. These factors include prey population demographic dynamics, nonhuman predator prey selection patterns, carcass consumption patterns by both humans and other carnivores, human carcass element transport variation, and postdepositional processes acting differentially on skeletal elements. Moreover, it remains unclear how representative of overall Neanderthal diet such episodes are. Consequently, current taphonomic analyses of these and other archeological faunal assemblages do not always permit assessment of the degrees to which the assemblages were accumulated through active predation versus scavenging.

Neanderthal predation has also been supported by the evidence for spears (stone-tipped and wooden) among both the Neanderthals and their Middle Pleistocene European predecessors (4549), combined with rare examples of such weapons in the remains of apparent prey animals [e.g., the wooden spear in the ribs of an Elephas skeleton at Lehringen, Germany, and the Levallois point embedded an Equus cervical vertebra from Umm el Tlel in Syria (45, 50)]. In addition, indirect measures of Neanderthal subsistence such as the Levallois point to core frequencies have been used to suggest that the Neanderthals were highly predatory in the Near East (ref. 51; but see refs. 52 and 53), despite the absence of evidence for the kind of projectile weaponry seen in the Upper Paleolithic that would increase the mechanical efficiency and safety of hunting or for the patterned variance in extractive technologies widely seen in Upper Paleolithic and more recent hunter–gatherer toolkits (54).

This inference of active predation on the part of the Neanderthals is further supported by their anatomical distribution of trauma, which suggests proximate encounters with large animals (55) of the kind necessitated by their predominantly heavy available weaponry (45, 47, 48, 56). Yet, their pattern of trauma does not permit distinctions between injuries sustained during hunting versus those suffered in competition with other carnivores for carcasses or space.

Consequently, although several lines of evidence support active mammalian predation by the Neanderthals and contradict the previous models of the Neanderthals acquiring their animal protein principally through scavenging, the archeological data nonetheless remain frequently ambiguous as to the extent to which these late archaic humans were the primary predators of the mammals whose remains they processed. The consistent stable isotope data indicating their position as top-level carnivores provides insight into this issue.

There are no true mammalian scavengers, as all are omnivores (ursids and canids) and/or actively hunt (hyenas) (57). This is because the search time for scavenging relative to the return is too expensive for terrestrial homeothermic vertebrates, and most predators actively defend their kills, thereby increasing risk to any potential terrestrial scavenger (57). If the Neanderthals were obtaining their animal protein principally through scavenging, they would have had to obtain most of their food from plants, as a reliable food source, and only supplemented this with scavenged animal products. Even though the isotope data cannot distinguish the species or even the sizes of the animals consumed, they clearly show that animal products were the overwhelming source of protein in European Neanderthal diets and that protein from plants was insignificant. It is therefore likely that scavenging, although undoubtedly practiced on an opportunistic basis by these European Neanderthals, must have been distinctly secondary to predation.

Summary and Conclusions 


Isotope analyses of two Neanderthals and associated fauna from Vindija Cave, Croatia, have indicated that the bulk of their dietary protein came from animal sources. Comparison with faunal remains from this and other sites of similar age indicates that the Vindija Neanderthal isotope values were similar to those of other carnivores. These results are very close to the results for earlier Late Pleistocene Neanderthals from France and Belgium.

Therefore, the emerging picture of the European Neanderthal diet indicates that although physiologically they were presumably omnivores, they behaved as carnivores, with animal protein being the main source of dietary protein. This finding is in agreement with the indirect archeological evidence and strongly points to the Neanderthals having been active predators.



Article published online before print: Proc. Natl. Acad. Sci. USA, 10.1073/pnas.120178997.

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5. J Hum Evol. 2002 Dec;43(6):831-72.

Male strategies and Plio-Pleistocene archaeology.

O'Connell JF, Hawkes K, Lupo KD, Blurton Jones NG.

