Editing Macroevolution

{{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>

== 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" />

== Microevolution vs Macroevolution ==

There has been considerable debate regarding the connection between microevolution and macroevolution.<ref name="Saupe2021a"></ref>

The '''‘Extrapolation’''' view holds that macroevolution is merely cumulative microevolution. 

The '''‘Decoupled’''' view holds that there are separate macroevolutionary processes that cannot be sufficiently explained by microevolutionary processes alone.<ref>{{cite book |last=Ayala Francisco J |date=1983|name-list-style= and |editor-last1 = Asquith | editor-first1= Peter D|  editor-last2=Nickles|editor-first2= Thomas |title=PSA 1982 |volume=2|publisher=Philosophy of Science Association |pages=118–132 |chapter=Beyond Darwinism? The Challenge of Macroevolution to the Synthetic Theory of Evolution |isbn=}}</ref><ref name="Levinton2001">{{cite book | vauthors = Levinton Jeffrey S | date = 2001 | title = Genetics, Paleontology, and Macroevolution 2nd edition | publisher = Cambridge University Press | place = Cambridge, UK | isbn = 0-521-80317-9}}</ref><ref name="Rolland2022a"></ref><ref name="Simons2002a">{{cite journal |last1=Simons |first1=Andrew M. |title=The continuity of microevolution and macroevolution |journal=Journal of Evolutionary Biology |date=August 21, 2002 |volume=15 |issue=5 |pages=688–701 |doi=10.1046/j.1420-9101.2002.00437.x}}</ref><ref name="Erwin2001a">{{cite journal |last1=Erwin |first1=Douglas H. |title=Macroevolution is more than repeated rounds of microevolution |journal=Evolution & Development |date=December 24, 2001 |volume=2 |issue=2 |pages=78–84 |doi=10.1046/j.1525-142x.2000.00045.x|pmid=11258393 }}</ref><ref name="Adams1990a">{{cite journal |last1=Adams |first1=Mark B |title=Filipchenko [Philiptschenko], Iurii Aleksandrovich. |journal=Dictionary of Scientific Biography |date=1990 |volume=17 |issue=297–303 |url=https://www.encyclopedia.com/science/dictionaries-thesauruses-pictures-and-press-releases/filipchenko-philiptschenko-iurii-aleksandrovich}}</ref><ref name="DAOAL1"></ref><ref name="Moran2022a">{{cite web |last1=Moran |first1=Laurence A. |title=Macroevolution |url=https://sandwalk.blogspot.com/2022/10/macroevolution.html |website=Sandwalk Blog |date=October 13, 2022}}</ref>  

Within microevolution, the evolutionary process of changing heritable characteristics (e.g. changes in allele frequencies) is described by [[population genetics]], with mechanisms such as [[mutation]], [[natural selection]], and [[genetic drift]],<ref name=":0"></ref> and [[speciation]] (e.g. [[sympatric]] and [[allopatric]] speciation), [[phyletic gradualism]] and [[punctuated equilibrium]].<ref name="Saupe2021a"></ref> Macroevolution asks how higher taxonomic groups ([[genera]], [[Family_(biology)|families]], [[Order_(biology)|orders]], etc) have evolved across geography and vast spans of [[geological time]]. Important questions and topics include:
* How different species are related to each other is addressed by [[phylogenetics]].
* The rates of evolutionary change and across time in the [[fossil record]].<ref name="Rolland2022a"></ref> Why do some groups experience a lot of change while others remain morphologically stable? The latter case are often called '[[living fossils]]'.<ref name="Kin2014a">{{Cite journal|last1=Kin|first1=Adrian|last2=Błażejowski|first2=Błażej|date=2014-10-02|title=The Horseshoe Crab of the Genus Limulus: Living Fossil or Stabilomorph?|journal=PLOS ONE|language=en|volume=9|issue=10|pages=e108036|doi=10.1371/journal.pone.0108036|issn=1932-6203|pmc=4183490|pmid=25275563|bibcode=2014PLoSO...9j8036K|doi-access=free}}</ref>
* [[Mass extinctions]] and [[adaptive radiation|evolutionary diversifications]],<ref name="GEOL331a"></ref> e.g. the [[Permian-Triassic]] and [[End Cretaceous|Cretaceous-Paleogene]] events, the [[Cambrian Explosion]] and [[Cretaceous Terrestrial Revolution]].
* Why different taxonomic groups (even in spite of having similar ages) exhibit different survival/extinction rates, [[species diversity]], and/or [[Phenotypic_disparity|morphological disparity]]. 
* Long-term trends in evolution. Are these trends directed in some way, e.g. towards complexity or simplicity.<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> 
* How species and higher taxa have evolved. Examples of this include [[gene duplication]], [[heterochrony]], [[Evolutionary_developmental_biology#The_origins_of_novelty|novelty in evodevo]] from [[facilitated variation]], and [[constructive neutral evolution]].

