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126-466: Fossils of early birds were poorly known until the late 20th century. Of those known, all fell into either the relatively modernly built birds with fused ribcage and the breastbone extended into a keel , or the " Urvogels " of the Solnhofen Plattenkalk of late Jurassic age. As the physiological and anatomical difference between the two was so great, the subclass Archaeornithes was erected for

252-487: 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

378-570: 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

504-514: 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

630-467: A constant rate. These " molecular clocks ", however, are fallible, and provide only approximate timing: for example, they are not sufficiently precise and reliable for estimating when the groups that feature in the Cambrian explosion first evolved, and estimates produced by different techniques may vary by a factor of two. Organisms are only rarely preserved as fossils in the best of circumstances, and only

756-426: A fraction of such fossils have been discovered. This is illustrated by the fact that the number of species known through the fossil record is less than 5% of the number of known living species, suggesting that the number of species known through fossils must be far less than 1% of all the species that have ever lived. Because of the specialized and rare circumstances required for a biological structure to fossilize, only

882-495: A hierarchical classification system still in use today. Darwin and his contemporaries first linked the hierarchical structure of the tree of life with the then very sparse fossil record. Darwin eloquently described a process of descent with modification, or evolution, whereby organisms either adapt to natural and changing environmental pressures, or they perish. When Darwin wrote On the Origin of Species by Means of Natural Selection, or

1008-402: 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

1134-418: A mid-Ordovician age. Such index fossils must be distinctive, be globally distributed and occupy a short time range to be useful. Misleading results are produced if the index fossils are incorrectly dated. Stratigraphy and biostratigraphy can in general provide only relative dating ( A was before B ), which is often sufficient for studying evolution. However, this is difficult for some time periods, because of

1260-526: A mutation first appeared. Phylogenetics and paleontology work together in the clarification of science's still dim view of the appearance of life and its evolution. Niles Eldredge 's study of the Phacops trilobite genus supported the hypothesis that modifications to the arrangement of the trilobite's eye lenses proceeded by fits and starts over millions of years during the Devonian . Eldredge's interpretation of

1386-405: 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

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1512-611: 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

1638-535: 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

1764-575: 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

1890-430: 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

2016-433: 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

2142-570: A portion of the deceased organism, usually that portion that was partially mineralized during life, such as the bones and teeth of vertebrates , or the chitinous or calcareous exoskeletons of invertebrates . Fossils may also consist of the marks left behind by the organism while it was alive, such as animal tracks or feces ( coprolites ). These types of fossil are called trace fossils or ichnofossils , as opposed to body fossils . Some fossils are biochemical and are called chemofossils or biosignatures . Gathering fossils dates at least to

2268-499: 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

2394-430: 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

2520-400: A richly diverse assembly of early multicellular eukaryotes . The fossil record and faunal succession form the basis of the science of biostratigraphy or determining the age of rocks based on embedded fossils. For the first 150 years of geology , biostratigraphy and superposition were the only means for determining the relative age of rocks. The geologic time scale was developed based on

2646-534: A series of nested monophyletic groups, and the name is rarely used in modern literature. Archaeopteryx Archaeornis other infraclasses Odontognathae order order In traditional classification, it is one of two subclasses of birds, the other subclass being the Neornithes , the birds with a short, modern tail. This classification was erected by Hans Friedrich Gadow in 1893 and followed by Alfred Romer (1933) and subsequent authors through most of

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2772-452: 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,

2898-419: 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

3024-503: A single species, Archaeopteryx lithographica , the Archaeornis being the Berlin specimen of Archaeopteryx . The Confuciusornithidae and Enantiornithes were found a century after Gradow's organization of birds into two subclasses. They fall between Romer's description of Archaeornithes and Neornithes, in that they have clawed wings, but reduced tails with a rod-like pygostyle (as opposed to

3150-448: A small percentage of life-forms can be expected to be represented in discoveries, and each discovery represents only a snapshot of the process of evolution. The transition itself can only be illustrated and corroborated by transitional fossils, which will never demonstrate an exact half-way point. The fossil record is strongly biased toward organisms with hard-parts, leaving most groups of soft-bodied organisms with little to no role. It

3276-420: 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

3402-544: 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

3528-411: 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

3654-472: 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

3780-428: 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

3906-454: 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

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4032-442: 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,

4158-427: 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

4284-449: Is a notable example of how knowledge encoded by the fossil record continues to contribute otherwise unattainable information on the emergence and development of life on Earth. For example, the research suggests Markuelia has closest affinity to priapulid worms, and is adjacent to the evolutionary branching of Priapulida , Nematoda and Arthropoda . Despite significant advances in uncovering and identifying paleontological specimens, it