Archaeological data are frequently cited in support of the idea that big game hunting drove the evolution of early Homo, mainly through its role in offspring provisioning. This argument has been disputed on two grounds: (1) ethnographic observations on modern foragers show that although hunting may contribute a large fraction of the overall diet, it is an unreliable day-to-day food source, pursued more for status than subsistence; (2) archaeological evidence from the Plio-Pleistocene, coincident with the emergence of Homo can be read to reflect low-yield scavenging, not hunting. Our review of the archaeology yields results consistent with these critiques: (1) early humans acquired large-bodied ungulates primarily by aggressive scavenging, not hunting; (2) meat was consumed at or near the point of acquisition, not at home bases, as the hunting hypothesis requires; (3) carcasses were taken at highly variable rates and in varying degrees of completeness, making meat from big game an even less reliable food source than it is among modern foragers. Collectively, Plio-Pleistocene site location and assemblage composition are consistent with the hypothesis that large carcasses were taken not for purposes of provisioning, but in the context of competitive male displays. Even if meat were acquired more reliably than the archaeology indicates, its consumption cannot account for the significant changes in life history now seen to distinguish early humans from ancestral australopiths. The coincidence between the earliest dates for Homo ergaster and an increase in the archaeological visibility of meat eating that many find so provocative instead reflects: (1) changes in the structure of the environment that concentrated scavenging opportunities in space, making evidence of their pursuit more obvious to archaeologists; (2) H. ergaster's larger body size (itself a consequence of other factors), which improved its ability at interference competition.


6. Proc Natl Acad Sci U S A. 2001 February 13; 98 (4): 1358–1363
Evidence of termite foraging by Swartkrans early hominids
Lucinda R. Backwell and Francesco d'Errico

Previous studies have suggested that modified bones from the Lower Paleolithic sites of Swartkrans and Sterkfontein in South Africa represent the oldest known bone tools and that they were used by Australopithecus robustus to dig up tubers. Macroscopic and microscopic analysis of the wear patterns on the purported bone tools, pseudo bone tools produced naturally by known taphonomic processes, and experimentally used bone tools confirm the anthropic origin of the modifications. However, our analysis suggests that these tools were used to dig into termite mounds, rather than to dig for tubers. This result indicates that early hominids from southern Africa maintained a behavioral pattern involving a bone tool material culture that may have persisted for a long period and strongly supports the role of insectivory in the early hominid diet.

7. Homo. 2003;54(1):1-28.
Early hominin speciation at the Plio/Pleistocene transition.
Cameron DW.

Over the last half-decade or so, there has been an explosion in the recognition of hominin genera and species. We now have the late Miocene genera Orrorin and Sahelanthropus, the mid Pliocene genus Kenyanthropus, three new Pliocene species of Australopithecus (A. anamensis, A. garhi and A. bahrelghazali) and a sub species of Ardipithecus (Ar. r. kadabba) to contend with. Excepting also the more traditional species allocated to Paranthropus, Australopithecus and early Homo we are approaching around 15 species over 5 million years (excluding hominin evolution over the last one million years). Can such a large number of hominin species be justified? An examination of extant hominid (Gorilla gorilla, Pan troglodytes, and Pan paniscus) anatomical variability indicates that the range of fossil hominin variability supports the recognition of this large number of fossil species. It is also shown that not all hominins are directly related to the emergence of early Homo and as such have become extinct. Indeed the traditional australopithecine species 'A'. anamensis, 'A'. afarensis and 'A'. garhi are considered here to belong to a distinct genus Praeanthropus. They are also argued not be hominins, but rather an as yet undefined hominid group from which the more derived hominins evolved. The first hominin is represented by A. africanus or a hominin very much like it. The Paranthropus clade is defined by a derived heterochronic condition of peramorphosis, associated with sequential progenesis (contraction of successive growth stages) in brain and dental development, but a mixture of peramorphic and paedomorphic features in its craniofacial anatomy. Conversely, Kenyanthropus and Homo both share a pattern of peramorphosis, associated with sequential hypermorphosis (prolongation of successive growth stages) in brain development, and paedomorphosis processes in cranial, facial and dental development. This suggests, that these two clades share an important synapomorphy not recognised in the parsimony analyses, suggesting that they may form a sister group relationship to the exclusion of Paranthropus. This highlights the need to re-interpret phylogenetic results in terms of function and development. The rapid speciation and extinction as argued here is in keeping with other fossil groups in Africa at the Plio/Pleistocene transition. This emphasises that we must approach the pre-australopithecines and hominins as part of the endemic African fauna, and not in isolation to the evolutionary and climatic processes that were operating all around them.


8. Первоописание Ardipithecus kadabba.

Nature. 2001 Jul 12;412(6843):178-81.
Late Miocene hominids from the Middle Awash, Ethiopia.
Haile-Selassie Y.

