Prehistoric demography, palaeodemography or archaeological demography is the study of human and hominid demography in prehistory. [1]
More specifically, palaeodemography looks at the changes in pre-modern populations in order to determine something about the influences on the lifespan and health of earlier peoples.[ citation needed ] Reconstructions of ancient population sizes and dynamics are based on bioarchaeology, [2] ancient DNA, and inference from modern population genetics.[ citation needed ]
Skeletal analysis can yield information such as an estimation of age at time of death. There are numerous methods that can be used; [3] in addition to age estimation and sex estimation, someone versed in basic osteology can ascertain a minimum number of individuals (or MNI) in cluttered contexts—such as in mass graves or an ossuary. This is important, as it is not always obvious how many bodies compose the bones sitting in a heap as they are excavated.
Occasionally, historical disease prevalence for illnesses such as leprosy can also be determined from bone restructuring and deterioration. Paleopathology, as these investigations are called, can be useful in accurate estimation of mortality rates.
The increasing availability of DNA sequencing since the late 1990s has allowed estimates on Paleolithic effective population sizes. [4] [5] [6] Such models suggest a human effective population size of the order of 10,000 individuals for the Late Pleistocene. This includes only the breeding population that produced descendants over the long term, and the actual population may have been substantially larger (in the six digits). [7] Sherry et al. (1997) based on Alu elements estimated a roughly constant effective population size of the order of 18,000 individuals for the population of Homo ancestral to modern humans over the past one to two million years. [8] Huff et al. (2010) rejected all models with an ancient effective population size larger than 26,000. [9] For ca. 130,000 years ago, Sjödin et al. (2012) estimate an effective population size of the order of 10,000 to 30,000 individuals, and infer an actual "census population" of early Homo sapiens of roughly 100,000 to 300,000 individuals. [10] The authors also note that their model disfavours the assumption of an early (pre- Out-of-Africa ) population bottleneck affecting all of Homo sapiens. [11]
According to a 2015 study, the total land area of Africa, Eurasia, and Sahul that was habitable to humans during the Last Glacial Maximum (LGM) was around 76,959,712.4 km2. Based on a dataset of average population density of hunter-gatherer groups collected by Lewis R. Binford, which indicate a mean density of 0.1223 humans per km2 and a median density of 0.0444 humans per km2, the combined human population of Africa and Eurasia at the time of the LGM would have been between 2,998,820 and 8,260,262 people. Alternatively, if a human population density based on that of modern medium to large-bodied carnivores, whose median density is 0.0275 individuals per km2 and whose mean density is 0.0384 individuals per km2, is used, a total Afro-Eurasian human population of 2,120,000 to 2,950,000 is obtained. Sahul's population density was significantly lower than that of Afro-Eurasia, being calculated as only 0.005 humans per km2 during the time just prior to the LGM. As a consequence, assuming Sahul possessed an estimated total habitable land area of 9,418,730.8 km2, its population was at most 47,000 at the time of the LGM, and probably less than that given that its population is believed to have declined by as much as 61% during the LGM, [12] a demographic trend supported by archaeological evidence, [13] and it thus would have possessed an even lower actual population density than the calculated density from just before the LGM. [14]
It is estimated by J. Lawrence Angel [15] that the average life span of hominids on the African savanna between 4,000,000 and 200,000 years ago was 20 years. This means that the population would be completely renewed about five times per century,[ citation needed ] assuming that infant mortality has already been accounted for[ clarification needed ]. It is further estimated that the population of hominids in Africa fluctuated between 10,000 and 100,000 individuals, thus averaging about 50,000 individuals[ clarification needed ]. Multiplying 40,000 centuries by 50,000 to 500,000 individuals per century yields a total of 2 billion to 20 billion hominids that lived during that approximately 4,000,000-year time span. [16]
Human evolution is the evolutionary process within the history of primates that led to the emergence of Homo sapiens as a distinct species of the hominid family, which includes all the great apes. This process involved the gradual development of traits such as human bipedalism, dexterity and complex language, as well as interbreeding with other hominins, indicating that human evolution was not linear but weblike. The study of human evolution involves several scientific disciplines, including physical and evolutionary anthropology, paleontology, and genetics.
Genetic drift, also known as allelic drift or the Wright effect, is the change in the frequency of an existing gene variant (allele) in a population due to random chance.
In human genetics, the Mitochondrial Eve is the matrilineal most recent common ancestor (MRCA) of all living humans. In other words, she is defined as the most recent woman from whom all living humans descend in an unbroken line purely through their mothers and through the mothers of those mothers, back until all lines converge on one woman.