4410-463: Is a process of fossilization that occurs when an organism is buried. The empty spaces within an organism (spaces filled with liquid or gas during life) become filled with mineral-rich groundwater . Minerals precipitate from the groundwater, occupying the empty spaces. This process can occur in very small spaces, such as within the cell wall of a plant cell . Small scale permineralization can produce very detailed fossils. For permineralization to occur,

4536-417: 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

4662-412: Is any preserved remains, impression, or trace of any once-living thing from a past geological age . Examples include bones , shells , exoskeletons , stone imprints of animals or microbes , objects preserved in amber , hair , petrified wood and DNA remnants. The totality of fossils is known as the fossil record . Though the fossil record is incomplete, numerous studies have demonstrated that there

4788-464: 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

4914-407: 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,

5040-692: 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

5166-538: Is enough information available to give a good understanding of the pattern of diversification of life on Earth. In addition, the record can predict and fill gaps such as the discovery of Tiktaalik in the arctic of Canada . Paleontology includes the study of fossils: their age, method of formation, and evolutionary significance. Specimens are usually considered to be fossils if they are over 10,000 years old. The oldest fossils are around 3.48 billion years to 4.1 billion years old. The observation in

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5292-479: Is generally accepted that the fossil record is vastly incomplete. Approaches for measuring the completeness of the fossil record have been developed for numerous subsets of species, including those grouped taxonomically, temporally, environmentally/geographically, or in sum. This encompasses the subfield of taphonomy and the study of biases in the paleontological record. Paleontology seeks to map out how life evolved across geologic time. A substantial hurdle

5418-594: 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

5544-438: Is replete with the mollusks , the vertebrates , the echinoderms , the brachiopods and some groups of arthropods . Fossil sites with exceptional preservation—sometimes including preserved soft tissues—are known as Lagerstätten —German for "storage places". These formations may have resulted from carcass burial in an anoxic environment with minimal bacteria, thus slowing decomposition. Lagerstätten span geological time from

5670-421: 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,

5796-429: 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

5922-447: 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

6048-500: 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 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

6174-470: Is the difficulty of working out fossil ages. Beds that preserve fossils typically lack the radioactive elements needed for radiometric dating . This technique is our only means of giving rocks greater than about 50 million years old an absolute age, and can be accurate to within 0.5% or better. Although radiometric dating requires careful laboratory work, its basic principle is simple: the rates at which various radioactive elements decay are known, and so

6300-495: 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

6426-439: 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

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6552-413: Is the science of deciphering the "layer-cake" that is the sedimentary record. Rocks normally form relatively horizontal layers, with each layer younger than the one underneath it. If a fossil is found between two layers whose ages are known, the fossil's age is claimed to lie between the two known ages. Because rock sequences are not continuous, but may be broken up by faults or periods of erosion , it

6678-459: Is very difficult to match up rock beds that are not directly adjacent. However, fossils of species that survived for a relatively short time can be used to match isolated rocks: this technique is called biostratigraphy . For instance, the conodont Eoplacognathus pseudoplanus has a short range in the Middle Ordovician period. If rocks of unknown age have traces of E. pseudoplanus , they have

6804-757: The Cambrian period to the present . Worldwide, some of the best examples of near-perfect fossilization are the Cambrian Maotianshan Shales and Burgess Shale , the Devonian Hunsrück Slates , the Jurassic Solnhofen Limestone , and the Carboniferous Mazon Creek localities. A fossil is said to be recrystallized when the original skeletal compounds are still present but in a different crystal form, such as from aragonite to calcite . Replacement occurs when

6930-581: The Phacops fossil record was that the aftermaths of the lens changes, but not the rapidly occurring evolutionary process, were fossilized. This and other data led Stephen Jay Gould and Niles Eldredge to publish their seminal paper on punctuated equilibrium in 1971. Synchrotron X-ray tomographic analysis of early Cambrian bilaterian embryonic microfossils yielded new insights of metazoan evolution at its earliest stages. The tomography technique provides previously unattainable three-dimensional resolution at

7056-559: The Renaissance . Leonardo da Vinci concurred with Aristotle's view that fossils were the remains of ancient life. For example, Leonardo noticed discrepancies with the biblical flood narrative as an explanation for fossil origins: If the Deluge had carried the shells for distances of three and four hundred miles from the sea it would have carried them mixed with various other natural objects all heaped up together; but even at such distances from