==Macroevolutionary processes==

=== Speciation ===
{{Main|speciation}}
According to the modern definition, the evolutionary transition from the ancestral to the daughter species is microevolutionary, because it results from selection (or, more generally, sorting) among varying organisms.  However, speciation has also a macroevolutionary aspect, because it produces the interspecific variation species selection operates on.<ref name=":1" /> Another macroevolutionary aspect of speciation is the rate at which it successfully occurs, analogous to reproductive success in microevolution.<ref name=":0" />

Speciation is the process in which populations within one species change to an extent at which they become [[Reproductive isolation|reproductively isolated]], that is, they cannot interbreed anymore. However, this classical concept has been challenged and more recently, a phylogenetic or evolutionary [[species]] concept has been adopted. Their main criteria for new species is to be diagnosable and [[Monophyly|monophyletic]], that is, they form a clearly defined lineage.<ref>{{Cite journal |last=Luckow |first=Melissa |date=1995 |title=Species Concepts: Assumptions, Methods, and Applications |url=https://www.jstor.org/stable/2419812 |journal=Systematic Botany |volume=20 |issue=4 |pages=589–605 |doi=10.2307/2419812 |jstor=2419812 |issn=0363-6445|url-access=subscription }}</ref><ref>{{Cite journal |last1=Frost |first1=Darrel R. |last2=Hillis |first2=David M. |date=1990 |title=Species in Concept and Practice: Herpetological Applications |url=https://www.jstor.org/stable/3892607 |journal=Herpetologica |volume=46 |issue=1 |pages=86–104 |jstor=3892607 |issn=0018-0831}}</ref>

[[Charles Darwin]] first discovered that speciation can be extrapolated so that species not only evolve into new species, but also into new [[Genus|genera]], families and other groups of animals. In other words, macroevolution is reducible to microevolution through selection of traits over long periods of time.<ref>{{Cite journal|last=Greenwood|first=P. H.|title=Macroevolution - myth or reality ?|journal=Biological Journal of the Linnean Society|year=1979|volume=12|issue=4|pages=293–304|doi=10.1111/j.1095-8312.1979.tb00061.x}}</ref> In addition, some scholars have argued that selection at the species level is important as well.<ref>{{Cite journal|last=Grantham|first=T A|date=November 1995|title=Hierarchical Approaches to Macroevolution: Recent Work on Species Selection and the "Effect Hypothesis"|journal=Annual Review of Ecology and Systematics|language=en|volume=26|issue=1|pages=301–321|doi=10.1146/annurev.es.26.110195.001505|bibcode=1995AnRES..26..301G |issn=0066-4162}}</ref> The advent of genome sequencing enabled the discovery of gradual genetic changes both during speciation but also across higher taxa. For instance, the evolution of humans from ancestral primates or other mammals can be traced to numerous but individual mutations.<ref>{{Cite journal |last1=Foley |first1=Nicole M. |last2=Mason |first2=Victor C. |last3=Harris |first3=Andrew J. |last4=Bredemeyer |first4=Kevin R. |last5=Damas |first5=Joana |last6=Lewin |first6=Harris A. |last7=Eizirik |first7=Eduardo |last8=Gatesy |first8=John |last9=Karlsson |first9=Elinor K. |last10=Lindblad-Toh |first10=Kerstin |last11=Zoonomia Consortium‡ |last12=Springer |first12=Mark S. |last13=Murphy |first13=William J. |last14=Andrews |first14=Gregory |last15=Armstrong |first15=Joel C. |date=2023-04-28 |title=A genomic timescale for placental mammal evolution |journal=Science |language=en |volume=380 |issue=6643 |pages=eabl8189 |doi=10.1126/science.abl8189 |issn=0036-8075 |pmc=10233747 |pmid=37104581}}</ref>