Molecular studies suggest that the lineages leading to humans and chimpanzees diverged approximately 6.5-5.5 million years (Myr) ago, in the Late Miocene. Hominid fossils from this interval, however, are fragmentary and of uncertain phylogenetic status, age, or both. Here I report new hominid specimens from the Middle Awash area of Ethiopia that date to 5.2-5.8 Myr and are associated with a wooded palaeoenvironment. These Late Miocene fossils are assigned to the hominid genus Ardipithecus and represent the earliest definitive evidence of the hominid clade. Derived dental characters are shared exclusively with all younger hominids. This indicates that the fossils probably represent a hominid taxon that postdated the divergence of lineages leading to modern chimpanzees and humans. However, the persistence of primitive dental and postcranial characters in these new fossils indicates that Ardipithecus was phylogenetically close to the common ancestor of chimpanzees and humans. These new findings raise additional questions about the claimed hominid status of Orrorin tugenensis, recently described from Kenya and dated to approximately 6 Myr.

9. Science. 2004 Mar 5;303(5663):1503-5.
Late Miocene teeth from Middle Awash, Ethiopia, and early hominid dental evolution.
Haile-Selassie Y, Suwa G, White TD.

Late Miocene fossil hominid teeth recovered from Ethiopia's Middle Awash are assigned to Ardipithecus kadabba. Their primitive morphology and wear pattern demonstrate that A. kadabba is distinct from Ardipithecus ramidus. These fossils suggest that the last common ancestor of apes and humans had a functionally honing canine-third premolar complex. Comparison with teeth of Sahelanthropus and Orrorin, the two other named late Miocene hominid genera, implies that these putative taxa are very similar to A. kadabba. It is therefore premature to posit extensive late Miocene hominid diversity on the basis of currently available samples.

10. Nature. 1997 Feb 27;385(6619):807-10.
Lower Palaeolithic hunting spears from Germany.
Thieme H.

Little is known about the organic component of Lower and Middle Palaeolithic technologies, particular with respect to wooden tools. Here I describe some wooden throwing spears about 400,000 years old that were discovered in 1995 at the Pleistocene site at Schöningen, Germany. They are thought to be the oldest complete hunting weapons so far discovered to have been used by humans. Found in association with stone tools and the butchered remains of more than ten horses, the spears strongly suggest that systematic hunting, involving foresight, planning and the use of appropriate technology, was part of the behavioural repertoire of pre-modern hominids. The use of sophisticated spears as early as the Middle Pleistocene may mean that many current theories on early human behaviour and culture must be revised.

11. Nature. 1998 May 7;393(6680):62-6.
New specimens and confirmation of an early age for Australopithecus anamensis.
Leakey MG, Feibel CS, McDougall I, Ward C, Walker A.
The discovery of Australopithecus anamensis fossils from strata lying between tephra dated at 4.17 and 4.12 million years ago, and from slightly higher strata not well constrained in age by overlying dated units, provoked the claim that more than one species might be represented: it was suggested that the stratigraphically higher fossils, which include the important tibia, humerus and a large, presumed male, mandible (KNM-KP 29287), might belong to a later, more derived hominid. We have recovered new fossils from Kanapoi and Allia Bay, Kenya, during field work in 1995-1997 that confirm the primitive status of Australopithecus anamensis, the earliest species of Australopithecus. Isotope dating confirms A. anamensis' intermediate age as being between those of Ardipithecus ramidus and Australopithecus afarensis. New specimens of maxilla, mandible and capitate show that this species is demonstrably more primitive than A. afarensis. A lower first deciduous molar (dm 1) is intermediate in morphology between that reported for Ardipithecus ramidus and A. afarensis. Single-crystal 40Ar-39Ar age determinations on the Kanapoi Tuff show that, except for a large mandible, all of the hominid fossils from Kanapoi are from sediments deposited between 4.17+/-0.03 and 4.07+/-0.02 million years ago.