7182-591: The Song dynasty during the 11th century, who kept a specific seashell fossil with his own poem engraved on it. In his Dream Pool Essays published in 1088, Song dynasty Chinese scholar-official Shen Kuo hypothesized that marine fossils found in a geological stratum of mountains located hundreds of miles from the Pacific Ocean was evidence that a prehistoric seashore had once existed there and shifted over centuries of time . His observation of petrified bamboos in

7308-402: The law of superposition ) preserved different assemblages of fossils, and that these assemblages succeeded one another in a regular and determinable order. He observed that rocks from distant locations could be correlated based on the fossils they contained. He termed this the principle of faunal succession . This principle became one of Darwin's chief pieces of evidence that biological evolution

7434-425: 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

7560-493: The ploughshare -shaped one in modern birds) and the presence of a small carina. While rarely used by palaenthologists today, the term was revived by the ornithologists Livezey and Zusi in 2007, for a group comprising Archaeopterygidae and the Confuciusornithidae . [REDACTED] [REDACTED] Fossils A fossil (from Classical Latin fossilis , lit.   ' obtained by digging ' )

7686-454: 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

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7812-403: The thunderbird . There is no such direct mythological connection known from prehistoric Africa, but there is considerable evidence of tribes there excavating and moving fossils to ceremonial sites, apparently treating them with some reverence. In Japan, fossil shark teeth were associated with the mythical tengu , thought to be the razor-sharp claws of the creature, documented some time after

7938-449: The 16th century. Roman naturalist Pliny the Elder wrote of " tongue stones ", which he called glossopetra . These were fossil shark teeth, thought by some classical cultures to look like the tongues of people or snakes. He also wrote about the horns of Ammon , which are fossil ammonites , whence the group of shelled octopus-cousins ultimately draws its modern name. Pliny also makes one of

8064-741: The 19th century that certain fossils were associated with certain rock strata led to the recognition of a geological timescale and the relative ages of different fossils. The development of radiometric dating techniques in the early 20th century allowed scientists to quantitatively measure the absolute ages of rocks and the fossils they host. There are many processes that lead to fossilization , including permineralization , casts and molds, authigenic mineralization , replacement and recrystallization, adpression, carbonization , and bioimmuration. Fossils vary in size from one- micrometre (1 μm) bacteria to dinosaurs and trees, many meters long and weighing many tons. A fossil normally preserves only

8190-609: The 20th century. Other mesozoic birds like the toothed, but otherwise modern, birds like Hesperornis were included under the latter in their own superorder, the Odontognathae . According to Romer , the Archaeornithes are characterised by having clawed wings, a reptilian style ribcage without a large carina and the presence of a long, bony tail. The known members of the group by the time of its erection were Archaeopteryx and Archaeornis . The two are now thought to represent

8316-413: The 8th century AD. In medieval China, the fossil bones of ancient mammals including Homo erectus were often mistaken for " dragon bones" and used as medicine and aphrodisiacs . In addition, some of these fossil bones are collected as "art" by scholars, who left scripts on various artifacts, indicating the time they were added to a collection. One good example is the famous scholar Huang Tingjian of

8442-597: The Preservation of Favoured Races in the Struggle for Life , the oldest animal fossils were those from the Cambrian Period, now known to be about 540 million years old. He worried about the absence of older fossils because of the implications on the validity of his theories, but he expressed hope that such fossils would be found, noting that: "only a small portion of the world is known with accuracy." Darwin also pondered

8568-555: The Proterozoic and deeper still in the Archean is only "recounted by microscopic fossils and subtle chemical signals." Molecular biologists, using phylogenetics , can compare protein amino acid or nucleotide sequence homology (i.e., similarity) to evaluate taxonomy and evolutionary distances among organisms, with limited statistical confidence. The study of fossils, on the other hand, can more specifically pinpoint when and in what organism

8694-519: 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

8820-402: 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

8946-507: 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

9072-413: The beach, indicating the fossils were once living animals. He had previously explained them in terms of vaporous exhalations , which Persian polymath Avicenna modified into the theory of petrifying fluids ( succus lapidificatus ). Recognition of fossil seashells as originating in the sea was built upon in the 14th century by Albert of Saxony , and accepted in some form by most naturalists by

9198-710: The beginning of recorded history. The fossils themselves are referred to as the fossil record. The fossil record was one of the early sources of data underlying the study of evolution and continues to be relevant to the history of life on Earth . Paleontologists examine the fossil record to understand the process of evolution and the way particular species have evolved. Fossils have been visible and common throughout most of natural history, and so documented human interaction with them goes back as far as recorded history, or earlier. There are many examples of paleolithic stone knives in Europe, with fossil echinoderms set precisely at