=== Evolution of new organs and tissues ===
One of the main questions in evolutionary biology is how new structures evolve, such as new [[Organ (biology)|organs]]. Macroevolution is often thought to require the evolution of structures that are 'completely new'. However, fundamentally novel structures are not necessary for dramatic evolutionary change. As can be seen in [[Vertebrate|vertebrate evolution]], most "new" organs are actually not new—they are simply modifications of previously existing organs. For instance, the evolution of [[mammal]] diversity in the past 100 million years has not required any major innovation.<ref>{{Cite journal |last1=Meredith |first1=R. W. |last2=Janecka |first2=J. E. |last3=Gatesy |first3=J. |last4=Ryder |first4=O. A. |last5=Fisher |first5=C. A. |last6=Teeling |first6=E. C. |last7=Goodbla |first7=A. |last8=Eizirik |first8=E. |last9=Simao |first9=T. L. L. |last10=Stadler |first10=T. |last11=Rabosky |first11=D. L. |last12=Honeycutt |first12=R. L. |last13=Flynn |first13=J. J. |last14=Ingram |first14=C. M. |last15=Steiner |first15=C. |date=2011-10-28 |title=Impacts of the Cretaceous Terrestrial Revolution and KPg Extinction on Mammal Diversification |url=https://www.sciencemag.org/lookup/doi/10.1126/science.1211028 |journal=Science |language=en |volume=334 |issue=6055 |pages=521–524 |doi=10.1126/science.1211028 |pmid=21940861 |bibcode=2011Sci...334..521M |s2cid=38120449 |issn=0036-8075|url-access=subscription }}</ref> All of this diversity can be explained by modification of existing organs, such as the evolution of [[Tusk|elephant tusks]] from [[Incisor|incisors]]. Other examples include [[Bird wing|wings]] (modified limbs), [[feather]]s (modified [[reptile scale]]s),<ref>{{Cite journal |last1=Wu |first1=Ping |last2=Yan |first2=Jie |last3=Lai |first3=Yung-Chih |last4=Ng |first4=Chen Siang |last5=Li |first5=Ang |last6=Jiang |first6=Xueyuan |last7=Elsey |first7=Ruth M |last8=Widelitz |first8=Randall |last9=Bajpai |first9=Ruchi |last10=Li |first10=Wen-Hsiung |last11=Chuong |first11=Cheng-Ming |date=2017-11-21 |title=Multiple Regulatory Modules Are Required for Scale-to-Feather Conversion |url=|journal=Molecular Biology and Evolution |volume=35 |issue=2 |pages=417–430 |doi=10.1093/molbev/msx295 |issn=0737-4038 |pmc=5850302 |pmid=29177513}}</ref> [[lung]]s (modified [[swim bladder]]s, e.g. found in [[fish]]),<ref>{{Cite journal |last=Brainerd |first=E. L. |date=1999-12-01 |title=New perspectives on the evolution of lung ventilation mechanisms in vertebrates |url=|journal=Experimental Biology Online |language=en |volume=4 |issue=2 |pages=1–28 |doi=10.1007/s00898-999-0002-1 |bibcode=1999EvBO....4b...1B |s2cid=35368264 |issn=1430-3418}}</ref><ref>{{Cite journal |last1=Hoffman |first1=M. |last2=Taylor |first2=B. E. |last3=Harris |first3=M. B. |date=2016-04-01 |title=Evolution of lung breathing from a lungless primitive vertebrate |journal=Respiratory Physiology & Neurobiology |series=Physiology of respiratory networks of non-mammalian vertebrates |language=en |volume=224 |pages=11–16 |doi=10.1016/j.resp.2015.09.016 |issn=1569-9048 |pmc=5138057 |pmid=26476056}}</ref> or even the [[heart]] (a muscularized segment of a [[vein]]).<ref>{{Cite journal |last1=Jensen |first1=Bjarke |last2=Wang |first2=Tobias |last3=Christoffels |first3=Vincent M. |last4=Moorman |first4=Antoon F. M. |date=2013-04-01 |title=Evolution and development of the building plan of the vertebrate heart |journal=Biochimica et Biophysica Acta (BBA) - Molecular Cell Research |series=Cardiomyocyte Biology: Cardiac Pathways of Differentiation, Metabolism and Contraction |language=en |volume=1833 |issue=4 |pages=783–794 |doi=10.1016/j.bbamcr.2012.10.004 |pmid=23063530 |s2cid=28787569 |issn=0167-4889|doi-access=free }}</ref>

The same concept applies to the evolution of "novel" tissues. Even fundamental tissues such as [[bone]] can evolve from combining existing [[protein]]s ([[collagen]]) with calcium phosphate (specifically, [[Hydroxyapatite|hydroxy-apatite]]). This probably happened when certain cells that make collagen also accumulated calcium phosphate to get a proto-bone cell.<ref>{{Cite journal |last1=Wagner |first1=Darja Obradovic |last2=Aspenberg |first2=Per |date=2011-08-01 |title=Where did bone come from? |url=|journal=Acta Orthopaedica |volume=82 |issue=4 |pages=393–398 |doi=10.3109/17453674.2011.588861 |issn=1745-3674 |pmc=3237026 |pmid=21657973}}</ref>

=== Molecular macroevolution ===
Microevolution is facilitated by [[mutation]]s, the vast majority of which have no or very small effects on gene or protein function. For instance, the activity of an [[enzyme]] may be slightly changed or the stability of a protein slightly altered. However, occasionally mutations can dramatically change the structure and functions of protein. This may be called "molecular macroevolution".

