Misplaced Pages

#10989

156-504: 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 that 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 (a tribe of the African hominid subfamily), indicating that human evolution

312-487: A DNA sequence within a chromosome is known as a locus . If the DNA sequence at a locus varies between individuals, the different forms of this sequence are called alleles. DNA sequences can change through mutations, producing new alleles. If a mutation occurs within a gene, the new allele may affect the trait that the gene controls, altering the phenotype of the organism. However, while this simple correspondence between an allele and

468-570: A GC-biased E. coli mutator strain in 1967, along with the proposal of the neutral theory , established the plausibility of mutational explanations for molecular patterns, which are now common in the molecular evolution literature. For instance, mutation biases are frequently invoked in models of codon usage. Such models also include effects of selection, following the mutation-selection-drift model, which allows both for mutation biases and differential selection based on effects on translation. Hypotheses of mutation bias have played an important role in

624-514: A chromosome becoming duplicated (usually by genetic recombination ), which can introduce extra copies of a gene into a genome. Extra copies of genes are a major source of the raw material needed for new genes to evolve. This is important because most new genes evolve within gene families from pre-existing genes that share common ancestors. For example, the human eye uses four genes to make structures that sense light: three for colour vision and one for night vision ; all four are descended from

780-487: A common ancestor about 660,000 years ago. However, a sequencing of the Neanderthal genome in 2010 indicated that Neanderthals did indeed interbreed with anatomically modern humans c. 45,000-80,000 years ago, around the time modern humans migrated out from Africa, but before they dispersed throughout Europe, Asia and elsewhere. The genetic sequencing of a 40,000-year-old human skeleton from Romania showed that 11% of its genome

936-598: A higher rate of depression. The flow of genes from Neanderthal populations to modern humans was not all one way. Sergi Castellano of the Max Planck Institute for Evolutionary Anthropology reported in 2016 that while Denisovan and Neanderthal genomes are more related to each other than they are to us, Siberian Neanderthal genomes show more similarity to modern human genes than do European Neanderthal populations. This suggests Neanderthal populations interbred with modern humans around 100,000 years ago, probably somewhere in

1092-428: A host of prefixes have been defined to describe various branches of the phylogenetic tree relative to extant organisms. A pan-group or total group is the crown group and all organisms more closely related to it than to any other extant organisms. In a tree analogy, it is the crown group and all branches back to (but not including) the split with the closest branch to have living members. The Pan-Aves thus contain

1248-402: A manner similar to a sequence of letters spelling out a sentence. Before a cell divides, the DNA is copied, so that each of the resulting two cells will inherit the DNA sequence. Portions of a DNA molecule that specify a single functional unit are called genes; different genes have different sequences of bases. Within cells, each long strand of DNA is called a chromosome . The specific location of

1404-485: A number of clear anatomical differences between anatomically modern humans (AMH) and Neanderthal specimens, many relating to the superior Neanderthal adaptation to cold environments. Neanderthal surface to volume ratio was even lower than that among modern Inuit populations, indicating superior retention of body heat. Neanderthals also had significantly larger brains, as shown from brain endocasts, casting doubt on their intellectual inferiority to modern humans. However,

1560-405: A nutrient in a long-term laboratory experiment , Flavobacterium evolving a novel enzyme that allows these bacteria to grow on the by-products of nylon manufacturing, and the soil bacterium Sphingobium evolving an entirely new metabolic pathway that degrades the synthetic pesticide pentachlorophenol . An interesting but still controversial idea is that some adaptations might increase

1716-611: A phenomenon termed de novo gene birth . The generation of new genes can also involve small parts of several genes being duplicated, with these fragments then recombining to form new combinations with new functions ( exon shuffling ). When new genes are assembled from shuffling pre-existing parts, domains act as modules with simple independent functions, which can be mixed together to produce new combinations with new and complex functions. For example, polyketide synthases are large enzymes that make antibiotics ; they contain up to 100 independent domains that each catalyse one step in

SECTION 10

#1732848005011

1872-535: A population are therefore more likely to be replaced by the offspring of parents with favourable characteristics for that environment. In the early 20th century, competing ideas of evolution were refuted and evolution was combined with Mendelian inheritance and population genetics to give rise to modern evolutionary theory. In this synthesis the basis for heredity is in DNA molecules that pass information from generation to generation. The processes that change DNA in

2028-575: A population include natural selection, genetic drift, mutation , and gene flow . All life on Earth—including humanity —shares a last universal common ancestor (LUCA), which lived approximately 3.5–3.8 billion years ago. The fossil record includes a progression from early biogenic graphite to microbial mat fossils to fossilised multicellular organisms . Existing patterns of biodiversity have been shaped by repeated formations of new species ( speciation ), changes within species ( anagenesis ), and loss of species ( extinction ) throughout

2184-430: A population is not critical, but instead a measure known as the effective population size. The effective population is usually smaller than the total population since it takes into account factors such as the level of inbreeding and the stage of the lifecycle in which the population is the smallest. The effective population size may not be the same for every gene in the same population. It is usually difficult to measure

2340-439: A population over successive generations. The process of evolution has given rise to biodiversity at every level of biological organisation . The scientific theory of evolution by natural selection was conceived independently by two British naturalists, Charles Darwin and Alfred Russel Wallace , in the mid-19th century as an explanation for why organisms are adapted to their physical and biological environments. The theory

2496-433: A population through neutral transitions due to the principles of excess capacity, presuppression, and ratcheting, and it has been applied in areas ranging from the origins of the spliceosome to the complex interdependence of microbial communities . The time it takes a neutral allele to become fixed by genetic drift depends on population size; fixation is more rapid in smaller populations. The number of individuals in

2652-499: A range of genes from bacteria, fungi and plants. Viruses can also carry DNA between organisms, allowing transfer of genes even across biological domains . Large-scale gene transfer has also occurred between the ancestors of eukaryotic cells and bacteria, during the acquisition of chloroplasts and mitochondria . It is possible that eukaryotes themselves originated from horizontal gene transfers between bacteria and archaea . Some heritable changes cannot be explained by changes to

2808-421: A recent species of the genus Homo that exhibits derived traits not shared with modern humans. In other words, H. floresiensis shares a common ancestor with modern humans, but split from the modern human lineage and followed a distinct evolutionary path. The main find was a skeleton believed to be a woman of about 30 years of age. Found in 2003, it has been dated to approximately 18,000 years old. The living woman

2964-430: A reduction in scope when it was discovered that (1) GC-biased gene conversion makes an important contribution to composition in diploid organisms such as mammals and (2) bacterial genomes frequently have AT-biased mutation. Contemporary thinking about the role of mutation biases reflects a different theory from that of Haldane and Fisher. More recent work showed that the original "pressures" theory assumes that evolution

3120-472: A related lineage. In September 2019, scientists reported the computerized determination, based on 260 CT scans , of a virtual skull shape of the last common human ancestor to modern humans ( H. sapiens ), representative of the earliest modern humans, and suggested that modern humans arose between 260,000 and 350,000 years ago through a merging of populations in East and South Africa. Between 400,000 years ago and

3276-452: A single ancestral gene. New genes can be generated from an ancestral gene when a duplicate copy mutates and acquires a new function. This process is easier once a gene has been duplicated because it increases the redundancy of the system; one gene in the pair can acquire a new function while the other copy continues to perform its original function. Other types of mutations can even generate entirely new genes from previously noncoding DNA,