Small populations can behave differently from larger populations. They are often the result of population bottlenecks from larger populations, leading to loss of heterozygosity and reduced genetic diversity and loss or fixation of alleles and shifts in allele frequencies. A small population is then more susceptible to demographic and genetic stochastic events, which can impact the long-term survival of the population. Therefore, small populations are often considered at risk of endangerment or extinction, and are often of conservation concern.
Early modern human (EMH) or anatomically modern human (AMH) are terms used to distinguish Homo sapiens that are anatomically consistent with the range of phenotypes seen in contemporary humans from extinct archaic human species. This distinction is useful especially for times and regions where anatomically modern and archaic humans co-existed, for example, in Paleolithic Europe. Among the oldest known remains of Homo sapiens are those found at the Omo-Kibish I archaeological site in south-western Ethiopia, dating to about 233,000 to 196,000 years ago, the Florisbad site in South Africa, dating to about 259,000 years ago, and the Jebel Irhoud site in Morocco, dated about 315,000 years ago.
The Youngest Toba eruption was a supervolcano eruption that occurred around 74,000 years ago at the site of present-day Lake Toba in Sumatra, Indonesia. It is one of the Earth's largest known explosive eruptions. The Toba catastrophe theory holds that this event caused a global volcanic winter of six to ten years and possibly a 1,000-year-long cooling episode, leading to agenetic bottleneck in humans.
A population bottleneck or genetic bottleneck is a sharp reduction in the size of a population due to environmental events such as famines, earthquakes, floods, fires, disease, and droughts; or human activities such as specicide, widespread violence or intentional culling, and human population planning. Such events can reduce the variation in the gene pool of a population; thereafter, a smaller population, with a smaller genetic diversity, remains to pass on genes to future generations of offspring through sexual reproduction. Genetic diversity remains lower, increasing only when gene flow from another population occurs or very slowly increasing with time as random mutations occur. This results in a reduction in the robustness of the population and in its ability to adapt to and survive selecting environmental changes, such as climate change or a shift in available resources. Alternatively, if survivors of the bottleneck are the individuals with the greatest genetic fitness, the frequency of the fitter genes within the gene pool is increased, while the pool itself is reduced.
The molecular clock is a figurative term for a technique that uses the mutation rate of biomolecules to deduce the time in prehistory when two or more life forms diverged. The biomolecular data used for such calculations are usually nucleotide sequences for DNA, RNA, or amino acid sequences for proteins. The benchmarks for determining the mutation rate are often fossil or archaeological dates. The molecular clock was first tested in 1962 on the hemoglobin protein variants of various animals, and is commonly used in molecular evolution to estimate times of speciation or radiation. It is sometimes called a gene clock or an evolutionary clock.
Archaeogenetics is the study of ancient DNA using various molecular genetic methods and DNA resources. This form of genetic analysis can be applied to human, animal, and plant specimens. Ancient DNA can be extracted from various fossilized specimens including bones, eggshells, and artificially preserved tissues in human and animal specimens. In plants, Ancient DNA can be extracted from seeds and tissue. Archaeogenetics provides us with genetic evidence of ancient population group migrations, domestication events, and plant and animal evolution. The ancient DNA cross referenced with the DNA of relative modern genetic populations allows researchers to run comparison studies that provide a more complete analysis when ancient DNA is compromised.
Behavioral modernity is a suite of behavioral and cognitive traits that distinguishes current Homo sapiens from other anatomically modern humans, hominins, and primates. Most scholars agree that modern human behavior can be characterized by abstract thinking, planning depth, symbolic behavior, music and dance, exploitation of large game, and blade technology, among others. Underlying these behaviors and technological innovations are cognitive and cultural foundations that have been documented experimentally and ethnographically by evolutionary and cultural anthropologists. These human universal patterns include cumulative cultural adaptation, social norms, language, and extensive help and cooperation beyond close kin.
In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The term is also used in reference to the ancestry of groups of genes (haplotypes) rather than organisms.
The Allee effect is a phenomenon in biology characterized by a correlation between population size or density and the mean individual fitness of a population or species.
The effective population size (Ne) is a number that, in some simplified scenarios, corresponds to the number of breeding individuals in the population. More generally, Ne is the number of individuals that an idealised population would need to have in order for some specified quantity of interest to be the same as in the real population. Idealised populations are based on unrealistic but convenient simplifications such as random mating, simultaneous birth of each new generation, constant population size, and equal numbers of children per parent. For most quantities of interest and most real populations, the effective population size Ne is usually smaller than the census population size N of a real population. The same population may have multiple effective population sizes, for different properties of interest, including for different genetic loci.
In population genetics and population ecology, population size is the number of individual organisms in a population. Population size is directly associated with amount of genetic drift, and is the underlying cause of effects like population bottlenecks and the founder effect. Genetic drift is the major source of decrease of genetic diversity within populations which drives fixation and can potentially lead to speciation events.