9324-441: 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

9450-488: 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

9576-660: The deity Sopdu , the Morning Star, equivalent of Venus in Roman mythology. Fossils appear to have directly contributed to the mythology of many civilizations, including the ancient Greeks. Classical Greek historian Herodotos wrote of an area near Hyperborea where gryphons protected golden treasure. There was indeed gold mining in that approximate region , where beaked Protoceratops skulls were common as fossils. A later Greek scholar, Aristotle , eventually realized that fossil seashells from rocks were similar to those found on

9702-403: 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

9828-400: 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

9954-490: 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

10080-466: The dry northern climate zone of what is now Yan'an , Shaanxi province, China, led him to advance early ideas of gradual climate change due to bamboo naturally growing in wetter climate areas. In medieval Christendom , fossilized sea creatures on mountainsides were seen as proof of the biblical deluge of Noah's Ark . After observing the existence of seashells in mountains, the ancient Greek philosopher Xenophanes (c. 570 – 478 BC) speculated that

10206-415: The earlier known references to toadstones , thought until the 18th century to be a magical cure for poison originating in the heads of toads, but which are fossil teeth from Lepidotes , a Cretaceous ray-finned fish. The Plains tribes of North America are thought to have similarly associated fossils, such as the many intact pterosaur fossils naturally exposed in the region, with their own mythology of

10332-517: The earliest known stromatolites are over 3.4 billion years old. The fossil record is life's evolutionary epic that unfolded over four billion years as environmental conditions and genetic potential interacted in accordance with natural selection. Paleontology has joined with evolutionary biology to share the interdisciplinary task of outlining the tree of life, which inevitably leads backwards in time to Precambrian microscopic life when cell structure and functions evolved. Earth's deep time in

10458-460: The earth during earthquake and subsidences, and petrifies whatever comes into contact with it. As a matter of fact, the petrifaction of the bodies of plants and animals is not more extraordinary than the transformation of waters. From the 13th century to the present day, scholars pointed out that the fossil skulls of Deinotherium giganteum , found in Crete and Greece, might have been interpreted as being

10584-489: 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

10710-478: The existence of a world previous to ours, destroyed by some kind of catastrophe. Interest in fossils, and geology more generally, expanded during the early nineteenth century. In Britain, Mary Anning 's discoveries of fossils, including the first complete ichthyosaur and a complete plesiosaurus skeleton, sparked both public and scholarly interest. Early naturalists well understood the similarities and differences of living species leading Linnaeus to develop

10836-414: 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

10962-427: 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

11088-472: 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

11214-479: 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

11340-519: The hand grip, dating back to Homo heidelbergensis and Neanderthals . These ancient peoples also drilled holes through the center of those round fossil shells, apparently using them as beads for necklaces. The ancient Egyptians gathered fossils of species that resembled the bones of modern species they worshipped. The god Set was associated with the hippopotamus , therefore fossilized bones of hippo-like species were kept in that deity's temples. Five-rayed fossil sea urchin shells were associated with

11466-640: 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

11592-450: 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;

11718-400: The latter. With the unearthing of several well preserved early bird fossils in the last decades of the 20th century and early 21st century, our knowledge of the evolution of birds has increased dramatically. The evolution of the modern avian traits such as the compact body, clawless wing and the alula are now known to appear over successive stages. Today the Archaeornithes are classified into

11844-512: 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

11970-535: The limits of fossilization. Fossils of two enigmatic bilaterians, the worm-like Markuelia and a putative, primitive protostome , Pseudooides , provide a peek at germ layer embryonic development. These 543-million-year-old embryos support the emergence of some aspects of arthropod development earlier than previously thought in the late Proterozoic. The preserved embryos from China and Siberia underwent rapid diagenetic phosphatization resulting in exquisite preservation, including cell structures. This research

12096-452: 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

12222-405: 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

12348-458: 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

12474-408: 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

12600-427: 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

12726-499: 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

12852-429: The organism must become covered by sediment soon after death, otherwise the remains are destroyed by scavengers or decomposition. The degree to which the remains are decayed when covered determines the later details of the fossil. Some fossils consist only of skeletal remains or teeth; other fossils contain traces of skin , feathers or even soft tissues. This is a form of diagenesis . Evolution Evolution