SECTION 20

#1732848005011

3432-419: A single chromosome compared to expectations , which is called their linkage disequilibrium . A set of alleles that is usually inherited in a group is called a haplotype . This can be important when one allele in a particular haplotype is strongly beneficial: natural selection can drive a selective sweep that will also cause the other alleles in the haplotype to become more common in the population; this effect

3588-420: A species or population, in particular shifts in allele frequency and adaptation. Macroevolution is the outcome of long periods of microevolution. Thus, the distinction between micro- and macroevolution is not a fundamental one—the difference is simply the time involved. However, in macroevolution, the traits of the entire species may be important. For instance, a large amount of variation among individuals allows

3744-544: A species to rapidly adapt to new habitats , lessening the chance of it going extinct, while a wide geographic range increases the chance of speciation, by making it more likely that part of the population will become isolated. In this sense, microevolution and macroevolution might involve selection at different levels—with microevolution acting on genes and organisms, versus macroevolutionary processes such as species selection acting on entire species and affecting their rates of speciation and extinction. A common misconception

3900-432: A stem group allows the order of these acquisitions to be established, and thus the ecological and functional setting of the evolution of the major features of the group in question. Stem groups thus offer a route to integrate unique palaeontological data into questions of the evolution of living organisms. Furthermore, they show that fossils that were considered to lie in their own separate group because they did not show all

4056-411: A striking example are people with the inherited trait of albinism , who do not tan at all and are very sensitive to sunburn . Heritable characteristics are passed from one generation to the next via DNA , a molecule that encodes genetic information. DNA is a long biopolymer composed of four types of bases. The sequence of bases along a particular DNA molecule specifies the genetic information, in

4212-468: A study of the life history of Ar. ramidus that the species provides evidence for a suite of anatomical and behavioral adaptations in very early hominins unlike any species of extant great ape. This study demonstrated affinities between the skull morphology of Ar. ramidus and that of infant and juvenile chimpanzees, suggesting the species evolved a juvenalised or paedomorphic craniofacial morphology via heterochronic dissociation of growth trajectories. It

4368-472: A system in which organisms interact with every other element, physical as well as biological , in their local environment. Eugene Odum , a founder of ecology, defined an ecosystem as: "Any unit that includes all of the organisms...in a given area interacting with the physical environment so that a flow of energy leads to clearly defined trophic structure, biotic diversity, and material cycles (i.e., exchange of materials between living and nonliving parts) within

4524-428: A trait works in some cases, most traits are influenced by multiple genes in a quantitative or epistatic manner. Evolution can occur if there is genetic variation within a population. Variation comes from mutations in the genome, reshuffling of genes through sexual reproduction and migration between populations ( gene flow ). Despite the constant introduction of new variation through mutation and gene flow, most of

4680-590: Is Kamoyapithecus from the uppermost Oligocene at Eragaleit in the northern Great Rift Valley in Kenya, dated to 24 million years ago. Its ancestry is thought to be species related to Aegyptopithecus , Propliopithecus , and Parapithecus from the Faiyum, at around 35 mya. In 2010, Saadanius was described as a close relative of the last common ancestor of the crown catarrhines, and tentatively dated to 29–28 mya, helping to fill an 11-million-year gap in

4836-454: Is directional selection , which is a shift in the average value of a trait over time—for example, organisms slowly getting taller. Secondly, disruptive selection is selection for extreme trait values and often results in two different values becoming most common, with selection against the average value. This would be when either short or tall organisms had an advantage, but not those of medium height. Finally, in stabilising selection there

Human evolution - Misplaced Pages Continue

4992-442: Is more noticeable . Indeed, the evolution of microorganisms is particularly important to evolutionary research since their rapid reproduction allows the study of experimental evolution and the observation of evolution and adaptation in real time. Adaptation is the process that makes organisms better suited to their habitat. Also, the term adaptation may refer to a trait that is important for an organism's survival. For example,

5148-427: Is a byproduct of this process that may sometimes be adaptively beneficial. Gene flow is the exchange of genes between populations and between species. It can therefore be a source of variation that is new to a population or to a species. Gene flow can be caused by the movement of individuals between separate populations of organisms, as might be caused by the movement of mice between inland and coastal populations, or

5304-417: Is an inherited characteristic and an individual might inherit the "brown-eye trait" from one of their parents. Inherited traits are controlled by genes and the complete set of genes within an organism's genome (genetic material) is called its genotype . The complete set of observable traits that make up the structure and behaviour of an organism is called its phenotype . Some of these traits come from

5460-464: Is based on standing variation: when evolution depends on events of mutation that introduce new alleles, mutational and developmental biases in the introduction of variation (arrival biases) can impose biases on evolution without requiring neutral evolution or high mutation rates. Several studies report that the mutations implicated in adaptation reflect common mutation biases though others dispute this interpretation. Recombination allows alleles on

5616-407: Is called deep homology . During evolution, some structures may lose their original function and become vestigial structures. Such structures may have little or no function in a current species, yet have a clear function in ancestral species, or other closely related species. Examples include pseudogenes , the non-functional remains of eyes in blind cave-dwelling fish, wings in flightless birds,

5772-692: Is called genetic hitchhiking or genetic draft. Genetic draft caused by the fact that some neutral genes are genetically linked to others that are under selection can be partially captured by an appropriate effective population size. A special case of natural selection is sexual selection, which is selection for any trait that increases mating success by increasing the attractiveness of an organism to potential mates. Traits that evolved through sexual selection are particularly prominent among males of several animal species. Although sexually favoured, traits such as cumbersome antlers, mating calls, large body size and bright colours often attract predation, which compromises

5928-443: Is claimed to have been discovered living at the same time period of A. afarensis . There is debate whether A. deyiremeda is a new species or is A. afarensis . Australopithecus prometheus , otherwise known as Little Foot has recently been dated at 3.67 million years old through a new dating technique, making the genus Australopithecus as old as afarensis . Given the opposable big toe found on Little Foot, it seems that

6084-454: Is consistent with recent studies indicating that the divergence of some human alleles dates to one Ma, although this interpretation has been questioned. Neanderthals and AMH Homo sapiens could have co-existed in Europe for as long as 10,000 years, during which AMH populations exploded, vastly outnumbering Neanderthals, possibly outcompeting them by sheer numbers. In 2008, archaeologists working at

6240-685: Is evidence of a wide diversity of forms across Africa and the Mediterranean basin during the relatively warm and equable climatic regimes of the Early and Middle Miocene. The youngest of the Miocene hominoids, Oreopithecus , is from coal beds in Italy that have been dated to 9 million years ago. Molecular evidence indicates that the lineage of gibbons diverged from the line of great apes some 18–12 mya, and that of orangutans (subfamily Ponginae ) diverged from

6396-405: Is limited; both poor preservation – rain forest soils tend to be acidic and dissolve bone – and sampling bias probably contribute to this problem. Other hominins probably adapted to the drier environments outside the equatorial belt; and there they encountered antelope, hyenas, dogs, pigs, elephants, horses, and others. The equatorial belt contracted after about 8 million years ago, and there