Coalescent theory is a model of how alleles sampled from a population may have originated from a common ancestor. In the simplest case, coalescent theory assumes no recombination, no natural selection, and no gene flow or population structure, meaning that each variant is equally likely to have been passed from one generation to the next. The model looks backward in time, merging alleles into a single ancestral copy according to a random process in coalescence events. Under this model, the expected time between successive coalescence events increases almost exponentially back in time. Variance in the model comes from both the random passing of alleles from one generation to the next, and the random occurrence of mutations in these alleles.
Human evolutionary genetics studies how one human genome differs from another human genome, the evolutionary past that gave rise to the human genome, and its current effects. Differences between genomes have anthropological, medical, historical and forensic implications and applications. Genetic data can provide important insights into human evolution.
This article lists current estimates of the world population in history. In summary, estimates for the progression of world population since the Late Middle Ages are in the following ranges:
The human mitochondrial molecular clock is the rate at which mutations have been accumulating in the mitochondrial genome of hominids during the course of human evolution. The archeological record of human activity from early periods in human prehistory is relatively limited and its interpretation has been controversial. Because of the uncertainties from the archeological record, scientists have turned to molecular dating techniques in order to refine the timeline of human evolution. A major goal of scientists in the field is to develop an accurate hominid mitochondrial molecular clock which could then be used to confidently date events that occurred during the course of human evolution.
The McDonald–Kreitman test is a statistical test often used by evolutionary and population biologists to detect and measure the amount of adaptive evolution within a species by determining whether adaptive evolution has occurred, and the proportion of substitutions that resulted from positive selection. To do this, the McDonald–Kreitman test compares the amount of variation within a species (polymorphism) to the divergence between species (substitutions) at two types of sites, neutral and nonneutral. A substitution refers to a nucleotide that is fixed within one species, but a different nucleotide is fixed within a second species at the same base pair of homologous DNA sequences. A site is nonneutral if it is either advantageous or deleterious. The two types of sites can be either synonymous or nonsynonymous within a protein-coding region. In a protein-coding sequence of DNA, a site is synonymous if a point mutation at that site would not change the amino acid, also known as a silent mutation. Because the mutation did not result in a change in the amino acid that was originally coded for by the protein-coding sequence, the phenotype, or the observable trait, of the organism is generally unchanged by the silent mutation. A site in a protein-coding sequence of DNA is nonsynonymous if a point mutation at that site results in a change in the amino acid, resulting in a change in the organism's phenotype. Typically, silent mutations in protein-coding regions are used as the "control" in the McDonald–Kreitman test.
A ghost population is a population that has been inferred through using statistical techniques.
The maximum preexpansion population size for the NorthCentral African population is 6,600, the lower bound for the postexpansion population size is 8,400, and the allowed dates are between 49,000 and 640,000 years ago
The relationship between census size and effective size is complex, but arguments based on an island model of migration show that if the effective population size reflects the number of breeding individuals and the effects of population subdivision, then an effective population size of 10,000 is inconsistent with the census size of 500,000 to 1,000,000 that has been suggested by archeological evidence. However, these models have ignored the effects of population extinction and recolonization, which increase the expected variance among demes and reduce the inbreeding effective population size. Using models developed for population extinction and recolonization, we show that a large census size consistent with the multiregional model can be reconciled with an effective population size of 10,000, but genetic variation among demes must be high, reflecting low interdeme migration rates and a colonization process that involves a small number of colonists or kin-structured colonization.
A small human effective population size, on the order of 10,000 individuals, which is smaller than the effective population size of most great apes, has been interpreted as a result of a very long history, starting ? 2 mya, of a small population size, coined as the long-necked bottle model (Harpending et al. 1998; Hawks et al. 2000). Our findings are consistent with this hypothesis, but, depending on the mutation rate, we find either an effective population size of NA = 12,000 (95% C.I. = 9,000–15,500 when averaging over all three demographic models) using the mutation rate calibrated with the human-chimp divergence or an effective population size of NA = 32,500 individuals (95% C.I. = 27,500–34,500) using the mutation rate given by whole-genome trio analysis (The 1000 Genomes Project Consortium 2010) (supplementary figure 4 and table 6, Supplementary Material online). Not surprisingly, the estimated effective mutation rates ? = 4NAµ are comparable for the two mutation rates we considered, and are equal to 1.4 × 10?3/bp/generation (95% C.I. = (1.1–1.7) × 10?3). Relating the estimated effective population size to the census population size during the Pleistocene is a difficult task because there are many factors affecting the effective population size (Charlesworth 2009). Nevertheless, based on published estimates of the ratio between effective and census population size, a comprehensive value on the order of 10% has been found by Frankham (1995). This 10% rule roughly predicts that 120,000–325,[0]00 individuals (depending on the assumed mutation rate)