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Macroevolution
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{{short description|Evolution on a scale at or above the level of species}} {{Use dmy dates|date=August 2021}} {{Evolutionary biology}} '''Macroevolution''' comprises the evolutionary processes and patterns which occur at and above the [[species]] level.<ref name="Saupe2021a">{{cite book |last1=Saupe |first1=Erin E. |last2=Myers |first2=Corinne E. |editor1-last=Nuño de la Rosa |editor1-first=Laura |editor2-last=Müller |editor2-first=Gerd B. |title=Chapter: Macroevolution, Book: Evolutionary Developmental Biology - A Reference Guide |date=April 1, 2021 |publisher=Springer, Cham. |isbn=978-3-319-32979-6 |pages=149–167 |edition=1 |chapter-url=https://doi.org/10.1007/978-3-319-32979-6_126 |chapter=Macroevolution|doi=10.1007/978-3-319-32979-6_126 }}</ref><ref name=":0">{{cite journal |last=Stanley|first=S. M. |date=1975-02-01 |title=A theory of evolution above the species level |journal=[[Proceedings of the National Academy of Sciences]] |language=en |volume=72 |issue=2 |pages=646–50 |doi=10.1073/pnas.72.2.646 |issn=0027-8424 |pmc=432371 |pmid=1054846 |bibcode=1975PNAS...72..646S |doi-access=free}}</ref><ref name="Gould2002a">{{cite book |last=Gould|first=Stephen Jay |title=The structure of evolutionary theory |date=2002 |publisher=Belknap Press of [[Harvard University Press]] |isbn=0-674-00613-5 |location=Cambridge, Mass. |oclc=47869352}}</ref> In contrast, [[microevolution]] is evolution occurring within the population(s) of a single species. In other words, microevolution is the scale of evolution that is limited to intraspecific (within-species) variation, while macroevolution extends to interspecific (between-species) variation.<ref name=":1">{{cite journal |last=Hautmann|first=Michael |date=2020 |title=What is macroevolution?|journal=[[Palaeontology (journal)|Palaeontology]] |language=en |volume=63 |issue=1 |pages=1–11 |doi=10.1111/pala.12465 |bibcode=2020Palgy..63....1H |issn=0031-0239 |doi-access=free}}</ref> The evolution of new species ([[speciation]]) is an example of macroevolution. This is the common definition for 'macroevolution' used by contemporary scientists.{{efn|Rolland et al. (2023)<ref name="Rolland2022a">{{cite journal |last1=Rolland |first1=J. |last2=Henao-Diaz |first2=L.F. |last3=Doebeli |first3=M.|last4=Germain |first4=Rachel |display-authors=3 |title=Conceptual and empirical bridges between micro- and macroevolution. |journal=Nature Ecology & Evolution |date=July 10, 2023 |volume=7 |issue=8 |pages=1181–1193 |doi=10.1038/s41559-023-02116-7 |pmid=37429904 |bibcode=2023NatEE...7.1181R |url=https://files.zoology.ubc.ca/mank-lab/pdf/2023NEEGaps.pdf |issn=2397-334X}}</ref> in the introduction describe ‘microevolution’ and ‘macroevolution’ occurring at two different scales; below the species level and at/above the species level respectively: ''“Since the modern synthesis, many evolutionary biologists have focused their attention on evolution at one of two different timescales: microevolution, that is, the evolution of populations below the species level (in fields such as population genetics, phylogeography and quantitative genetics), or macroevolution, that is, the evolution of species or higher taxonomic levels (for example, phylogenetics, palaeobiology and biogeography).”''}}{{efn| Saupe & Myers (2021)<ref name="Saupe2021a"></ref> states: ''“Macroevolution is the study of patterns and processes associated with evolutionary change at and above the species level, and includes investigations of both evolutionary tempo and mode.”''}}{{efn| Michael Hautmann (2019)<ref name=":1"></ref> discusses 3 categories of definitions that have been historically used. He argues in favor of the following definition [added clarity]: ''"Macroevolution is evolutionary change that is guided by sorting of interspecific [between-species] variation."''}}{{efn| David Jablonski (2017)<ref name="Jablosnki2017a">{{cite journal |last1=Jablonski |first1=D. |title=Approaches to Macroevolution: 1. General Concepts and Origin of Variation. |journal=Springer, Evolutionary Biology |date=June 3, 2017 |volume=44 |issue=4 |pages=427–450 |doi=10.1007/s11692-017-9420-0|pmid=29142333 |pmc=5661017 |bibcode=2017EvBio..44..427J }}</ref><ref name="Jablosnki2017b">{{cite journal |last1=Jablonski |first1=D. |title=Approaches to Macroevolution: 2. Sorting of Variation, Some Overarching Issues, and General Conclusions. |journal=Springer, Evolutionary Biology |date=October 24, 2017 |volume=44 |issue=4 |pages=451–475 |doi=10.1007/s11692-017-9434-7|pmid=29142334 |pmc=5661022 |bibcode=2017EvBio..44..451J }}</ref> states: “Macroevolution, defined broadly as evolution above the species level, is thriving as a field.”