Human evolution - Misplaced Pages Continue

6552-594: Is measured by an organism's ability to survive and reproduce, which determines the size of its genetic contribution to the next generation. However, fitness is not the same as the total number of offspring: instead fitness is indicated by the proportion of subsequent generations that carry an organism's genes. For example, if an organism could survive well and reproduce rapidly, but its offspring were all too small and weak to survive, this organism would make little genetic contribution to future generations and would thus have low fitness. If an allele increases fitness more than

6708-618: Is no consensus phylogeny. Stem arthropods constitute a group that has seen attention in connection with the Burgess Shale fauna. Several of the finds , including the enigmatic Opabinia and Anomalocaris have some, though not all, features associated with arthropods , and are thus considered stem arthropods. The sorting of the Burgess Shale fauna into various stem groups finally enabled phylogenetic sorting of this enigmatic assemblage and also allowed for identifying velvet worms as

6864-693: Is seen most completely in the Upper Eocene and lowermost Oligocene fossil beds of the Faiyum depression southwest of Cairo —gave rise to all extant primate species, including the lemurs of Madagascar , lorises of Southeast Asia, galagos or "bush babies" of Africa, and to the anthropoids , which are the Platyrrhines or New World monkeys, the Catarrhines or Old World monkeys, and the great apes, including humans and other hominids. The earliest known catarrhine

7020-421: Is selection against extreme trait values on both ends, which causes a decrease in variance around the average value and less diversity. This would, for example, cause organisms to eventually have a similar height. Natural selection most generally makes nature the measure against which individuals and individual traits, are more or less likely to survive. "Nature" in this sense refers to an ecosystem , that is,

7176-425: Is shown below: † Archaeopteryx other extinct groups Neornithes (modern birds, some extinct like the dodo) In this diagram, the clade labelled "Neornithes" is the crown group of birds: it includes the most recent common ancestor of all living birds and its descendants, living or not. Although considered to be birds (i.e. members of the clade Aves), Archaeopteryx and other extinct groups are not included in

7332-518: Is still some debate among academics whether certain African hominid species of this time, such as A. robustus and A. boisei , constitute members of the same genus; if so, they would be considered to be "robust australopiths" while the others would be considered "gracile australopiths". However, if these species do indeed constitute their own genus, then they may be given their own name, Paranthropus . A new proposed species Australopithecus deyiremeda

7488-686: Is supported in part, because some modern humans who live on Flores , the Indonesian island where the skeleton was found, are pygmies . This, coupled with pathological dwarfism, could have resulted in a significantly diminutive human. The other major attack on H. floresiensis as a separate species is that it was found with tools only associated with H. sapiens . The hypothesis of pathological dwarfism, however, fails to explain additional anatomical features that are unlike those of modern humans (diseased or not) but much like those of ancient members of our genus. Aside from cranial features, these features include

7644-429: Is that evolution has goals, long-term plans, or an innate tendency for "progress", as expressed in beliefs such as orthogenesis and evolutionism; realistically, however, evolution has no long-term goal and does not necessarily produce greater complexity. Although complex species have evolved, they occur as a side effect of the overall number of organisms increasing, and simple forms of life still remain more common in

7800-447: Is the nearly neutral theory , according to which a mutation that would be effectively neutral in a small population is not necessarily neutral in a large population. Other theories propose that genetic drift is dwarfed by other stochastic forces in evolution, such as genetic hitchhiking, also known as genetic draft. Another concept is constructive neutral evolution (CNE), which explains that complex systems can emerge and spread into

7956-495: Is the more common means of reproduction among eukaryotes and multicellular organisms. The Red Queen hypothesis has been used to explain the significance of sexual reproduction as a means to enable continual evolution and adaptation in response to coevolution with other species in an ever-changing environment. Another hypothesis is that sexual reproduction is primarily an adaptation for promoting accurate recombinational repair of damage in germline DNA, and that increased diversity

SECTION 50

#1732848005011

8112-439: Is the process by which traits that enhance survival and reproduction become more common in successive generations of a population. It embodies three principles: More offspring are produced than can possibly survive, and these conditions produce competition between organisms for survival and reproduction. Consequently, organisms with traits that give them an advantage over their competitors are more likely to pass on their traits to

8268-453: Is thought by some to make the Cambrian explosion easier to understand without invoking unusual evolutionary mechanisms; however, application of the stem group concept does nothing to ameliorate the difficulties that phylogenetic telescoping poses to evolutionary theorists attempting to understand both macroevolutionary change and the abrupt character of the Cambrian explosion . Overemphasis on

8424-408: Is very little fossil evidence for the split—thought to have occurred around that time—of the hominin lineage from the lineages of gorillas and chimpanzees. The earliest fossils argued by some to belong to the human lineage are Sahelanthropus tchadensis (7 Ma) and Orrorin tugenensis (6 Ma), followed by Ardipithecus (5.5–4.4 Ma), with species Ar. kadabba and Ar. ramidus . It has been argued in

8580-610: The Gorillini tribe between 8 and 9 mya; Australopithecine (including the extinct biped ancestors of humans) separated from the Pan genus (containing chimpanzees and bonobos ) 4–7 mya. The Homo genus is evidenced by the appearance of H. habilis over 2 mya, while anatomically modern humans emerged in Africa approximately 300,000 years ago. The evolutionary history of primates can be traced back 65 million years. One of

8736-604: The Late Cretaceous period, with their earliest fossils appearing over 55 mya, during the Paleocene . Primates produced successive clades leading to the ape superfamily, which gave rise to the hominid and the gibbon families; these diverged some 15–20 mya. African and Asian hominids (including orangutans ) diverged about 14 mya. Hominins (including the Australopithecine and Panina subtribes) parted from

8892-486: The Late Pliocene or Early Pleistocene , 2.5–2 Ma, when it diverged from the australopithecines with the development of smaller molars and larger brains. One of the first known hominins, it made tools from stone and perhaps animal bones, leading to its name homo habilis (Latin 'handy man') bestowed by discoverer Louis Leakey . Some scientists have proposed moving this species from Homo into Australopithecus due to

9048-459: The Pleistocene played a significant role in human evolution observed via human transitions between subsistence systems. The most significant of these adaptations are bipedalism, increased brain size, lengthened ontogeny (gestation and infancy), and decreased sexual dimorphism . The relationship between these changes is the subject of ongoing debate. Other significant morphological changes included

9204-405: The dinosaurs and the pterosaurs . The last common ancestor of birds and crocodilians—the first crown group archosaur—was neither bird nor crocodilian and possessed none of the features unique to either. As the bird stem group evolved, distinctive bird features such as feathers and hollow bones appeared. Finally, at the base of the crown group, all traits common to extant birds were present. Under

9360-464: The fossil record , cranial capacity had doubled to 850 cm. (Such an increase in human brain size is equivalent to each generation having 125,000 more neurons than their parents.) It is believed that H. erectus and H. ergaster were the first to use fire and complex tools, and were the first of the hominin line to leave Africa, spreading throughout Africa, Asia, and Europe between 1.3 to 1.8 million years ago . According to

9516-580: The gorillas and chimpanzees, diverged from the hominin line over a period covering the same time, so either Sahelanthropus or Orrorin may be our last shared ancestor. Ardipithecus , a full biped, arose approximately 5.6 million years ago. Evolution Evolution is the change in the heritable characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, resulting in certain characteristics becoming more or less common within

