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Asmari Formation

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The Asmari Formation is a geological formation in western Iran . Located in the western Zagros Mountains , it is a carbonate formation that largely consists of limestone , dolomitic limestone , dolomite , and marly limestone. There are also smaller amounts of anhydrite , lithic sandstone , and limey sandstone. The formation dates to the Oligocene and Miocene periods and is a major hydrocarbon reserve. It has been producing oil since the 1930s. It also forms a major aquifer , which discharges at various springs in the Zagros region.

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91-537: The Asmari Formation's basic biostratigraphy was established in the 1950s and it was formally described in 1965. It consists of carbonate platforms that were formed in 6 distinct stages. The formation is named after the Asmari Mountains southeast of Masjed Soleyman , and its type sample was taken from the Tang-e Gel Torsh in these mountains. The Asmari Formation was deposited at a time when

182-407: A faunal assemblage , rather than an individual species — this allows greater precision as the time span in which all of the species in the assemblage existed together is narrower than the time spans of any of the members. Furthermore, if only one species is present in a sample, it can mean either that (1) the strata were formed in the known fossil range of that organism; or (2) that the fossil range of

273-490: 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

364-570: 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

455-514: 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

546-402: 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

637-405: 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

728-399: A particular span of geologic time or environment, and can be used to identify and date the containing rocks. To be practical, index fossils must have a limited vertical time range, wide geographic distribution, and rapid evolutionary trends. Rock formations separated by great distances but containing the same index fossil species are thereby known to have both formed during the limited time that

819-611: 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

910-538: 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

1001-575: 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

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1092-430: 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

1183-439: 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

1274-433: 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

1365-499: 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

1456-430: 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

1547-452: 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,

1638-419: 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

1729-420: 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

1820-544: 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

1911-411: 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

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2002-472: 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

2093-428: 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

2184-404: A unique association of three or more taxa within it. Abundance biozones are strata in which the abundance of a particular taxon or group of taxa is significantly greater than in the adjacent part of the section. Index fossils (also known as guide fossils , indicator fossils , or dating fossils ) are the fossilized remains or traces of particular plants or animals that are characteristic of

2275-454: 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

2366-442: 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,

2457-427: 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

2548-417: 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

2639-464: 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

2730-407: 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,

2821-692: 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

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2912-594: 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

3003-421: 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,

3094-429: 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

3185-447: 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

3276-430: Is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock strata by using the fossil assemblages contained within them. The primary objective of biostratigraphy is correlation , demonstrating that a particular horizon in one geological section represents the same period of time as another horizon at a different section. Fossils within these strata are useful because sediments of

3367-495: 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

3458-439: 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

3549-714: The Tethys Ocean was finally closing and the Zagros Mountains were first rising. During this period, the region was a shallow ocean gradually getting less deep. By the time of the succeeding Gachsaran Formation , this process had culminated in the sea shrinking to lagoons. In the southwestern Zagros, the Asmari Formation sits on top of the Pabdeh Formation . In the Fars and Luristan regions,

3640-425: 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

3731-454: 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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3822-539: The 18th century it began to be accepted that fossils were remains left by species that had become extinct, but were then preserved within the rock record. The method was well-established before Charles Darwin explained the mechanism behind it— evolution . Scientists William Smith , George Cuvier , and Alexandre Brongniart came to the conclusion that fossils then indicated a series of chronological events, establishing layers of rock strata as some type of unit, later termed biozone . From here on, scientists began relating

3913-706: The Asmari Formation sits on top of the Jahrum and Shahbazan Formations. The Asmari Formation is thickest in the northeastern part of the Dezful Embayment . The formation has various microfossils including planktonic foraminifera , echinoids , red algae , ostracoda , and oncoids . Individual species include Globigerina , Turborotalia cerroazulensis , Hantkenina , Nummulites , Lepidocyclina , Miogypsina , Elphidium , Peneroplis farsenensis , and Borelis melo curdica and Borelis melo melo . Biostratigraphy Biostratigraphy

4004-519: 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

4095-402: 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

4186-507: 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

4277-501: The basic biostratigraphy units, and define geological time periods based upon the fossil species found within each section. Basic concepts of biostratigraphic principles were introduced centuries ago, going as far back as the early 1800s. A Danish scientist and bishop by the name of Nicolas Steno was one of the first geologists to recognize that rock layers correlate to the Law of Superposition . With advancements in science and technology, by

4368-441: 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

4459-451: The boundaries of the different eras ( Paleozoic , Mesozoic , Cenozoic ), as well as Periods ( Cambrian , Ordovician , Silurian ) through the isotopes found within fossils via radioactive decay. Current 21st century uses of biostratigraphy involve interpretations of age for rock layers, which are primarily used by oil and gas industries for drilling workflows and resource allocations. Fossil assemblages were traditionally used to designate

4550-488: 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

4641-503: The changes in strata and biozones to different geological eras, establishing boundaries and time periods within major faunal changes. By the late 18th century the Cambrian and Carboniferous periods were internationally recognized due to these findings. During the early 20th century, advancements in technology gave scientists the ability to study radioactive decay . Using this methodology, scientists were able to establish geological time,