[[File:PDB 2aj4 EBI.png|thumb|The metabolic enzyme [[galactokinase]] can be converted to a [[transcription factor]] (in [[Saccharomyces cerevisiae|yeast]]) by just a 2 amino-acid insertion.]]

'''Protein function'''. There are countless cases in which protein function is dramatically altered by mutations. For instance, a mutation in [[acetaldehyde dehydrogenase]] (EC:1.2.1.10) can change it to a [[4-hydroxy-2-oxovalerate aldolase|4-hydroxy-2-oxopentanoate pyruvate lyase]] (EC:4.1.3.39), i.e., a mutation that changes an [[enzyme]] from one to another [[Enzyme Commission number|EC]] class (there are only 7 main classes of enzymes).<ref>{{Cite journal |last1=Tyzack |first1=Jonathan D |last2=Furnham |first2=Nicholas |last3=Sillitoe |first3=Ian |last4=Orengo |first4=Christine M |last5=Thornton |first5=Janet M |date=2017-12-01 |title=Understanding enzyme function evolution from a computational perspective |journal=Current Opinion in Structural Biology |series=Protein–nucleic acid interactions • Catalysis and regulation |language=en |volume=47 |pages=131–139 |doi=10.1016/j.sbi.2017.08.003 |pmid=28892668 |issn=0959-440X|doi-access=free }}</ref> Another example is the conversion of a [[yeast]] [[galactokinase]] (Gal1) to a [[transcription factor]] (Gal3) which can be achieved by an insertion of only two amino acids.<ref>{{Cite journal |last1=Platt |first1=A. |last2=Ross |first2=H. C. |last3=Hankin |first3=S. |last4=Reece |first4=R. J. |date=2000-03-28 |title=The insertion of two amino acids into a transcriptional inducer converts it into a galactokinase |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=97 |issue=7 |pages=3154–3159 |doi=10.1073/pnas.97.7.3154 |issn=0027-8424 |pmc=16208 |pmid=10737789|bibcode=2000PNAS...97.3154P |doi-access=free }}</ref>

While some mutations may not change the molecular function of a protein significantly, their biological function may be dramatically changed. For instance, most brain receptors recognize specific neurotransmitters, but that specificity can easily be changed by mutations. This has been shown by [[acetylcholine receptor]]s that can be changed to [[serotonin]] or [[glycine receptor]]s which actually have very different functions. Their similar gene structure also indicates that they must have arisen from [[gene duplication]]s.<ref>{{Cite journal |last1=Uetz |first1=Peter |last2=Abdelatty |first2=Fawzy |last3=Villarroel |first3=Alfredo |last4=Rappold |first4=Gudrun |author-link4=Gudrun Rappold |last5=Weiss |first5=Birgit |last6=Koenen |first6=Michael |date=1994-02-21 |title=Organisation of the murine 5-HT 3 receptor gene and assignment tohuman chromosome 11 |journal=FEBS Letters |language=en |volume=339 |issue=3 |pages=302–306 |doi=10.1016/0014-5793(94)80435-4 |pmid=8112471 |s2cid=28979681 |doi-access=free|bibcode=1994FEBSL.339..302U }}</ref>

'''Protein structure'''. Although protein structures are highly conserved, sometimes one or a few mutations can dramatically change a protein. For instance, an [[Insulin-like growth factor-binding protein|IgG-binding]], 4<math>\beta</math>+<math>\alpha</math> fold can be transformed into an [[albumin]]-binding, 3-α fold via a single amino-acid mutation. This example also shows that such a transition can happen with neither function nor native structure being completely lost.<ref>{{Cite journal |last1=Alexander |first1=Patrick A. |last2=He |first2=Yanan |last3=Chen |first3=Yihong |last4=Orban |first4=John |last5=Bryan |first5=Philip N. |date=2009-12-15 |title=A minimal sequence code for switching protein structure and function |journal=Proceedings of the National Academy of Sciences |language=en |volume=106 |issue=50 |pages=21149–21154 |doi=10.1073/pnas.0906408106 |issn=0027-8424 |pmc=2779201 |pmid=19923431|doi-access=free }}</ref> In other words, even when multiple mutations are required to convert one protein or structure into another, the structure and function is at least partially retained in the intermediary sequences. Similarly, [[Protein domain|domains]] can be converted into other domains (and thus other functions). For instance, the structures of [[SH3 domain|SH3]] folds can evolve into [[OB-fold|OB folds]] which in turn can evolve into CLB folds.<ref>{{Cite journal |last1=Alvarez-Carreño |first1=Claudia |last2=Gupta |first2=Rohan J. |last3=Petrov |first3=Anton S. |last4=Williams |first4=Loren Dean |date=2022-12-27 |title=Creative destruction: New protein folds from old |journal=Proceedings of the National Academy of Sciences |language=en |volume=119 |issue=52 |pages=e2207897119 |doi=10.1073/pnas.2207897119 |doi-access=free |pmid=36534803 |pmc=9907106 |bibcode=2022PNAS..11907897A |s2cid=254907939 |issn=0027-8424}}</ref>