12. J Hum Evol. 2004 May;46(5):605-22.
Dental topography and diets of Australopithecus afarensis and early Homo.
Ungar P.
Diet is key to understanding the paleoecology of early hominins. We know little about the diets of these fossil taxa, however, in part because of a limited fossil record, and in part because of limitations in methods available to infer their feeding adaptations. This paper applies a new method, dental topographic analysis, to the inference of diet from fossil hominin teeth. This approach uses laser scanning to generate digital 3D models of teeth and geographic information systems software to measure surface attributes, such as slope and occlusal relief. Because it does not rely on specific landmarks that change with wear, dental topographic analysis allows measurement and comparison of variably worn teeth, greatly increasing sample sizes compared with techniques that require unworn teeth. This study involved comparison of occlusal slope and relief of the lower second molars of Australopithecus afarensis (n=15) and early Homo (n=8) with those of Gorilla gorilla gorilla (n=47) and Pan troglodytes troglodytes (n=54). Results indicate that while all groups show reduced slope and relief in progressively more worn specimens, there are consistent differences at given wear stages among the taxa. Early Homo shows steeper slopes and more relief than chimpanzees, whereas A. afarensis shows less slope and relief than any of the other groups. The differences between the two hominin taxa are on the same order as those between the extant apes, suggesting similar degrees of difference in diet. Because these chimpanzees and gorillas differ mostly in fallback foods where they are sympatric, results suggest that the early hominins may likewise have differed mostly in fallback foods, with A. afarensis emphasizing harder, more brittle foods, and early Homo relying on tougher, more elastic foods.


13. J Hum Evol. 2003 May;44(5):581-97.
The carbon isotope ecology and diet of Australopithecus africanus at Sterkfontein, South Africa.
van der Merwe NJ, Thackeray JF, Lee-Thorp JA, Luyt J.
The stable carbon isotope ratio of fossil tooth enamel carbonate is determined by the photosynthetic systems of plants at the base of the animal's foodweb. In subtropical Africa, grasses and many sedges have C(4)photosynthesis and transmit their characteristically enriched 13C/(12)C ratios (more positive delta13C values) along the foodchain to consumers. We report here a carbon isotope study of ten specimens of Australopithecus africanus from Member 4, Sterkfontein (ca. 2.5 to 2.0Ma), compared with other fossil mammals from the same deposit. This is the most extensive isotopic study of an early hominin species that has been achieved so far. The results show that this hominin was intensively engaged with the savanna foodweb and that the dietary variation between individuals was more pronounced than for any other early hominin or non-human primate species on record. Suggestions that more than one species have been incuded in this taxon are not supported by the isotopic evidence. We conclude that Australopithecus africanus was highly opportunistic and adaptable in its feeding habits.


14. Nutr Health. 2002;16(4):267-89. 
Wading for food the driving force of the evolution of bipedalism?
Kuliukas A.
Evidence is accumulating that suggests that the large human brain is most likely to have evolved in littoral and estuarine habitats rich in naturally occurring essential fatty acids. This paper adds further weight to this view, suggesting that another key human trait, our bipedality might also be best explained as an adaptation to a water-side niche. Evidence is provided here that extant apes, although preferring to keep dry, go into water when driven to do so by hunger. The anecdotal evidence has suggested that they tend to do this bipedally. Here, a new empirical study of captive bonobos found them to exhibit 2% or less bipedality on the ground or in trees but over 90% when wading in water to collect food. The skeletal morphology of AL 288-1 ("Lucy") is shown to indicate a strong ability to abduct and adduct the femur. These traits, together with a remarkably platypelloid pelvis, have not yet been adequately explained by terrestrial or arboreal models for early bipedalism but are consistent with those expected in an ape that adopted a specialist side-to-side 'ice-skating' or sideways wading mode. It is argued that this explanation of A. afarensis locomotor morphology is more parsimonious than others which have plainly failed to produce a consensus. Microwear evidence of Australopithecus dentition is also presented as evidence that the drive for such a wading form of locomotion might well have been waterside foods. This model obtains further support from the paleo-habitats of the earliest known bipeds, which are consistent with the hypothesis that wading contributed to the adaptive pressure towards bipedality.

15. J Hum Evol. 2000 Dec;39(6):565-76.
The hunters and the hunted revisited.
Lee-Thorp J, Thackeray JF, van der Merwe N.
The dietary niches of extinct animals, including hominids and predators, may be constrained using stable carbon isotope ratios in fossil tooth enamel.(13)C/(12)C ratios of many of the primates abundant in the faunal assemblages of Members 1 and 2 at Swartkrans, including cercopithecoids and Australopithecus (Paranthropus) robustus, and a range of other possible prey species, have been reported previously. Resulting suggestions of a mixed, or omnivorous, diet for A. robustus raise questions about niche overlap with coeval, larger brained Homo. Here we present(13)C/(12)C data from Homo and several large predators including Panthera pardus, Dinofelis sp., Megantereon cultridens and Chasmoporthetes nitidula in Member 1, and P. pardus and P. leo in Member 2, in order to compare the two hominid species and to determine likely predators of the various primates and other macrovertebrates. Results for three Homo cf. ergaster individuals are indistinguishable from those of A. robustus, showing that proportions of C(3)- and C(4)-based foods in their diets did not differ. P. pardus, Megantereon and Crocuta are shown to be likely predators of the hominids and Papio baboons in Member 1, while the Dinofelis individual concentrated on prey which consumed C(4)grasses. The hunting hyaenid C. nitidula preyed on either mixed feeders or on a range of animals across the spectrum of C(3)and C(4)variation. The data from Members 1 and 2 confirm a shift in leopard diets towards animals that consumed C(4)grasses. Copyright 2000 Academic Press.

