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Macroevolution
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== Origin and changing meaning of the term == After [[Charles Darwin]] published his book ''On the Origin of Species''<ref>{{Cite book|last=Darwin|first=C.|title=On the origin of species by means of natural selection|publisher=John Murray|year=1859|location=London}}</ref> in 1859, evolution was widely accepted to be real phenomenon. However, many scientists still disagreed with Darwin that [[natural selection]] was the primary mechanism to explain evolution. Prior to the [[Modern synthesis (20th century)|modern synthesis]], during the period between the 1880s to the 1930s (dubbed the ‘[[Eclipse of Darwinism]]’) many scientists argued in favor of alternative explanations. These included ‘[[orthogenesis]]’, and among its proponents was the Russian entomologist [[Yuri Filipchenko|Yuri A. Filipchenko]]. Filipchenko appears to have been the one who coined the term ‘macroevolution’ in his book ''Variabilität und Variation'' (1927).<ref name=":2" /> While introducing the concept, he claimed that the field of genetics is insufficient to explain ''“the origin of higher systematic units”'' above the species level. {{Text and translation | Auf die Weise hebt die heutige Genetik zweifellos den Schleier von der Evolution der Biotypen, Jordanone und Linneone (eine Art Mikroevolution), dagegen jene Evolution der höheren systematischen Gruppen, welche von jeher die Geister besonders für sich in Anspruch genommen hat (eine Art Makroevolution), liegt gänzlich außerhalb ihres Gesichtsfeldes, und dieser Umstand scheint uns die von uns oben angeführten Erwägungen über das Fehlen einer inneren Beziehung zwischen der Genetik und der Deszendenzlehre, die sich ja hauptsächlich mit der Makroevolution befaßt, nur zu unterstreichen. Bei einer solchen Sachlage muß zugegeben werden, daß die Entscheidung der Frage über die Faktoren der größeren Züge der Evolution, d. h. dessen, was wir Makroevolution nennen, unabhängig von den Ergebnissen der gegenwärtigen Genetik geschehen muß. So vorteilhaft es für uns auch wäre, uns auch in dieser Frage auf die exakten Resultate der Genetik zu stützen, so sind sie doch, unserer Meinung nach, zu diesem Zweck ganz unbrauchbar, da die Frage über die Entstehung der höheren systematischen Einheiten ganz außerhalb des Forschungsgebietes der Genetik liegt. Infolgedessen ist letztere auch eine exakte Wissenschaft, während die Deszendenzlehre heute, ebenso wie auch im XIX. Jahrhundert, einen einen spekulativen Charakter trägt. | In this way, modern genetics undoubtedly lifts the veil from the evolution of biotypes, Jordanones and Linneones [i.e. variations within a species]{{efn|name="BiotJordLinn" | The terms ('biotypes', 'Jordanone', and 'Linneone') used here by Filipchenko were/are rarely used among non-Russian speaking scientists. According to Krasil'nikov (1958),<ref name="Krasil1958a">{{cite book |last1=Krasilʹnikov |first1=Nikolaĭ Aleksandrovich |title=Soil microorganisms and higher plants |date=1958 |publisher=Academy of Sciences of the USSR |location=Moscow |url=https://www.soilandhealth.org/wp-content/uploads/01aglibrary/010112.krasilnikov.pdf}}</ref> these terms were used to describe the variety of forms observed within a single species: ''"With the development of genetics the concept of species widened according to the ideas of variability and heredity of organisms. New terms were introduced for the determination of species subdivision, such as "biotype", "pure line", "jardanon", "linneon", etc. ["Jardanon"--a simple means of classification of lower organisms. "Linneon"--the complex of "jardanons"--according to the Russian concept, the inner species variety of forms does not exceed the limits of qualitative unity of the species.]"''}} (a kind of microevolution), but that evolution of the higher systematic groups, which has always particularly occupied the minds of men (a kind of macroevolution), lies entirely outside its field of vision, and this circumstance seems to us only to emphasize the considerations we have given above about the lack of an inner relationship between genetics and the theory of descent, which is mainly concerned with macroevolution. In such a state of affairs, it must be admitted that the decision of the question depends on the factors of the larger features of evolution, of what we call macroevolution, must occur independently of the results of current genetics. As advantageous as it would be for us to rely on the exact results of genetics in this question, they are, in our opinion, completely useless for this purpose, since the question about the origin of the higher systematic units lies entirely outside the field research area of genetics. As a result, the latter is also an exact science, while the doctrine of descent today, as well as in the 19th century, has a speculative character. | Yuri Filipchenko, ''Variabilität und Variation'' (1927), pages 93-94<ref name=":2" /> }} Regarding the origin of higher systematic units, Filipchenko stated his claim that ‘like-produces-like’. A taxon must originate from other taxa of equivalent rank. A new species must come from an old species, a genus from an older genus, a family from another family, etc. {{Text and translation | Hier scheint uns ein wesentliches Mißverständnis obzuwalten. Davon schon gar nicht zu reden, daß es kaum richtig ist, in den Jardanonen Spaltungsprodukte eines Linneone zu sehen, ist es noch unrichtiger anzunehmen, daß nach den heutigen Anschauungen ein Jordanon sich im Evolutionsprozeß in ein neues Linneon verwandeln kann oder muß. Im Gegenteil, uns scheint, daß sich bei der Evolution die verschiedenen taxonomischen Einheiten so verhalten, daß Gleiches Gleiches erzeugt. Aus einem Biotyp entsteht durch Mutation ein neuer Biotypus, aus einem Jordanon bildet sich - durch eine Neugruppierung der ihn bildenden Biotypen, sowie durch das Auftreten einiger neuer - ein zweites Jordanon; endlich