==Examples==

=== Evolutionary faunas ===
A macroevolutionary benchmark study is Sepkoski's<ref>{{Cite journal|last=Sepkoski|first=J. John|date=1981|title=A factor analytic description of the Phanerozoic marine fossil record|journal=Paleobiology|volume=7|issue=1|pages=36–53|doi=10.1017/s0094837300003778|bibcode=1981Pbio....7...36S |issn=0094-8373}}</ref><ref>{{Cite journal|last=Sepkoski|first=J. John|date=1984|title=A kinetic model of Phanerozoic taxonomic diversity. III. Post-Paleozoic families and mass extinctions|journal=Paleobiology|volume=10|issue=2|pages=246–267|doi=10.1017/s0094837300008186|bibcode=1984Pbio...10..246S |issn=0094-8373}}</ref> work on marine animal diversity through the Phanerozoic. His iconic diagram of the numbers of marine families from the Cambrian to the Recent illustrates the successive expansion and dwindling of three "[[evolutionary fauna]]s" that were characterized by differences in origination rates and carrying capacities. Long-term ecological changes and major geological events are postulated to have played crucial roles in shaping these evolutionary faunas.<ref name="Rojas2021a">{{cite journal |last1=Rojas |first1=A. |last2=Calatayud |first2=J. |last3=Kowalewski |first3=M. |last4=Neuman |first4=M. |last5=Rosvall |first5=M. |title=A multiscale view of the Phanerozoic fossil record reveals the three major biotic transitions. |journal=Communications Biology |date=March 8, 2021 |volume=4 |issue=1 |page=309 |doi=10.1038/s42003-021-01805-y |pmid=33686149 |issn=2399-3642|pmc=7977041 }}</ref>

=== Stanley's rule ===
Macroevolution is driven by differences between species in origination and extinction rates. Remarkably, these two factors are generally positively correlated: taxa that have typically high diversification rates also have high extinction rates. This observation has been described first by [[Steven M. Stanley|Steven Stanley]], who attributed it to a variety of ecological factors.<ref>{{Cite book|last=Stanley, Steven M.|title=Macroevolution, pattern and process|date=1979|publisher=W.H. Freeman|isbn=0-7167-1092-7|location=San Francisco|oclc=5101557}}</ref> Yet, a positive correlation of origination and extinction rates is also a prediction of the [[Red Queen hypothesis]], which postulates that evolutionary progress (increase in fitness) of any given species causes a decrease in fitness of other species, ultimately driving to extinction those species that do not adapt rapidly enough.<ref>{{Cite journal|last=Van Valen|first=L.|date=1973|title=A new evolutionary law|journal=Evolutionary Theory|volume=1|pages=1–30}}</ref> High rates of origination must therefore correlate with high rates of extinction.<ref name=":1" /> Stanley's rule, which applies to almost all taxa and geologic ages, is therefore an indication for a dominant role of biotic interactions in macroevolution.

=== "Macromutations": Single mutations leading to dramatic change ===
{{Multiple image
| image1            = 202208 Fruit fly female adult from a overhead view.svg
| image2            = 202208 Fruit fly bithorax complex.svg
| footer            = Mutations in the [[Ultrabithorax]] gene lead to a duplication of wings in fruit flies.
| total_width       = 300
| caption1          = Normal phenotype
| caption2          = Bithorax phenotype
| caption_align     = center
}}
While the vast majority of mutations are inconsequential, some can have a dramatic effect on morphology or other features of an organism. One of the best studied cases of a single mutation that leads to massive structural change is the [[Ultrabithorax]] mutation in [[Drosophila melanogaster|fruit flies.]] The mutation duplicates the wings of a fly to make it look like a [[dragonfly]], a different order of insect.