16. Science. 1999 Jan 15;283(5400):368-70. 
Isotopic evidence for the diet of an early hominid, Australopithecus africanus.
Sponheimer M, Lee-Thorp JA.
M. Sponheimer, Department of Anthropology, Rutgers University, New Brunswick NJ 08901-1414, USA.
Current consensus holds that the 3-million-year-old hominid Australopithecus africanus subsisted on fruits and leaves, much as the modern chimpanzee does. Stable carbon isotope analysis of A. africanus from Makapansgat Limeworks, South Africa, demonstrates that this early hominid ate not only fruits and leaves but also large quantities of carbon-13-enriched foods such as grasses and sedges or animals that ate these plants, or both. The results suggest that early hominids regularly exploited relatively open environments such as woodlands or grasslands for food. They may also suggest that hominids consumed high-quality animal foods before the development of stone tools and the origin of the genus Homo.


17. Asia Pac J Clin Nutr. 2004;13(Suppl):S17. 
Paleolithic nutrition: what can we learn from the past?
Mann NJ.
Department of Food Science, RMIT University, Melbourne, 3001, Australia.
Background - Anthropologists and some nutritionists have long recognised that the diets of Paleolithic and recent hunter-gatherers (HG) may represent a reference standard for modern human nutrition and a model for defense against certain western lifestyle diseases. Boyd Eaton of Emory University (Atlanta) has spent over 20 years reconstructing prehistoric diets from anthropological evidence and observations of surviving HG societies, put this succinctly: "We are the heirs of inherited characteristics accrued over millions of years, the vast majority of our biochemistry and physiology are tuned to life conditions that existed prior to the advent of agriculture some 10,000 years ago. Genetically our bodies are virtually the same as they were at the end of paleolithic some 20,000 years ago. The appearance of agriculture and domestication of animals some 10,000 years ago and the Industrial Revolution some 200 years ago introduced new dietary pressures for which no adaptation has been possible in such a short time span. Thus an inevitable discordance exists between our dietary intake and that which our genes are suited to". This discordance hypothesis postulated by Eaton, could explain many of the chronic "diseases of civilisation". But what did hunter-gatherer populations actually eat? Review - The lines of investigation used by anthropologists to deduce the evolutionary diet of hominids include the study of: (i) changes in cranio-dental features, (ii) isotopic chemical tracer methods, including carbon isotope (13C/12C), strontium isotope (87Sr/86Sr) and trace element Sr/Ca ratios in enamel and bone of fossils,(iii) comparative gut morphology of modern humans and other mammals, (iv) the energetic requirements of a developing a large brain:body size ratio, (v) optimal foraging theory and food selection, (vi) the study of dietary patterns of surviving hunter-gatherer societies. Findings show clear cranio-dental changes including, a decrease in molar teeth size, jaws/skull became more gracile and front teeth became well-buttressed, all indicative of less emphasis on grinding course foliage and more on biting and tearing. Carbon isotope studies indicate the dietary intake of C4 grasses, undoubtedly in the form of herbivorous animals, at a level which increased substantially during the progression of our genus from A. aferensis to H. sapiens. Even as far back as 3.5 million years, the Sr/Ca ratio falls in between those typical for herbivores and carnivores. Gut morphology studies indicate a closer structural analogy with carnivores than the folivorous or frugivorous mammals. Energetic requirements of a relatively enlarged brain have been balanced by reduction in size and energy requirement of the digestive system, a phenomena requiring a high quality diet. Investigation of food procurement habits of hunter-gatherer societies indicates the advantage of hunting of game animals compared with plant foraging in terms of energy gain versus expenditure. Study of macronutrient energy proportions in the diet of HG societies (n=229) show a relatively high protein intake 19-35%, highly variable fat intake 28-47% and low carbohydrate level 22-40%. Conclusions - It is postulated that changes in food staples and food processing procedures introduced during the Neolithic and Industrial era have fundamentally altered seven crucial nutritional characteristics of our ancestral diet: (i) glycaemic load, (ii) fatty acid balance, (iii) macronutrient balance, (iv) trace nutrient density, (v) acid-base balance, (vi) sodium-potassium balance, (vii) fiber content.