zerfällt ein aus mehreren Jordanonen bestehendes Linneon infolge des Verschwindens einiger von ihnen in zwei selbständige Linneone. Es ist vollkommen richtig, daß niemand eine Umwandlung der Rassen in eine Art beobachtet hat, aber das braucht auch nicht zu sein, da im Prozeß der Evolution eine neue Art oder Arten gewöhnlich aus einer alten Art, eine neue Gattung aus einer anderen Gattung usw. entstehen. | There seems to be a fundamental misunderstanding here. Not to mention that it is hardly correct to see the Jardanones{{efn|name="BiotJordLinn"}} as products of the fission of a Linneone,{{efn|name="BiotJordLinn"}} it is even more incorrect to assume that, according to modern views, a Jordanone can or must transform into a new Linneone in the process of evolution. On the contrary, it seems to us that in evolution the various taxonomic units behave in such a way that like produces like. A new biotype{{efn|name="BiotJordLinn"}} arises from one biotype through mutation; a Jordanone forms a second Jordanone through a regrouping of the biotypes that make up it and the appearance of some new ones; finally, a Linneone consisting of several Jordanones splits into two independent Linneones as a result of the disappearance of some of them. It is quite true that no one has observed a transformation of the races into a species, but that need not be the case, since in the process of evolution a new species or species usually arise from an old species, a new genus from another genus, etc. | Yuri Filipchenko, ''Variabilität und Variation'' (1927), page 89 <ref name=":2" /> }} Filipchenko believed this was the only way to explain the origin of the major characters that define species and especially higher taxonomic groups ([[genera]], [[Family_(biology)|families]], [[Order_(biology)|orders]], etc). For example, the origin of families must require the sudden appearance of new traits which are different in greater magnitude compared to the characters required for the origin of a genus or species. However, this view is no longer consistent with contemporary understanding of evolution. Furthermore, the [[Taxonomic_rank#Significance_and_problems|Linnaean ranks]] of ‘genus’ (and higher) are not real entities but artificial concepts which [[Taxonomic_boundary_paradox|break down]] when they are combined with the process of evolution.<ref name="Hendricks2014a">{{cite journal |last1=Hendricks |first1=Jonathan R. |last2=Saupe |first2=Erin E |last3=Myers |first3=Corinne E. |last4=Hermsen |first4=Elizabeth J. |last5=Allmon |first5=Warren D. |title=he generification of the fossil record. |journal=Paleobiology |date=2014 |volume=40 |issue=4 |pages=511–528 |doi=10.1666/13076}}</ref><ref name="DAOAL1"></ref> Nevertheless, Filipchenko’s distinction between microevolution and macroevolution had a major impact on the development of evolutionary science. The term was adopted by Filipchenko's protégé [[Theodosius Dobzhansky]] in his book ''‘Genetics und the Origin of Species’'' (1937), a seminal piece that contributed to the development of the [[Modern Synthesis]]. ‘Macroevolution’ was also adopted by those who used it to criticize the Modern Synthesis. A notable example of this was the book ''The Material Basis of Evolution'' (1940) by the geneticist [[Richard Goldschmidt]], a close friend of Filipchenko.<ref name="Adams1990a"></ref> Goldschmidt suggested [[Saltational evolution|saltational evolutionary changes]] either due to mutations that affect the rates of developmental processes<ref>{{Cite journal|last=Goldschmidt|first=R.|title=Some aspects of evolution|journal=Science|year=1933|volume=78|issue=2033|pages=539–547|doi=10.1126/science.78.2033.539|pmid=17811930|bibcode=1933Sci....78..539G}}</ref> or due to alterations in the chromosomal pattern.<ref>{{Cite book|last=Goldschmidt|first=R.|title=The material basis of evolution|publisher=Yale University Press|year=1940}}</ref> Particularly the latter idea was widely rejected by the [[Modern synthesis (20th century)|modern synthesis]], but the hopeful monster concept based on [[Evolutionary developmental biology]] (or evo-devo) explanations found a moderate revival in recent times.<ref>{{Cite journal|last=Theißen|first=Günter|date=March 2009|title=Saltational evolution: hopeful monsters are here to stay|journal=Theory in Biosciences|language=en|volume=128|issue=1|pages=43–51|doi=10.1007/s12064-009-0058-z|pmid=19224263|s2cid=4983539|issn=1431-7613}}</ref><ref>{{Cite book|last=Rieppel, Olivier|title=Turtles as hopeful monsters : origins and evolution|date=13 March 2017|isbn=978-0-253-02507-4|location=Bloomington, Indiana|oclc=962141060}}</ref> Occasionally such dramatic changes can lead to novel features that survive. As an alternative to saltational evolution, [[Dobzhansky]]<ref>{{Cite book|last=Dobzhanski|first=T.|title=Genetics and the origin of species.|publisher=Columbia University Press|year=1937}}</ref> suggested that the difference between macroevolution and microevolution reflects essentially a difference in time-scales, and that macroevolutionary changes were simply the sum of microevolutionary changes over geologic time. This view became broadly accepted, and accordingly, the term macroevolution has been used widely as a neutral label for the study of evolutionary changes that take place over a very large time-scale.<ref>{{Cite book|last=Dawkins, Richard, 1941-|title=The extended phenotype : the gene as the unit of selection|date=1982|publisher=Freeman|isbn=0-7167-1358-6|location=Oxford [Oxfordshire]|oclc=7652745}}</ref> Further, species selection<ref name=":0" /> suggests that selection among species is a major evolutionary factor that is independent from and complementary to selection among organisms. Accordingly, the level of selection has become the conceptual basis of a third definition, which defines macroevolution as evolution through selection among [[Interspecific competition|interspecific]] variation.<ref name=":1" />
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