=== Evolution of multicellularity ===
{{Main|Multicellular organism}}
The evolution of multicellular organisms is one of the major breakthroughs in evolution. The first step of converting a unicellular organism into a [[Animal|metazoan]] (a multicellular organism) is to allow cells to attach to each other. This can be achieved by one or a few [[mutation]]s. In fact, many [[bacteria]] form multicellular assemblies, e.g. [[cyanobacteria]] or [[myxobacteria]]. Another species of bacteria, ''Jeongeupia sacculi'', form well-ordered sheets of cells, which ultimately develop into a bulbous structure.<ref>{{Cite journal |last1=Datta |first1=Sayantan |last2=Ratcliff |first2=William C |date=2022-10-11 |title=Illuminating a new path to multicellularity |journal=eLife |volume=11 |pages=e83296 |doi=10.7554/eLife.83296 |pmid=36217823 |issn=2050-084X |pmc=9553208 |doi-access=free }}</ref><ref>{{Cite journal |last1=Mizuno |first1=Kouhei |last2=Maree |first2=Mais |last3=Nagamura |first3=Toshihiko |last4=Koga |first4=Akihiro |last5=Hirayama |first5=Satoru |last6=Furukawa |first6=Soichi |last7=Tanaka |first7=Kenji |last8=Morikawa |first8=Kazuya |date=2022-10-11 |editor-last=Goldstein |editor-first=Raymond E |editor2-last=Weigel |editor2-first=Detlef |title=Novel multicellular prokaryote discovered next to an underground stream |journal=eLife |volume=11 |pages=e71920 |doi=10.7554/eLife.71920 |pmid=36217817 |pmc=9555858 |issn=2050-084X |doi-access=free }}</ref> Similarly, unicellular yeast cells can become multicellular by a single mutation in the ACE2 gene, which causes the cells to form a branched multicellular form.<ref>{{Cite journal |last1=Ratcliff |first1=William C. |last2=Fankhauser |first2=Johnathon D. |last3=Rogers |first3=David W. |last4=Greig |first4=Duncan |last5=Travisano |first5=Michael |date=May 2015 |title=Origins of multicellular evolvability in snowflake yeast |journal=Nature Communications |language=en |volume=6 |issue=1 |pages=6102 |doi=10.1038/ncomms7102 |issn=2041-1723 |pmc=4309424 |pmid=25600558|bibcode=2015NatCo...6.6102R }}</ref>

=== Evolution of bat wings ===
The wings of [[bat]]s have the same structural elements (bones) as any other five-fingered mammal (see [[Limb development|periodicity in limb development]]). However, the finger bones in bats are dramatically elongated, so the question is how these bones became so long. It has been shown that certain growth factors such as [[bone morphogenetic protein]]s (specifically [[Bone morphogenetic protein 2|Bmp2]]) is over expressed so that it stimulates an elongation of certain bones. Genetic changes in the bat genome identified the changes that lead to this phenotype and it has been recapitulated in mice: when specific bat DNA is inserted in the mouse genome, recapitulating these mutations, the bones of mice grow longer.<ref name=":4">{{Cite journal |last1=Sears |first1=Karen E. |last2=Behringer |first2=Richard R. |last3=Rasweiler |first3=John J. |last4=Niswander |first4=Lee A. |date=2006-04-25 |title=Development of bat flight: Morphologic and molecular evolution of bat wing digits |journal=Proceedings of the National Academy of Sciences |language=en |volume=103 |issue=17 |pages=6581–6586 |doi=10.1073/pnas.0509716103 |issn=0027-8424 |pmc=1458926 |pmid=16618938|bibcode=2006PNAS..103.6581S |doi-access=free }}</ref>

=== Limb loss in lizards and snakes ===
{{main|Limbless vertebrates}}
[[File:Vine-thicket Fine-lined Slider (Lerista cinerea).jpg|thumb|Limbloss in lizards can be observed in the genus ''[[Lerista]]'' which shows many intermediary steps with increasing loss of digits and toes. The species shown here, ''[[Lerista cinerea]]'', has no digits and only 1 toe left.]]
[[Snake]]s evolved from [[lizard]]s. [[Phylogenetics|Phylogenetic]] analysis shows that snakes are actually nested within the [[phylogenetic tree]] of lizards, demonstrating that they have a common ancestor.<ref>{{Cite journal |last1=Streicher |first1=Jeffrey W. |last2=Wiens |first2=John J. |date=2017-09-30 |title=Phylogenomic analyses of more than 4000 nuclear loci resolve the origin of snakes among lizard families |journal=Biology Letters |volume=13 |issue=9 |pages=20170393 |doi=10.1098/rsbl.2017.0393 |pmc=5627172 |pmid=28904179}}</ref> This split happened about 180 million years ago and several intermediary [[fossil]]s are known to document the origin. In fact, limbs have been lost in numerous clades of [[reptile]]s, and there are cases of recent [[Limbless vertebrate|limb loss]]. For instance, the [[skink]] genus ''[[Lerista]]'' has lost limbs in multiple cases, with all possible intermediary steps, that is, there are species which have fully developed limbs, shorter limbs with 5, 4, 3, 2, 1 or no toes at all.<ref>{{Cite journal |last1=Skinner |first1=Adam |last2=Lee |first2=Michael SY |last3=Hutchinson |first3=Mark N |date=2008 |title=Rapid and repeated limb loss in a clade of scincid lizards |journal=BMC Evolutionary Biology |language=en |volume=8 |issue=1 |pages=310 |doi=10.1186/1471-2148-8-310 |issn=1471-2148 |pmc=2596130 |pmid=19014443 |doi-access=free |bibcode=2008BMCEE...8..310S }}</ref>

=== Human evolution ===
While human evolution from their primate ancestors did not require massive morphological changes, our brain has sufficiently changed to allow human consciousness and intelligence. While the latter involves relatively minor morphological changes it did result in dramatic changes to [[Brain|brain function]].<ref>{{Cite book |url=https://www.worldcat.org/oclc/903489046 |title=Macroevolution: explanation, interpretation and evidence |date=2015 |first1=Emanuele |last1=Serrelli |first2=Nathalie |last2=Gontier |isbn=978-3-319-15045-1 |location=Cham |oclc=903489046}}</ref> Thus, macroevolution does not have to be morphological, it can also be functional.