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4732-403: 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

4823-400: 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

4914-490: 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

5005-413: The duration of periods. Since a large change in fauna was required to make early stratigraphers create a new period, most of the periods we recognize today are terminated by a major extinction event or faunal turnover. A stage is a major subdivision of strata, each systematically following the other each bearing a unique assemblage of fossils. Therefore, stages can be defined as a group of strata containing

5096-489: 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

5187-414: 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

5278-427: 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

5369-472: 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

5460-479: 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

5551-640: 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

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5642-450: 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;

5733-481: The known stratigraphic and geographic range of occurrence of a single taxon. Concurrent range biozone includes the concurrent, coincident, or overlapping part of the range of two specified taxa. Interval biozones include the strata between two specific biostratigraphic surfaces and can be based on lowest or highest occurrences. Lineage biozones are strata containing species representing a specific segment of an evolutionary lineage. Assemblage biozones are strata that contain

5824-512: 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

5915-452: 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

6006-405: 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

6097-458: 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

6188-663: The most fundamental unit of measurement. The thickness and range of these zones can be a few meters, up to hundreds of meters. They can also range from local to worldwide, as the extent of which they can reach in the horizontal plane relies on tectonic plates and tectonic activity. Two of the tectonic processes that run the risk of changing these zones' ranges are metamorphic folding and subduction . Furthermore, biostratigraphic units are divided into six principal kinds of biozones: Taxon range biozone , Concurrent range biozone, Interval biozone, Lineage biozone, Assemblage biozone, and Abundance biozone . The Taxon range biozone represents

6279-408: 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

6370-427: 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

6461-499: 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

6552-466: The organism was incompletely known, and the strata extend the known fossil range. For instance, the presence of the trace fossil Treptichnus pedum was used to define the base of the Cambrian period, but it has since been found in older strata. If the fossil is easy to preserve and easy to identify, more precise time estimating of the stratigraphic layers is possible. The concept of faunal succession

6643-456: 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,

6734-578: 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,

6825-430: The overlapping range of fossils. They represent the time between the appearance of species chosen at the base of the zone and the appearance of other species chosen at the base of the next succeeding zone. Oppel's zones are named after a particular distinctive fossil species, called an index fossil. Index fossils are one of the species from the assemblage of species that characterize the zone. Biostratigraphy uses zones for

6916-527: 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

7007-517: The principle of faunal succession, where fossil organisms succeed one another in a definite and determinable order, and therefore any time period can be categorized by its fossil extent. 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

7098-469: 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

7189-494: 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

7280-408: 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

7371-423: The same age can look completely different, due to local variations in the sedimentary environment . For example, one section might have been made up of clays and marls , while another has more chalky limestones . However, if the fossil species recorded are similar, the two sediments are likely to have been laid down around the same time. Ideally these fossils are used to help identify biozones , as they make up

7462-443: The same major fossil assemblages. French palaeontologist Alcide d'Orbigny is credited for the invention of this concept. He named stages after geographic localities with particularly good sections of rock strata that bear the characteristic fossils on which the stages are based. In 1856 German palaeontologist Albert Oppel introduced the concept of zone (also known as biozones or Oppel zone). A zone includes strata characterized by

7553-417: 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

7644-552: 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

7735-832: The species lived. Index fossils were originally used to define and identify geologic units, then became a basis for defining geologic periods , and then for faunal stages and zones. Ammonites , graptolites , archeocyathids , inoceramids , and trilobites are groups of animals from which many species have been identified as index fossils that are widely used in biostratigraphy. Species of microfossils such as acritarchs , chitinozoans , conodonts , dinoflagellate cysts, ostracods , pollen , spores and foraminiferans are also frequently used. Different fossils work well for sediments of different ages; trilobites, for example, are particularly useful for sediments of Cambrian age. A long series of ammonite and inoceramid species are particularly useful for correlating environmental events around

7826-406: 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

7917-640: 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

8008-642: 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

8099-642: The world during the super-greenhouse of the Late Cretaceous . To work well, the fossils used must be widespread geographically, so that they can be found in many different places. They must also be short-lived as a species, so that the period of time during which they could be incorporated in the sediment is relatively narrow. The longer lived the species, the poorer the stratigraphic precision, so fossils that evolve rapidly, such as ammonites, are favored over forms that evolve much more slowly, like nautiloids . Often biostratigraphic correlations are based on

8190-663: 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

8281-619: Was theorized at the beginning of the 19th century by William Smith . When Smith was studying rock strata, he began to recognize that rock outcrops contained a unique collection of fossils. The idea that these distant rock outcrops contained similar fossils allowed for Smith to order rock formations throughout England. With Smith's work on these rock outcrops and mapping around England, he began to notice some beds of rock may contain mostly similar species, however there were also subtle differences within or between these fossil groups. This difference in assemblages that appeared identical at first, lead to

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