SECTION 60

#1732848005011

9672-466: The last common ancestor of the crown group and their closest living relatives. It follows from the definition that all members of a stem group are extinct. The "stem group" is the most used and most important of the concepts linked to crown groups, as it offers a means to reify and name paraphyletic assemblages of fossils that otherwise do not fit into systematics based on living organisms. While often attributed to Jefferies (1979), Willmann (2003) traced

9828-426: The lungfish , our nearest relatives among the fishes. In addition to a series of lobe-finned fishes , they also include some of the early labyrinthodonts . Exactly what labyrinthodonts are in the stem group tetrapods rather than the corresponding crown group is uncertain, as the phylogeny of early tetrapods is not well understood. This example shows that crown and stem group definitions are of limited value when there

9984-425: The neutral theory of molecular evolution most evolutionary changes are the result of the fixation of neutral mutations by genetic drift. In this model, most genetic changes in a population are thus the result of constant mutation pressure and genetic drift. This form of the neutral theory has been debated since it does not seem to fit some genetic variation seen in nature. A better-supported version of this model

10140-454: The product of a gene , or prevent the gene from functioning, or have no effect. About half of the mutations in the coding regions of protein-coding genes are deleterious — the other half are neutral. A small percentage of the total mutations in this region confer a fitness benefit. Some of the mutations in other parts of the genome are deleterious but the vast majority are neutral. A few are beneficial. Mutations can involve large sections of

10296-400: The "crown" and "stem" group terminology was coined by R. P. S. Jefferies in 1979. Though formulated in the 1970s, the term was not commonly used until its reintroduction in 2000 by Graham Budd and Sören Jensen . It is not necessary for a species to have living descendants in order for it to be included in the crown group. Extinct side branches on the family tree that are descended from

10452-515: The Australopithecine and Panina subtribes) parted from the Gorillini tribe (gorillas) between 8 and 9 mya; Australopithecine (including the extinct biped ancestors of humans) separated from the Pan genus (containing chimpanzees and bonobos) 4–7 mya. The Homo genus is evidenced by the appearance of H. habilis over 2 mya, while anatomically modern humans emerged in Africa approximately 300,000 years ago. Species close to

10608-521: The Crocodilia branch. Basal branch names such as Avemetatarsalia are usually more obscure. However, not so advantageous are the facts that "Pan-Aves" and "Aves" are not the same group, the circumscription of the concept of "Pan-Aves" (synonymous with Avemetatarsalia) is only evident by examination of the above tree, and calling both groups "birds" is ambiguous. Stem mammals are those in the lineage leading to living mammals, together with side branches, from

10764-824: The Early Pleistocene, 1.5–1 Ma, some populations of Homo habilis are thought to have evolved larger brains and to have made more elaborate stone tools; these differences and others are sufficient for anthropologists to classify them as a new species, Homo erectus —in Africa. The evolution of locking knees and the movement of the foramen magnum are thought to be likely drivers of the larger population changes. This species also may have used fire to cook meat. Richard Wrangham notes that Homo seems to have been ground dwelling, with reduced intestinal length, smaller dentition, and "brains [swollen] to their current, horrendously fuel-inefficient size", and hypothesizes that control of fire and cooking, which released increased nutritional value,

10920-513: The Indonesian island of Java. He originally named the material Anthropopithecus erectus (1892–1893, considered at this point as a chimpanzee-like fossil primate) and Pithecanthropus erectus (1893–1894, changing his mind as of based on its morphology, which he considered to be intermediate between that of humans and apes). Years later, in the 20th century, the German physician and paleoanthropologist Franz Weidenreich (1873–1948) compared in detail

11076-513: The Near East. Studies of a Neanderthal child at Gibraltar show from brain development and tooth eruption that Neanderthal children may have matured more rapidly than Homo sapiens . H. floresiensis , which lived from approximately 190,000 to 50,000 years before present (BP), has been nicknamed the hobbit for its small size, possibly a result of insular dwarfism . H. floresiensis is intriguing both for its size and its age, being an example of

11232-651: The Toba catastrophe; however, nearby H. floresiensis survived it. The early phase of H. erectus , from 1.8 to 1.25 Ma, is considered by some to be a separate species, H. ergaster , or as H. erectus ergaster , a subspecies of H. erectus . Many paleoanthropologists now use the term Homo ergaster for the non-Asian forms of this group, and reserve H. erectus only for those fossils that are found in Asia and meet certain skeletal and dental requirements which differ slightly from H. ergaster . In Africa in

11388-519: The ability of organisms to generate genetic diversity and adapt by natural selection (increasing organisms' evolvability). Adaptation occurs through the gradual modification of existing structures. Consequently, structures with similar internal organisation may have different functions in related organisms. This is the result of a single ancestral structure being adapted to function in different ways. The bones within bat wings, for example, are very similar to those in mice feet and primate hands, due to

11544-468: The absence of aggressive canine morphology in Ar. ramidus and the implications this has for the evolution of hominin social psychology, they wrote: Of course Ar. ramidus differs significantly from bonobos, bonobos having retained a functional canine honing complex. However, the fact that Ar. ramidus shares with bonobos reduced sexual dimorphism, and a more paedomorphic form relative to chimpanzees, suggests that

11700-402: The adaptation of horses' teeth to the grinding of grass. By using the term adaptation for the evolutionary process and adaptive trait for the product (the bodily part or function), the two senses of the word may be distinguished. Adaptations are produced by natural selection. The following definitions are due to Theodosius Dobzhansky: Adaptation may cause either the gain of a new feature, or

11856-507: The alleles are subject to sampling error . This drift halts when an allele eventually becomes fixed, either by disappearing from the population or by replacing the other alleles entirely. Genetic drift may therefore eliminate some alleles from a population due to chance alone. Even in the absence of selective forces, genetic drift can cause two separate populations that begin with the same genetic structure to drift apart into two divergent populations with different sets of alleles. According to

12012-436: The ancestral hominin line. There is yet no consensus as to which of these groups should be considered a separate species and which should be subspecies; this may be due to the dearth of fossils or to the slight differences used to classify species in the genus Homo . The Sahara pump theory (describing an occasionally passable "wet" Sahara desert) provides one possible explanation of the intermittent migration and speciation in

12168-405: The ancient forest and woodland ecosystems of late Miocene and early Pliocene Africa. Consequently, they argue that humans may not represent evolution from a chimpanzee-like ancestor as has traditionally been supposed. This suggests many modern human adaptations represent phylogenetically deep traits and that the behavior and morphology of chimpanzees may have evolved subsequent to the split with

12324-441: The biosphere. For example, the overwhelming majority of species are microscopic prokaryotes , which form about half the world's biomass despite their small size and constitute the vast majority of Earth's biodiversity. Simple organisms have therefore been the dominant form of life on Earth throughout its history and continue to be the main form of life up to the present day, with complex life only appearing more diverse because it

12480-488: The change over time in this genetic variation. The frequency of one particular allele will become more or less prevalent relative to other forms of that gene. Variation disappears when a new allele reaches the point of fixation —when it either disappears from the population or replaces the ancestral allele entirely. Mutations are changes in the DNA sequence of a cell's genome and are the ultimate source of genetic variation in all organisms. When mutations occur, they may alter