18.  J Hum Evol. 2004 Feb;46(2):119-62.
Patterns of resource use in early Homo and Paranthropus.
Wood B, Strait D.
CASHP and Department of Anthropology, The George Washington University, 2110 G St. NW, Washington, DC 20052, USA.
Conventional wisdom concerning the extinction of Paranthropus suggests that these species developed highly derived morphologies as a consequence of specializing on a diet consisting of hard and/or low-quality food items. It goes on to suggest that these species were so specialized or stenotopic that they were unable to adapt to changing environments in the period following 1.5 Ma. The same conventional wisdom proposes that early Homo species responded very differently to the same environmental challenges. Instead of narrowing their niche it was the dietary and behavioral flexibility (eurytopy) exhibited by early Homo that enabled that lineage to persist. We investigate whether evidence taken across eleven criteria supports a null hypothesis in which Paranthropus is more stenotopic than early Homo. In six instances (most categories of direct evidence of dietary breadth, species diversity, species duration, susceptibility to dispersal, dispersal direction, and non-dietary adaptations) the evidence is inconsistent with the hypothesis. Only one line of indirect evidence for dietary breadth-occlusal morphology-is unambiguously consistent with the null hypothesis that Paranthropus' ability to process tough, fibrous food items (e.g., leaves) was reduced relative to early Homo. Other criteria (habitat preference, population density, direct and indirect evidence of dietary breadth related to incisor use) are only consistent with the hypothesis under certain conditions. If those conditions are not met, then the evidence is either inconsistent with the hypothesis, or ambiguous. On balance, Paranthropus and early Homo were both likely to have been ecological generalists. These data are inconsistent with the conventional wisdom that stenotopy was a major contributing factor in the extinction of the Paranthropus clade. Researchers will need to explore other avenues of research in order to generate testable hypotheses about the demise of Paranthropus. Ecological models that may explain the evolution of eurytopy in early hominins are discussed.


19. J Hum Evol. 2002 Sep;43(3):291-321.

Examining time trends in the Oldowan technology at Beds I and II, Olduvai Gorge.
Kimura Y.
Institute of History and Anthropology, University of Tsukuba, 1-1-1 Ten'noudai, Tsukuba, Ibaraki, Japan 305-8571.
The lithic analysis of the Bed I and II assemblages from Olduvai Gorge reveals both static and dynamic time trends in early hominids' technology from 1.8 to 1.2 m.y.a. The Bed I Oldowan (1.87-1.75 m.y.a.) is characterized by the least effort strategy in terms of raw material exploitation and tool production. The inclusion of new raw material, chert, for toolmaking in the following Developed Oldowan A (DOA, 1.65-1.53 m.y.a.) facilitated more distinctive and variable flaking strategies depending on the kind of raw materials. The unique characters of DOA are explainable by this raw material factor, rather than technological development of hominids. The disappearance of chert in the subsequent Developed Oldowan B and Acheulian (1.53-1.2 m.y.a.) necessitated a shift in tool production strategy more similar to that of Bed I Oldowan than DOA. However, the evidence suggests that Bed II hominids might have been more skillful toolmakers, intensive tool-users, and engaged in more active transport of stone tools than the Bed I predecessors. Koobi Fora hominids maintained a more static tool-using behavior than their Olduvai counterparts due mainly to a stable supply of raw materials. They differed from Olduvai hominids in terms of less battering of cores, consistent transport behavior, and few productions of side-struck flakes, indicating a regional variation of toolmaking and using practice. However, they shared with Olduvai hominids a temporal trend toward the production of larger flakes from larger cores after 1.6 m.y.a. Increased intake of animal resources and the expansion of ranging area of Homo ergaster would have led to the development of technological organization. Technological changes in the Oldowan industry are attested at Olduvai Gorge, Koobi Fora, and Sterkfontein, suggesting that it was a pan-African synchronous phenomenon, beginning at 1.5 m.y.a.


20. Am J Phys Anthropol. 2002 Oct;119(2):192-7.
Revised age estimates of Australopithecus-bearing deposits at Sterkfontein, South Africa.
Berger LR, Lacruz R, De Ruiter DJ.