=== Evolution of viviparity in lizards ===
[[File:Zootoca vivipara. 3epo.Post.jpg|thumb|The European Common Lizard (''[[Viviparous lizard|Zootoca vivipara]]'') consists of populations that are egg-laying or live-bearing, demonstrating that this dramatic difference can even evolve within a species.]]
Most lizards are egg-laying and thus need an environment that is warm enough to incubate their eggs. However, some species have evolved [[viviparity]], that is, they give birth to live young, as almost all [[mammal]]s do. In several clades of lizards, egg-laying (oviparous) species have evolved into live-bearing ones, apparently with very little genetic change. For instance, a European common lizard, [[Viviparous lizard|''Zootoca vivipara'']], is viviparous throughout most of its range, but oviparous in the extreme southwest portion.<ref>{{Cite journal |last=Heulin |first=Benoît |date=1990-05-01 |title=Étude comparative de la membrane coquillère chez les souches ovipare et vivipare du lézard Lacerta vivipara |url=http://www.nrcresearchpress.com/doi/10.1139/z90-147 |journal=Canadian Journal of Zoology |language=en |volume=68 |issue=5 |pages=1015–1019 |doi=10.1139/z90-147 |bibcode=1990CaJZ...68.1015H |issn=0008-4301|url-access=subscription }}</ref><ref>{{Cite journal |last1=Arrayago |first1=Maria-Jesus |last2=Bea |first2=Antonio |last3=Heulin |first3=Benoit |date=1996 |title=Hybridization Experiment between Oviparous and Viviparous Strains of Lacerta vivipara: A New Insight into the Evolution of Viviparity in Reptiles |url=https://www.jstor.org/stable/3892653 |journal=Herpetologica |volume=52 |issue=3 |pages=333–342 |jstor=3892653 |issn=0018-0831}}</ref> That is, within a single species, a radical change in reproductive behavior has happened. Similar cases are known from South American lizards of the genus ''[[Liolaemus]]'' which have egg-laying species at lower altitudes, but closely related viviparous species at higher altitudes, suggesting that the switch from oviparous to viviparous reproduction does not require many genetic changes.<ref>{{Cite journal |last1=Ii |first1=James A. Schulte |last2=Macey |first2=J. Robert |last3=Espinoza |first3=Robert E. |last4=Larson |first4=Allan |date=January 2000 |title=Phylogenetic relationships in the iguanid lizard genus Liolaemus: multiple origins of viviparous reproduction and evidence for recurring Andean vicariance and dispersal |journal=Biological Journal of the Linnean Society |language=en |volume=69 |issue=1 |pages=75–102 |doi=10.1111/j.1095-8312.2000.tb01670.x|doi-access=free }}</ref>

=== Behavior: Activity pattern in mice ===
Most animals are either active at night or during the day. However, some species switched their activity pattern from day to night or vice versa. For instance, the African striped mouse (''[[Four-striped grass mouse|Rhabdomys pumilio]]''), transitioned from the ancestrally [[Nocturnality|nocturnal]] behavior of its close relatives to a [[Diurnality|diurnal]] one. [[Whole genome sequencing|Genome sequencing]] and [[Transcriptomics technologies|transcriptomics]] revealed that this transition was achieved by modifying genes in the [[Rod cell|rod]] [[Visual phototransduction|phototransduction]] pathway, among others.<ref>{{Cite journal |last1=Richardson |first1=Rose |last2=Feigin |first2=Charles Y. |last3=Bano-Otalora |first3=Beatriz |last4=Johnson |first4=Matthew R. |last5=Allen |first5=Annette E. |last6=Park |first6=Jongbeom |last7=McDowell |first7=Richard J. |last8=Mereby |first8=Sarah A. |last9=Lin |first9=I-Hsuan |last10=Lucas |first10=Robert J. |last11=Mallarino |first11=Ricardo |date=August 2023 |title=The genomic basis of temporal niche evolution in a diurnal rodent |url=|journal=Current Biology |volume=33 |issue=15 |pages=3289–3298.e6 |doi=10.1016/j.cub.2023.06.068 |pmid=37480852 |pmc=10529858 |bibcode=2023CBio...33E3289R |issn=0960-9822 }}</ref>