12978-456: 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,

13104-578: 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,

13230-527: 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

13356-545: The problems involved in matching rocks of the same age across continents . Family-tree relationships also help to narrow down the date when lineages first appeared. For instance, if fossils of B or C date to X million years ago and the calculated "family tree" says A was an ancestor of B and C, then A must have evolved earlier. It is also possible to estimate how long ago two living clades diverged, in other words approximately how long ago their last common ancestor must have lived, by assuming that DNA mutations accumulate at

13482-464: 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

13608-494: 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

13734-443: The ratio of the radioactive element to its decay products shows how long ago the radioactive element was incorporated into the rock. Radioactive elements are common only in rocks with a volcanic origin, and so the only fossil-bearing rocks that can be dated radiometrically are volcanic ash layers, which may provide termini for the intervening sediments. Consequently, palaeontologists rely on stratigraphy to date fossils. Stratigraphy

13860-462: The relative ages of rock strata as determined by the early paleontologists and stratigraphers . Since the early years of the twentieth century, absolute dating methods, such as radiometric dating (including potassium/argon , argon/argon , uranium series , and, for very recent fossils, radiocarbon dating ) have been used to verify the relative ages obtained by fossils and to provide absolute ages for many fossils. Radiometric dating has shown that

13986-408: 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

14112-417: 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

14238-429: The sea and that they were still living when the strait of Gibraltar was cut through. In the mountains of Parma and Piacenza multitudes of shells and corals with holes may be seen still sticking to the rocks.... In 1666, Nicholas Steno examined a shark, and made the association of its teeth with the "tongue stones" of ancient Greco-Roman mythology, concluding that those were not in fact the tongues of venomous snakes, but

14364-423: The sea we see the oysters all together and also the shellfish and the cuttlefish and all the other shells which congregate together, found all together dead; and the solitary shells are found apart from one another as we see them every day on the sea-shores. And we find oysters together in very large families, among which some may be seen with their shells still joined together, indicating that they were left there by

14490-552: 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

14616-456: The shell, bone, or other tissue is replaced with another mineral. In some cases mineral replacement of the original shell occurs so gradually and at such fine scales that microstructural features are preserved despite the total loss of original material. Scientists can use such fossils when researching the anatomical structure of ancient species. Several species of saurids have been identified from mineralized dinosaur fossils. Permineralization

14742-581: The skulls of the Cyclopes of Greek mythology , and are possibly the origin of that Greek myth. Their skulls appear to have a single eye-hole in the front, just like their modern elephant cousins, though in fact it's actually the opening for their trunk. In Norse mythology , echinoderm shells (the round five-part button left over from a sea urchin) were associated with the god Thor , not only being incorporated in thunderstones , representations of Thor's hammer and subsequent hammer-shaped crosses as Christianity

14868-406: 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

14994-467: The sudden appearance of many groups (i.e. phyla ) in the oldest known Cambrian fossiliferous strata. Since Darwin's time, the fossil record has been extended to between 2.3 and 3.5 billion years. Most of these Precambrian fossils are microscopic bacteria or microfossils . However, macroscopic fossils are now known from the late Proterozoic. The Ediacara biota (also called Vendian biota) dating from 575 million years ago collectively constitutes

15120-640: 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

15246-642: 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

15372-451: The teeth of some long-extinct species of shark. Robert Hooke (1635–1703) included micrographs of fossils in his Micrographia and was among the first to observe fossil forams . His observations on fossils, which he stated to be the petrified remains of creatures some of which no longer existed, were published posthumously in 1705. William Smith (1769–1839) , an English canal engineer, observed that rocks of different ages (based on

15498-511: The world was once inundated in a great flood that buried living creatures in drying mud. In 1027, the Persian Avicenna explained fossils' stoniness in The Book of Healing : If what is said concerning the petrifaction of animals and plants is true, the cause of this (phenomenon) is a powerful mineralizing and petrifying virtue which arises in certain stony spots, or emanates suddenly from

15624-458: Was adopted, but also kept in houses to garner Thor's protection. These grew into the shepherd's crowns of English folklore, used for decoration and as good luck charms, placed by the doorway of homes and churches. In Suffolk , a different species was used as a good-luck charm by bakers, who referred to them as fairy loaves , associating them with the similarly shaped loaves of bread they baked. More scientific views of fossils emerged during

15750-838: 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 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

15876-408: Was real. Georges Cuvier came to believe that most if not all the animal fossils he examined were remains of extinct species. This led Cuvier to become an active proponent of the geological school of thought called catastrophism . Near the end of his 1796 paper on living and fossil elephants he said: All of these facts, consistent among themselves, and not opposed by any report, seem to me to prove

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