12636-563: The characters of Dubois' Java Man , then named Pithecanthropus erectus , with the characters of the Peking Man , then named Sinanthropus pekinensis . Weidenreich concluded in 1940 that because of their anatomical similarity with modern humans it was necessary to gather all these specimens of Java and China in a single species of the genus Homo , the species H. erectus . Homo erectus lived from about 1.8 Ma to about 70,000 years ago – which would indicate that they were probably wiped out by

12792-514: The closest living relatives of arthropods. Stem priapulids are other early Cambrian to middle Cambrian faunas, appearing in Chengjiang to Burgess Shale. The genus Ottoia has more or less the same build as modern priapulids , but phylogenetic analysis indicates that it falls outside the crown group, making it a stem priapulid. The name plesion has a long history in biological systematics, and plesion group has acquired several meanings over

12948-401: The common ancestor they share with humans. The genus Australopithecus evolved in eastern Africa around 4 million years ago before spreading throughout the continent and eventually becoming extinct 2 million years ago. During this time period various forms of australopiths existed, including Australopithecus anamensis , A. afarensis , A. sediba , and A. africanus . There

13104-582: The crown group, as they fall outside the Neornithes clade, being descended from an earlier ancestor. An alternative definition does not require any members of a crown group to be extant, only to have resulted from a "major cladogenesis event". The first definition forms the basis of this article. Often, the crown group is given the designation "crown-", to separate it from the group as commonly defined. Both birds and mammals are traditionally defined by their traits, and contain fossil members that lived before

13260-403: The descent of all these structures from a common mammalian ancestor. However, since all living organisms are related to some extent, even organs that appear to have little or no structural similarity, such as arthropod , squid and vertebrate eyes, or the limbs and wings of arthropods and vertebrates, can depend on a common set of homologous genes that control their assembly and function; this

13416-400: The development of thinking about the evolution of genome composition, including isochores. Different insertion vs. deletion biases in different taxa can lead to the evolution of different genome sizes. The hypothesis of Lynch regarding genome size relies on mutational biases toward increase or decrease in genome size. However, mutational hypotheses for the evolution of composition suffered

13572-406: The developmental and social adaptations evident in bonobos may be of assistance in future reconstructions of early hominin social and sexual psychology. In fact the trend towards increased maternal care, female mate selection and self-domestication may have been stronger and more refined in Ar. ramidus than what we see in bonobos. The authors argue that many of the basic human adaptations evolved in

13728-483: The diagnostic features of a living clade, can nevertheless be related to it by lying in its stem group. Such fossils have been of particular importance in considering the origins of the tetrapods , mammals , and animals . The application of the stem group concept also influenced the interpretation of the organisms of the Burgess shale . Their classification in stem groups to extant phyla, rather than in phyla of their own,

13884-490: The direct control of genes include the inheritance of cultural traits and symbiogenesis . From a neo-Darwinian perspective, evolution occurs when there are changes in the frequencies of alleles within a population of interbreeding organisms, for example, the allele for black colour in a population of moths becoming more common. Mechanisms that can lead to changes in allele frequencies include natural selection, genetic drift, and mutation bias. Evolution by natural selection

14040-401: The distribution of stone tools), and faster spread of social and technological innovations. All these may have all contributed to modern Homo sapiens replacing Neanderthal populations by 28,000 BP. Earlier evidence from sequencing mitochondrial DNA suggested that no significant gene flow occurred between H. neanderthalensis and H. sapiens , and that the two were separate species that shared

14196-509: The divergence of the lineage from the Sauropsida to the last common ancestor of the living mammals. This group includes the synapsids as well as mammaliaforms like the morganucodonts and the docodonts ; the latter groups have traditionally and anatomically been considered mammals even though they fall outside the crown group mammals. Stem tetrapods are the animals belonging to the lineage leading to tetrapods from their divergence from

14352-469: The divergence point of the mtDNA was unexpectedly deep in time, the full genomic sequence suggested the Denisovans belonged to the same lineage as Neanderthals, with the two diverging shortly after their line split from the lineage that gave rise to modern humans. Modern humans are known to have overlapped with Neanderthals in Europe and the Near East for possibly more than 40,000 years, and the discovery raises

14508-465: The earlier dispersed H. erectus . This migration and origin theory is usually referred to as the "recent single-origin hypothesis" or "out of Africa" theory. H. sapiens interbred with archaic humans both in Africa and in Eurasia, in Eurasia notably with Neanderthals and Denisovans. The Toba catastrophe theory , which postulates a population bottleneck for H. sapiens about 70,000 years ago,

14664-482: The evolution of a power and precision grip , a change first occurring in H. erectus . Bipedalism , (walking on two legs), is the basic adaptation of the hominid and is considered the main cause behind a suite of skeletal changes shared by all bipedal hominids. The earliest hominin, of presumably primitive bipedalism, is considered to be either Sahelanthropus or Orrorin , both of which arose some 6 to 7 million years ago. The non-bipedal knuckle-walkers,

14820-489: The evolutionary history of life on Earth. Morphological and biochemical traits tend to be more similar among species that share a more recent common ancestor , which historically was used to reconstruct phylogenetic trees , although direct comparison of genetic sequences is a more common method today. Evolutionary biologists have continued to study various aspects of evolution by forming and testing hypotheses as well as constructing theories based on evidence from

14976-505: The extinct moa ) The crown group here is Neornithes , all modern bird lineages back to their last common ancestor. The closest living relatives of birds are crocodilians . If we follow the phylogenetic lineage leading to Neornithes to the left, the line itself and all side branches belong to the stem birds until the lineage merges with that of the crocodilians. In addition to non-crown group primitive birds like Archaeopteryx , Hesperornis and Confuciusornis , stem group birds include

15132-414: The field or laboratory and on data generated by the methods of mathematical and theoretical biology . Their discoveries have influenced not just the development of biology but also other fields including agriculture, medicine, and computer science . Evolution in organisms occurs through changes in heritable characteristics—the inherited characteristics of an organism. In humans, for example, eye colour

15288-427: The fitness of an allele is not a fixed characteristic; if the environment changes, previously neutral or harmful traits may become beneficial and previously beneficial traits become harmful. However, even if the direction of selection does reverse in this way, traits that were lost in the past may not re-evolve in an identical form. However, a re-activation of dormant genes, as long as they have not been eliminated from

15444-536: The forerunner of anatomically modern humans , evolved in the Middle Paleolithic between 400,000 and 250,000 years ago. Recent DNA evidence suggests that several haplotypes of Neanderthal origin are present among all non-African populations, and Neanderthals and other hominins, such as Denisovans, may have contributed up to 6% of their genome to present-day humans, suggestive of a limited interbreeding between these species . According to some anthropologists,

15600-591: The form of bones in the wrist, forearm, shoulder, knees, and feet. Additionally, this hypothesis fails to explain the find of multiple examples of individuals with these same characteristics, indicating they were common to a large population, and not limited to one individual. In 2016, fossil teeth and a partial jaw from hominins assumed to be ancestral to H. floresiensis were discovered at Mata Menge , about 74 km (46 mi) from Liang Bua. They date to about 700,000 years ago and are noted by Australian archaeologist Gerrit van den Bergh for being even smaller than