==Research topics==
Subjects studied within macroevolution include:<ref>Grinin, L., Markov, A. V., Korotayev, A. Aromorphoses in Biological and Social Evolution: Some General Rules for Biological and Social Forms of Macroevolution / Social evolution & History, vol.8, num. 2, 2009 [http://www.socionauki.ru/journal/articles/129272/]</ref>
* [[Adaptive radiation]]s such as the [[Cambrian Explosion]].
* Changes in [[Biodiversity#Evolution and history|biodiversity]] through time.
* [[Evolutionary developmental biology|Evo-devo]] (the connection between evolution and [[developmental biology]])
* [[Genome evolution]], like [[horizontal gene transfer]], genome fusions in endosymbioses, and adaptive changes in genome size.
* [[Extinction event|Mass extinctions]].
* Estimating [[diversification rates]], including rates of [[speciation]] and [[extinction]].
* The debate between [[punctuated equilibrium]] and [[gradualism]].
* The role of development in shaping evolution, particularly such topics as [[heterochrony]] and [[phenotypic plasticity]].

==See also==
* [[Extinction event]]
* [[Interspecific competition]]
* [[Microevolution]]
* [[Molecular evolution]]
* [[Punctuated equilibrium]]
* [[Red Queen hypothesis]]
* [[Speciation]]
* [[Transitional fossil]]
* [[Unit of selection]]

==Notes==
{{notelist}}


==References==
{{Reflist}}

==Further reading==
* What is marcroevolution? (pdf) https://onlinelibrary.wiley.com/doi/full/10.1111/pala.12465
* {{cite web|last=AAAS |first=American Association for the Advancement of Science |author-link=American Association for the Advancement of Science |date=16 February 2006 |title=Statement on the Teaching of Evolution |publisher=aaas.org |url=http://www.aaas.org/news/releases/2006/pdf/0219boardstatement.pdf |access-date=2007-01-14 |url-status=dead |archive-url=https://web.archive.org/web/20060221125539/http://www.aaas.org/news/releases/2006/pdf/0219boardstatement.pdf |archive-date=21 February 2006}}
* {{cite book|last=IAP |first=Interacademy Panel |date=2006-06-21 |title=IAP Statement on the Teaching of Evolution |publisher=interacademies.net |url=http://www.interacademies.net/Object.File/Master/6/150/Evolution%20statement.pdf |access-date=2007-01-14 |url-status=dead |archive-url=https://web.archive.org/web/20060705140010/http://www.interacademies.net/Object.File/Master/6/150/Evolution%20statement.pdf |archive-date=5 July 2006}}
* {{cite journal |last=Myers |first=P.Z. |author-link=PZ Myers |date=2006-06-18 |title=Ann Coulter: No Evidence for Evolution? |journal=[[Pharyngula (blog)|Pharyngula]] |publisher=[[ScienceBlogs]] |url=http://scienceblogs.com/pharyngula/2006/06/ann_coulter_no_evidence_for_ev.php |access-date=2007-09-12 |url-status=dead |archive-url=https://web.archive.org/web/20060622031856/http://scienceblogs.com/pharyngula/2006/06/ann_coulter_no_evidence_for_ev.php |archive-date=22 June 2006 }}
* {{Cite web |last=NSTA |first=National Science Teachers Association |author-link=National Science Teachers Association |year=2007 |title=An NSTA Evolution Q&A |url=http://www.nsta.org/publications/evolution.aspx |access-date=2008-02-01 |url-status=dead |archive-url=https://web.archive.org/web/20080202043206/http://www.nsta.org/publications/evolution.aspx |archive-date=2 February 2008 }}
* {{cite web |last=Pinholster |first=Ginger |date=19 February 2006 |title=AAAS Denounces Anti-Evolution Laws as Hundreds of K-12 Teachers Convene for 'Front Line' Event |publisher=aaas.org |url=http://www.aaas.org/news/releases/2006/0219boardstatement.shtml |access-date=2007-01-14 |archive-date=19 October 2013 |archive-url=https://web.archive.org/web/20131019171834/http://www.aaas.org/news/releases/2006/0219boardstatement.shtml |url-status=dead }}

==External links==
* [http://evolution.berkeley.edu/evolibrary/article/0_0_0/evo_47 Introduction to macroevolution]
* [http://www.talkorigins.org/faqs/comdesc/ Macroevolution as the common descent of all life]
* [http://www.nhm.ac.uk/hosted_sites/paleonet/paleo21/mevolution.html Macroevolution in the 21st century] Macroevolution as an independent discipline.
* [http://www.talkorigins.org/faqs/macroevolution.html Macroevolution FAQ]

{{evolution}}

[[Category:Evolutionary biology]]