}}{{efn| In his book “The Structure of Evolutionary Theory” (2002)<ref name="Gould2002a"></ref> page 612, Stephen J. Gould describes the species as the basic unit of macroevolution, and compares speciation and extinction to birth and death in microevolutionary processes respectively: ''“In particular, and continuing to use species as a “type” example of individuality at higher levels, all evolutionary criteria apply to the species as a basic unit of macro-evolution. Species have children by branching (in our professional jargon, we even engender these offspring as “daughter species”). Speciation surely obeys principles of hereditary, for daughters, by strong constraints of homology, originate with phenotypes and genotypes closer to those of their parent than to any other species of a collateral lineage. Species certainly vary, for the defining property of reproductive isolation demands genetic differentiation from parents and collateral relatives. Finally, species interact with the environment in a causal way that can influence rates of birth (speciation) and death (extinction).”''}}{{efn| In his paper proposing the theory of [[species selection]], Steven M. Stanly (1974)<ref name=":0"></ref> described macroevolution as being evolution above the species level and decoupled from microevolution: ''“In reaction to the arguments of macromutationists who opposed Neo-Darwinism, modern evolutionists have forcefully asserted that the process of natural selection is responsible for both microevolution, or evolution within species, and evolution above the species level, which is also known as macroevolution or transpecific evolution. [...] Macroevolution is decoupled from microevolution, and we must envision the process governing its course as being analogous to natural selection but operating at a higher level of biological organization. In this higher-level process species become analogous to individuals, and speciation replaces reproduction”''}}{{efn| The ‘Understanding Evolution’ website<ref name="BerkeleyEdu1">{{cite web |title=Evolution at different scales |url=https://evolution.berkeley.edu/evolution-at-different-scales-micro-to-macro/ |website=Understanding Evolution |publisher=UCMP, Berkely}}</ref> by [[UCMP]]: ''“Microevolution happens on a small scale (within a single population), while macroevolution happens on a scale that transcends the boundaries of a single species”''}}{{efn| Thomas Holtz’s course GEOL331 lecture notes<ref name="GEOL331a">{{cite web |title=Macroevolution in the Fossil Record? |url=https://www.geol.umd.edu/~tholtz/G331/lectures/331macroevo.html |website=GEOL331 Lecture Notes |publisher=University of Maryland Department of Geology}}</ref> discusses macroevolution observed in the fossil record:''“Following these early attempted modifications of Darwinism, the rest of the 20th Century onward stayed largely within a Darwinian model. However, there were different major schools of thought. Many of these differences hinged on views of microevolution (evolutionary change within a species) and macroevolution (evolutionary change above the species level). While most agreed that the ultimate processes in macroevolution were ultimately microevolutionary, there were disagreement[s] whether the patterns produced were actually reducible to microevolutionary changes.”''}}{{efn| The ‘Digital Atlas of Ancient Life’ website<ref name="DAOAL1">{{cite web |title=What is Macroevolution? |url=https://www.digitalatlasofancientlife.org/learn/evolution/macroevolution/ |website=Digital Atlas of Ancient Life |publisher=PRI}}</ref> by [[Paleontological Research Institution|PRI]] provides a very detailed historical overview for the definition of ‘macroevolution’: “The meaning of the term “macroevolution” has shifted over time. Indeed, early definitions do to not necessarily make much sense in light of our current understanding of evolution, yet are still worth considering to show how the field itself has evolved. Here we will consider usage of the term macroevolution in a few key works, as well as present a definition of macroevolution that we endorse. [...] Lieberman and Eldredge (2014) defined macroevolution as “the patterns and processes pertaining to the birth, death, and persistence of species” and we adopt this definition here.”}} Although, the exact usage of the term has varied throughout history.<ref name=":1"></ref><ref name="DAOAL1"></ref><ref name=":2">{{cite book |last=Filipchenko|first=J. |title=Variabilität und Variation |publisher=[[Gebrüder Borntraeger Verlagsbuchhandlung|Borntraeger]] |year=1927 |location=Berlin}}</ref> Macroevolution addresses the evolution of species and higher taxonomic groups ([[genera]], [[Family_(biology)|families]], [[Order_(biology)|orders]], etc) and uses evidence from [[phylogenetics]],<ref name="Rolland2022a"></ref> the fossil record,<ref name="GEOL331a"></ref> and molecular biology to answer how different taxonomic groups exhibit different [[species diversity]] and/or [[Phenotypic_disparity|morphological disparity]].<ref name="Gregory2008a">{{cite journal |last1=Gregory |first1=T.R. |title=Evolutionary Trends |journal=Evo Edu Outreach |date=June 25, 2008 |volume=1 |issue=3 |pages=259–273 |doi=10.1007/s12052-008-0055-6 |issn=1936-6434|doi-access=free }}</ref>
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