15756-989: The fossil record. In the Early Miocene , about 22 million years ago, the many kinds of arboreally -adapted (tree-dwelling) primitive catarrhines from East Africa suggest a long history of prior diversification. Fossils at 20 million years ago include fragments attributed to Victoriapithecus , the earliest Old World monkey. Among the genera thought to be in the ape lineage leading up to 13 million years ago are Proconsul , Rangwapithecus , Dendropithecus , Limnopithecus , Nacholapithecus , Equatorius , Nyanzapithecus , Afropithecus , Heliopithecus , and Kenyapithecus , all from East Africa. The presence of other generalized non-cercopithecids of Middle Miocene from sites far distant, such as Otavipithecus from cave deposits in Namibia, and Pierolapithecus and Dryopithecus from France, Spain and Austria,

15912-432: The full bifurcating phylogeny. Stem birds perhaps constitute the most cited example of a stem group, as the phylogeny of this group is fairly well known. The following cladogram, based on Benton (2005), illustrates the concept: Crocodilia Pterosauria Hadrosauridae Stegosauria Sauropoda Tyrannosauridae Archaeopteryx Neognathae (including the extinct dodo ) Paleognathae (including

16068-472: The genome and were only suppressed perhaps for hundreds of generations, can lead to the re-occurrence of traits thought to be lost like hindlegs in dolphins, teeth in chickens, wings in wingless stick insects, tails and additional nipples in humans etc. "Throwbacks" such as these are known as atavisms . Natural selection within a population for a trait that can vary across a range of values, such as height, can be categorised into three different types. The first

16224-479: The genome of a species is very similar among all individuals of that species. However, discoveries in the field of evolutionary developmental biology have demonstrated that even relatively small differences in genotype can lead to dramatic differences in phenotype both within and between species. An individual organism's phenotype results from both its genotype and the influence of the environment it has lived in. The modern evolutionary synthesis defines evolution as

16380-529: The genomes of modern humans outside Africa. HLA haplotypes from Denisovans and Neanderthal represent more than half the HLA alleles of modern Eurasians, indicating strong positive selection for these introgressed alleles. Corinne Simoneti at Vanderbilt University, in Nashville and her team have found from medical records of 28,000 people of European descent that the presence of Neanderthal DNA segments may be associated with

16536-621: The genus Homo . Based on archaeological and paleontological evidence, it has been possible to infer, to some extent, the ancient dietary practices of various Homo species and to study the role of diet in physical and behavioral evolution within Homo . Some anthropologists and archaeologists subscribe to the Toba catastrophe theory , which posits that the supereruption of Lake Toba on Sumatra in Indonesia some 70,000 years ago caused global starvation, killing

16692-817: The higher body mass of Neanderthals may have required larger brain mass for body control. Also, recent research by Pearce, Stringer , and Dunbar has shown important differences in brain architecture. The larger size of the Neanderthal orbital chamber and occipital lobe suggests that they had a better visual acuity than modern humans, useful in the dimmer light of glacial Europe. Neanderthals may have had less brain capacity available for social functions . Inferring social group size from endocranial volume (minus occipital lobe size) suggests that Neanderthal groups may have been limited to 120 individuals, compared to 144 possible relationships for modern humans. Larger social groups could imply that modern humans had less risk of inbreeding within their clan, trade over larger areas (confirmed in

16848-640: The idea of developmental bias . Haldane and Fisher argued that, because mutation is a weak pressure easily overcome by selection, tendencies of mutation would be ineffectual except under conditions of neutral evolution or extraordinarily high mutation rates. This opposing-pressures argument was long used to dismiss the possibility of internal tendencies in evolution, until the molecular era prompted renewed interest in neutral evolution. Noboru Sueoka and Ernst Freese proposed that systematic biases in mutation might be responsible for systematic differences in genomic GC composition between species. The identification of

17004-450: The interaction of its genotype with the environment while others are neutral. Some observable characteristics are not inherited. For example, suntanned skin comes from the interaction between a person's genotype and sunlight; thus, suntans are not passed on to people's children. The phenotype is the ability of the skin to tan when exposed to sunlight. However, some people tan more easily than others, due to differences in genotypic variation;

17160-598: The last common ancestor of gorillas, chimpanzees and humans may be represented by Nakalipithecus fossils found in Kenya and Ouranopithecus found in Greece. Molecular evidence suggests that between 8 and 4 million years ago, first the gorillas, and then the chimpanzees (genus Pan ) split off from the line leading to the humans. Human DNA is approximately 98.4% identical to that of chimpanzees when comparing single nucleotide polymorphisms (see human evolutionary genetics ). The fossil record, however, of gorillas and chimpanzees

17316-522: The last common ancestors of the living groups or, like the mammal Haldanodon , were not descended from that ancestor although they lived later. Crown-Aves and Crown-Mammalia therefore differ slightly in content from the common definition of Aves and Mammalia. This has caused some confusion in the literature. The cladistic idea of strictly using the topology of the phylogenetic tree to define groups necessitates other definitions than crown groups to adequately define commonly discussed fossil groups. Thus,

17472-411: The later fossils. A small number of specimens from the island of Luzon , dated 50,000 to 67,000 years ago, have recently been assigned by their discoverers, based on dental characteristics, to a novel human species, H. luzonensis . H. sapiens (the adjective sapiens is Latin for "wise" or "intelligent") emerged in Africa around 300,000 years ago, likely derived from H. heidelbergensis or

17628-512: The level of the individual organism are genes called transposons , which can replicate and spread throughout a genome. Selection at a level above the individual, such as group selection , may allow the evolution of cooperation. Genetic drift is the random fluctuation of allele frequencies within a population from one generation to the next. When selective forces are absent or relatively weak, allele frequencies are equally likely to drift upward or downward in each successive generation because

17784-628: The living birds and all (fossil) organisms more closely related to birds than to crocodilians (their closest living relatives). The phylogenetic lineage leading back from Neornithes to the point where it merges with the crocodilian lineage, along with all side branches, constitutes pan-birds. In addition to non-crown group primitive birds like Archaeopteryx , Hesperornis and Confuciusornis , therefore, pan-group birds would include all dinosaurs and pterosaurs as well as an assortment of non-crocodilian animals like Marasuchus . Pan-Mammalia consists of all mammals and their fossil ancestors back to

17940-452: The longer term, evolution produces new species through splitting ancestral populations of organisms into new groups that cannot or will not interbreed. These outcomes of evolution are distinguished based on time scale as macroevolution versus microevolution. Macroevolution refers to evolution that occurs at or above the level of species, in particular speciation and extinction, whereas microevolution refers to smaller evolutionary changes within

18096-405: The loss of an ancestral feature. An example that shows both types of change is bacterial adaptation to antibiotic selection, with genetic changes causing antibiotic resistance by both modifying the target of the drug, or increasing the activity of transporters that pump the drug out of the cell. Other striking examples are the bacteria Escherichia coli evolving the ability to use citric acid as

18252-589: The majority of humans and creating a population bottleneck that affected the genetic inheritance of all humans today. The genetic and archaeological evidence for this remains in question however. A 2023 genetic study suggests that a similar human population bottleneck of between 1,000 and 100,000 survivors occurred "around 930,000 and 813,000 years ago ... lasted for about 117,000 years and brought human ancestors close to extinction." Homo habilis lived from about 2.8 to 1.4 Ma. The species evolved in South and East Africa in

18408-458: The mechanics in developmental plasticity and canalisation . Heritability may also occur at even larger scales. For example, ecological inheritance through the process of niche construction is defined by the regular and repeated activities of organisms in their environment. This generates a legacy of effects that modify and feed back into the selection regime of subsequent generations. Other examples of heritability in evolution that are not under