Palaeoanthropology Unit for Research and Exploration, Bernard Price Institute for Palaeontology, University of the Witwatersrand, Johannesburg 2050, South Africa.

The Sterkfontein fossil site in South Africa has produced the largest concentration of early hominin fossils from a single locality. Recent reports suggest that Australopithecus from this site is found within a broad paleontological age of between 2.5-3.5 Ma (Partridge [2000] The Cenozoic of Southern Africa, Oxford: Oxford Monographs, p. 100-125; Partridge et al. [2000a], The Cenozoic of Southern Africa, Oxford: Oxford Monographs, p. 129-130; Kuman and Clarke [2000] J Hum Evol 38:827-847). Specifically, the hominin fossil commonly referred to as the "Little Foot" skeleton from Member 2, which is arguably the most complete early hominin skeleton yet discovered, has been magnetostratigraphically dated to 3.30-3.33 Ma (Partridge [2000] The Cenozoic of Southern Africa, Oxford: Oxford Monographs, p. 100-125; Partridge et al. [2000a], The Cenozoic of Southern Africa, Oxford: Oxford Monographs, p. 129-130). More recent claims suggest that hominin fossils from the Jacovec Cavern are even older, being dated to approximately 3.5 Ma. Our interpretation of the fauna, the archeometric results, and the magnetostratigraphy of Sterkfontein indicate that it is unlikely that any Members yet described from Sterkfontein are in excess of 3.04 Ma in age. We estimate that Member 2, including the Little Foot skeleton, is younger than 3.0 Ma, and that Member 4, previously dated to between 2.4-2.8 Ma, is more likely to fall between 1.5-2.5 Ma. Our results suggest that Australopithecus africanus should not be considered as a temporal contemporary of Australopithecus afarensis, Australopithecus bahrelghazali, and Kenyanthropus platyops. Copyright 2002 Wiley-Liss, Inc.

21. Nature. 2004 Apr 29;428(6986):936-9.
Surprisingly rapid growth in Neanderthals.
Ramirez Rozzi FV, Bermudez De Castro JM.
UPR 2147, Dyamique de l'Evolution Humaine, CNRS, 44, Rue de l'Amiral Mouchez, 75014 Paris, France.
Life-history traits correlate closely with dental growth, so differences in dental growth within Homo can enable us to determine how somatic development has evolved and to identify developmental shifts that warrant species-level distinctions. Dental growth can be determined from the speed of enamel formation (or extension rate). We analysed the enamel extension rate in Homo antecessor (8 teeth analysed), Homo heidelbergensis (106), Homo neanderthalensis ('Neanderthals'; 146) and Upper Palaeolithic-Mesolithic Homo sapiens (100). Here we report that Upper Palaeolithic-Mesolithic H. sapiens shared an identical dental development pattern with modern humans, but that H. antecessor and H. heidelbergensis had shorter periods of dental growth. Surprisingly, Neanderthals were characterized by having the shortest period of dental growth. Because dental growth is an excellent indicator of somatic development, our results suggest that Neanderthals developed faster even than their immediate ancestor, H. heidelbergensis. Dental growth became longer and brain size increased from the Plio-Pleistocene in hominid evolution. Neanderthals, despite having a large brain, were characterized by a short period of development. This autapomorphy in growth is an evolutionary reversal, and points strongly to a specific distinction between H. sapiens and H. neanderthalensis.

22. Am J Phys Anthropol. 2002 Nov;119(3):199-204.
Does Homo neanderthalensis play a role in modern human ancestry? The mandibular evidence.
Rak Y, Ginzburg A, Geffen E.
Department of Anatomy and Anthropology, Sackler Faculty of Medicine, Tel Aviv University, Tel Aviv 69978, Israel.
Data obtained from quantifying the upper part of the mandibular ramus (the coronoid process, the condylar process, and the notch between them) lead us to conclude that Neanderthals (both European and Middle Eastern) differ more from Homo sapiens (early specimens such as Tabun II, Skhul, and Qafzeh, as well as contemporary populations from as far apart as Alaska and Australia) than the latter differs from Homo erectus. The specialized Neanderthal mandibular ramus morphology emerges as yet another element constituting the derived complex of morphologies of the mandible and face that are unique to Neanderthals. These morphologies provide further support for the contention that Neanderthals do not play a role in modern human biological ancestry, either through "regional continuity" or through any other form of anagenetic progression. Copyright 2002 Wiley-Liss, Inc.