18564-486: The morphology of its skeleton being more adapted to living in trees rather than walking on two legs like later hominins. In May 2010, a new species, Homo gautengensis , was discovered in South Africa. These are proposed species names for fossils from about 1.9–1.6 Ma, whose relation to Homo habilis is not yet clear. The first fossils of Homo erectus were discovered by Dutch physician Eugene Dubois in 1891 on

18720-442: The most recent common ancestor of living members will still be part of a crown group. For example, if we consider the crown-birds (i.e. all extant birds and the rest of the family tree back to their most recent common ancestor), extinct side branches like the dodo or great auk are still descended from the most recent common ancestor of all living birds , so fall within the bird crown group. One very simplified cladogram for birds

18876-408: The movement of pollen between heavy-metal-tolerant and heavy-metal-sensitive populations of grasses. Gene transfer between species includes the formation of hybrid organisms and horizontal gene transfer . Horizontal gene transfer is the transfer of genetic material from one organism to another organism that is not its offspring; this is most common among bacteria. In medicine, this contributes to

19032-464: The narrower one. Often, an (extinct) grouping is identified as belonging together. Later, it may be realized other (extant) groupings actually emerged within such grouping, rendering them a stem grouping. Cladistically , the new groups should then be added to the group, as paraphyletic groupings are not natural. In any case, stem groupings with living descendants should not be viewed as a cohesive group, but their tree should be further resolved to reveal

19188-427: The next generation than those with traits that do not confer an advantage. This teleonomy is the quality whereby the process of natural selection creates and preserves traits that are seemingly fitted for the functional roles they perform. Consequences of selection include nonrandom mating and genetic hitchhiking . The central concept of natural selection is the evolutionary fitness of an organism. Fitness

19344-499: The offspring of sexual organisms contain random mixtures of their parents' chromosomes that are produced through independent assortment. In a related process called homologous recombination , sexual organisms exchange DNA between two matching chromosomes. Recombination and reassortment do not alter allele frequencies, but instead change which alleles are associated with each other, producing offspring with new combinations of alleles. Sex usually increases genetic variation and may increase

19500-635: The oldest known primate-like mammal species, the Plesiadapis , came from North America; another, Archicebus , came from China . Other similar basal primates were widespread in Eurasia and Africa during the tropical conditions of the Paleocene and Eocene . David R. Begun concluded that early primates flourished in Eurasia and that a lineage leading to the African apes and humans, including to Dryopithecus , migrated south from Europe or Western Asia into Africa. The surviving tropical population of primates—which

19656-474: The origin of the stem group concept to Austrian systematist Othenio Abel (1914), and it was discussed and diagrammed in English as early as 1933 by A. S. Romer . Alternatively, the term "stem group" is sometimes used in a wider sense to cover any members of the traditional taxon falling outside the crown group. Permian synapsids like Dimetrodon or Anteosaurus are stem mammals in the wider sense but not in

19812-456: The other alleles of that gene, then with each generation this allele has a higher probability of becoming common within the population. These traits are said to be "selected for ." Examples of traits that can increase fitness are enhanced survival and increased fecundity . Conversely, the lower fitness caused by having a less beneficial or deleterious allele results in this allele likely becoming rarer—they are "selected against ." Importantly,

19968-485: The other great apes at about 12 million years; there are no fossils that clearly document the ancestry of gibbons, which may have originated in a so-far-unknown Southeast Asian hominoid population, but fossil proto-orangutans may be represented by Sivapithecus from India and Griphopithecus from Turkey, dated to around 10 mya. Hominidae subfamily Homininae (African hominids) diverged from Ponginae (orangutans) about 14 mya. Hominins (including humans and

20124-578: The overall process, like a step in an assembly line. One example of mutation is wild boar piglets. They are camouflage coloured and show a characteristic pattern of dark and light longitudinal stripes. However, mutations in the melanocortin 1 receptor ( MC1R ) disrupt the pattern. The majority of pig breeds carry MC1R mutations disrupting wild-type colour and different mutations causing dominant black colouring. In asexual organisms, genes are inherited together, or linked , as they cannot mix with genes of other organisms during reproduction. In contrast,

20280-493: The phylogenetic split from the remaining amniotes (the Sauropsida ). Pan-Mammalia is thus an alternative name for Synapsida . A stem group is a paraphyletic assemblage composed of the members of a pan-group or total group, above, minus the crown group itself (and therefore minus all living members of the pan-group). This leaves primitive relatives of the crown groups , back along the phylogenetic line to (but not including)

20436-690: The possibility that Neanderthals, Denisovans, and modern humans may have co-existed and interbred. The existence of this distant branch creates a much more complex picture of humankind during the Late Pleistocene than previously thought. Evidence has also been found that as much as 6% of the DNA of some modern Melanesians derive from Denisovans, indicating limited interbreeding in Southeast Asia. Alleles thought to have originated in Neanderthals and Denisovans have been identified at several genetic loci in

20592-527: The presence of hip bones in whales and snakes, and sexual traits in organisms that reproduce via asexual reproduction. Examples of vestigial structures in humans include wisdom teeth , the coccyx , the vermiform appendix , and other behavioural vestiges such as goose bumps and primitive reflexes . However, many traits that appear to be simple adaptations are in fact exaptations : structures originally adapted for one function, but which coincidentally became somewhat useful for some other function in

20748-414: The process. One example is the African lizard Holaspis guentheri , which developed an extremely flat head for hiding in crevices, as can be seen by looking at its near relatives. However, in this species, the head has become so flattened that it assists in gliding from tree to tree—an exaptation. Within cells, molecular machines such as the bacterial flagella and protein sorting machinery evolved by

20904-494: The rate of evolution. The two-fold cost of sex was first described by John Maynard Smith . The first cost is that in sexually dimorphic species only one of the two sexes can bear young. This cost does not apply to hermaphroditic species, like most plants and many invertebrates . The second cost is that any individual who reproduces sexually can only pass on 50% of its genes to any individual offspring, with even less passed on as each new generation passes. Yet sexual reproduction

21060-472: The recent African origin theory, modern humans evolved in Africa possibly from H. heidelbergensis , H. rhodesiensis or H. antecessor and migrated out of the continent some 50,000 to 100,000 years ago, gradually replacing local populations of H. erectus , Denisova hominins , H. floresiensis , H. luzonensis and H. neanderthalensis , whose ancestors had left Africa in earlier migrations. Archaic Homo sapiens ,

21216-516: The recruitment of several pre-existing proteins that previously had different functions. Another example is the recruitment of enzymes from glycolysis and xenobiotic metabolism to serve as structural proteins called crystallins within the lenses of organisms' eyes. Crown group The concept was developed by Willi Hennig , the formulator of phylogenetic systematics , as a way of classifying living organisms relative to their extinct relatives in his "Die Stammesgeschichte der Insekten", and

21372-408: The relative importance of selection and neutral processes, including drift. The comparative importance of adaptive and non-adaptive forces in driving evolutionary change is an area of current research . Mutation bias is usually conceived as a difference in expected rates for two different kinds of mutation, e.g., transition-transversion bias, GC-AT bias, deletion-insertion bias. This is related to

21528-417: The same strand of DNA to become separated. However, the rate of recombination is low (approximately two events per chromosome per generation). As a result, genes close together on a chromosome may not always be shuffled away from each other and genes that are close together tend to be inherited together, a phenomenon known as linkage . This tendency is measured by finding how often two alleles occur together on

21684-575: The second interglacial period in the Middle Pleistocene , around 250,000 years ago, the trend in intra-cranial volume expansion and the elaboration of stone tool technologies developed, providing evidence for a transition from H. erectus to H. sapiens . The direct evidence suggests there was a migration of H. erectus out of Africa , then a further speciation of H. sapiens from H. erectus in Africa. A subsequent migration (both within and out of Africa) eventually replaced

21840-552: The sequence of nucleotides in the DNA. These phenomena are classed as epigenetic inheritance systems. DNA methylation marking chromatin , self-sustaining metabolic loops, gene silencing by RNA interference and the three-dimensional conformation of proteins (such as prions ) are areas where epigenetic inheritance systems have been discovered at the organismic level. Developmental biologists suggest that complex interactions in genetic networks and communication among cells can lead to heritable variations that may underlay some of

21996-549: The site of Denisova Cave in the Altai Mountains of Siberia uncovered a small bone fragment from the fifth finger of a juvenile member of another human species, the Denisovans. Artifacts, including a bracelet, excavated in the cave at the same level were carbon dated to around 40,000 BP. As DNA had survived in the fossil fragment due to the cool climate of the Denisova Cave, both mtDNA and nuclear DNA were sequenced. While

22152-403: The specimen was a good climber. It is thought given the night predators of the region that he built a nesting platform at night in the trees in a similar fashion to chimpanzees and gorillas. The earliest documented representative of the genus Homo is Homo habilis , which evolved around 2.8  million years ago , and is arguably the earliest species for which there is positive evidence of

22308-406: The spread of antibiotic resistance , as when one bacteria acquires resistance genes it can rapidly transfer them to other species. Horizontal transfer of genes from bacteria to eukaryotes such as the yeast Saccharomyces cerevisiae and the adzuki bean weevil Callosobruchus chinensis has occurred. An example of larger-scale transfers are the eukaryotic bdelloid rotifers , which have received

22464-640: The survival of individual males. This survival disadvantage is balanced by higher reproductive success in males that show these hard-to-fake , sexually selected traits. Evolution influences every aspect of the form and behaviour of organisms. Most prominent are the specific behavioural and physical adaptations that are the outcome of natural selection. These adaptations increase fitness by aiding activities such as finding food, avoiding predators or attracting mates. Organisms can also respond to selection by cooperating with each other, usually by aiding their relatives or engaging in mutually beneficial symbiosis . In

22620-642: The system...." Each population within an ecosystem occupies a distinct niche , or position, with distinct relationships to other parts of the system. These relationships involve the life history of the organism, its position in the food chain and its geographic range. This broad understanding of nature enables scientists to delineate specific forces which, together, comprise natural selection. Natural selection can act at different levels of organisation , such as genes, cells, individual organisms, groups of organisms and species. Selection can act at multiple levels simultaneously. An example of selection occurring below

22776-650: The transition to behavioral modernity with the development of symbolic culture, language, and specialized lithic technology happened around 50,000 years ago (beginning of the Upper Paleolithic ), although others point to evidence of a gradual change over a longer time span during the Middle Paleolithic. Homo sapiens is the only extant species of its genus, Homo . While some (extinct) Homo species might have been ancestors of Homo sapiens , many, perhaps most, were likely "cousins", having speciated away from

22932-402: The use of stone tools. The brains of these early hominins were about the same size as that of a chimpanzee, although it has been suggested that this was the time in which the human SRGAP2 gene doubled, producing a more rapid wiring of the frontal cortex. During the next million years a process of rapid encephalization occurred, and with the arrival of Homo erectus and Homo ergaster in

23088-597: The widely used total-group perspective, the Crocodylomorpha would become synonymous with the Crocodilia, and the Avemetatarsalia would become synonymous with the birds, and the above tree could be summarized as Crocodilia Birds An advantage of this approach is that declaring Theropoda to be birds (or Pan-aves ) is more specific than declaring it to be a member of the Archosauria, which would not exclude it from

23244-487: The years. One use is as "nearby group" (plesion means close to in Greek ), i.e. sister group to a given taxon , whether that group is a crown group or not. The term may also mean a group, possibly paraphyletic , defined by primitive traits (i.e. symplesiomorphies ). It is generally taken to mean a side branch splitting off earlier on the phylogenetic tree than the group in question. Placing fossils in their right order in

23400-524: Was Neanderthal, implying the individual had a Neanderthal ancestor 4–6 generations previously, in addition to a contribution from earlier interbreeding in the Middle East. Though this interbred Romanian population seems not to have been ancestral to modern humans, the finding indicates that interbreeding happened repeatedly. All modern non-African humans have about 1% to 4% (or 1.5% to 2.6% by more recent data) of their DNA derived from Neanderthals. This finding

23556-478: Was also argued that the species provides support for the notion that very early hominins, akin to bonobos ( Pan paniscus ) the less aggressive species of the genus Pan , may have evolved via the process of self-domestication . Consequently, arguing against the so-called "chimpanzee referential model" the authors suggest it is no longer tenable to use chimpanzee ( Pan troglodytes ) social and mating behaviors in models of early hominin social evolution. When commenting on

23712-547: Was controversial from its first proposal in the 1990s and by the 2010s had very little support. Distinctive human genetic variability has arisen as the result of the founder effect , by archaic admixture and by recent evolutionary pressures . Since Homo sapiens separated from its last common ancestor shared with chimpanzees , human evolution is characterized by a number of morphological , developmental , physiological , behavioral , and environmental changes. Environmental (cultural) evolution discovered much later during

23868-406: Was estimated to be one meter in height, with a brain volume of just 380 cm (considered small for a chimpanzee and less than a third of the H. sapiens average of 1400 cm). However, there is an ongoing debate over whether H. floresiensis is indeed a separate species. Some scientists hold that H. floresiensis was a modern H. sapiens with pathological dwarfism. This hypothesis

24024-655: Was first set out in detail in Darwin's book On the Origin of Species . Evolution by natural selection is established by observable facts about living organisms: (1) more offspring are often produced than can possibly survive; (2) traits vary among individuals with respect to their morphology , physiology , and behaviour; (3) different traits confer different rates of survival and reproduction (differential fitness ); and (4) traits can be passed from generation to generation ( heritability of fitness). In successive generations, members of

24180-465: Was not linear but weblike. The study of the origins of humans involves several scientific disciplines, including physical and evolutionary anthropology , paleontology , and genetics ; the field is also known by the terms anthropogeny , anthropogenesis , and anthropogony . (The latter two terms are sometimes used to refer to the related subject of hominization .) Primates diverged from other mammals about 85  million years ago ( mya ), in

24336-526: Was the key adaptation that separated Homo from tree-sleeping Australopithecines. These are proposed as species intermediate between H. erectus and H. heidelbergensis . H. heidelbergensis ("Heidelberg Man") lived from about 800,000 to about 300,000 years ago. Also proposed as Homo sapiens heidelbergensis or Homo sapiens paleohungaricus . Homo neanderthalensis , alternatively designated as Homo sapiens neanderthalensis , lived in Europe and Asia from 400,000 to about 28,000 years ago. There are

#10989