23. Am J Reprod Immunol. 2002 Jul;48(1):43-9.
Evolutionary genetics and infertility.
Gill TJ 3rd.
Department of Pathology, University of Pittsburgh (Emeritus), Duxbury, MA 02332-4801, USA.
PROBLEM: Mapping the human genome and advances in human evolution indicate a critical role for genetics in the study of reproduction. METHOD OF STUDY: Literature in human evolution, genetics and reproduction. RESULTS: This paper will focus on three points: (1) the course of primate evolution, (2) implications of this course for reproduction in humans, and (3) evolutionary mechanisms. Reproduction is the driving force of evolution, and sex selection is the important factor in determining reproductive activity. The hypothesis will be presented that the reproductive inefficiency of Homo sapiens (one of four pregnancies succeeds) is the consequence of the development of genetic isolation from other species of Homo that was necessary to drive the evolution of Homo sapiens. This hypothesis is based on the evidence that Homo sapiens and Homo neanderthalensis are separate species and that intermating would not yield fertile offspring (Haldane's Rule). This mating barrier involves zoological and linguistic differences and genetic mechanisms preventing fertile interspecies mating (chromosomal incompatibilities, disruption of genomic imprinting and recessive lethal defects). CONCLUSIONS: The medical and philosophical implications of modern reproductive technologies that may circumvent mating barriers must be addressed prospectively: propagation of genetic defects that would be eliminated by natural selection; the specter of 'genetic enhancement'; human cloning; and attempts to control the future direction of the evolution of Homo sapiens.


24. Proc Natl Acad Sci U S A. 2002 Feb 5;99(3):1134-9. Epub 2002 Jan 22.
The evolution and development of cranial form in Homo sapiens.
Lieberman DE, McBratney BM, Krovitz G.
Department of Anthropology, Harvard University, 11 Divinity Avenue, Cambridge, MA 02138, USA.
Despite much data, there is no unanimity over how to define Homo sapiens in the fossil record. Here, we examine cranial variation among Pleistocene and recent human fossils by using a model of cranial growth to identify unique derived features (autapomorphies) that reliably distinguish fossils attributed to "anatomically modern" H. sapiens (AMHS) from those attributed to various taxa of "archaic" Homo spp. (AH) and to test hypotheses about the changes in cranial development that underlie the origin of modern human cranial form. In terms of pattern, AMHS crania are uniquely characterized by two general structural autapomorphies: facial retraction and neurocranial globularity. Morphometric analysis of the ontogeny of these autapomorphies indicates that the developmental changes that led to modern human cranial form derive from a combination of shifts in cranial base angle, cranial fossae length and width, and facial length. These morphological changes, some of which may have occurred because of relative size increases in the temporal and possibly the frontal lobes, occur early in ontogeny, and their effects on facial retraction and neurocranial globularity discriminate AMHS from AH crania. The existence of these autapomorphies supports the hypothesis that AMHS is a distinct species from taxa of "archaic" Homo (e.g., Homo neanderthalensis).

25. J Hum Evol. 1999 Sep-Oct;37(3-4):459-99. Related Articles, Links
Axial and appendicular skeleton of Homo antecessor.
Carretero JM, Lorenzo C, Arsuaga JL.
Departamento de Ciencias Historicas y Geografia, Facultad de Humanidades y Educacion, Universidad de Burgos, Carretera Villadiego s/n, Burgos, 09001, Spain.
The human trunk and limb bones recovered from the Gran Dolina site, in the Sierra de Atapuerca (Burgos, Spain) are studied. All these fossils were excavated at the level called TD6 between 1994 and 1995 and have been dated in excess of 780,000 years ago. These remains have been recently attributed to a new Homo species named Homo antecessor. Axial (vertebrae and ribs) and part of the appendicular (clavicles, radii, femur and patellae) skeleton are studied here. Hand and foot bones have been studied elsewhere (Lorenzo et al., 1999). Four is the minimum number of individuals represented by the postcranial remains recovered up to now. All elements are briefly described anatomically, measured and compared with other fossil hominids and modern humans in order to establish, as far as possible, what postcranial morphology characterized this new species of our genus.The H. antecessor postcrania, generally, display a set of morphological traits that are more similar to modern humans than to the Middle and Upper Pleistocene European hominids. Our results do not contradict the previous phylogenetic analysis, i.e., that H. antecessor represents the last common ancestor for H. sapiens (modern humans) and H. neanderthalensis (Neandertals). Copyright 1999 Academic Press.



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