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Convergent evolution

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Convergent evolutionis the independentevolutionof similar features in species of different periods or epochs in time. Convergent evolution createsanalogous structuresthat have similar form or function but were not present in thelast common ancestorof those groups. Thecladisticterm for the same phenomenon ishomoplasy.Therecurrent evolutionof flight is a classic example, as flyinginsects,birds,pterosaurs,andbatshave independently evolved the useful capacity of flight. Functionally similar features that have arisen through convergent evolution areanalogous,whereashomologousstructures or traits have a common origin but can have dissimilar functions. Bird, bat, and pterosaurwingsare analogous structures, but their forelimbs are homologous, sharing an ancestral state despite serving different functions.

Twosucculent plantgenera,EuphorbiaandAstrophytum,are only distantly related, but the species within each have converged on a similar body form.

The opposite of convergence isdivergent evolution,where related species evolve different traits. Convergent evolution is similar toparallel evolution,which occurs when two independent species evolve in the same direction and thus independently acquire similar characteristics; for instance,gliding frogshave evolved in parallel from multiple types oftree frog.

Many instances of convergent evolution are known inplants,including the repeated development ofC4photosynthesis,seed dispersalby fleshyfruitsadapted to be eaten by animals, andcarnivory.

Overview

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Homologyand analogy in mammals and insects: on the horizontal axis, the structures are homologous in morphology, but different in function due to differences in habitat. On the vertical axis, the structures are analogous in function due to similar lifestyles but anatomically different with differentphylogeny.[a]
Further information:List of examples of convergent evolution

In morphology, analogous traits arise when different species live in similar ways and/or a similar environment, and so face the same environmental factors. When occupying similarecological niches(that is, a distinctive way of life) similar problems can lead to similar solutions.[1][2][3]The British anatomistRichard Owenwas the first to identify the fundamental difference between analogies andhomologies.[4]

In biochemistry, physical and chemical constraints onmechanismshave caused someactive sitearrangements such as thecatalytic triadto evolve independently in separateenzyme superfamilies.[5]

In his 1989 bookWonderful Life,Stephen Jay Gouldargued that if one could "rewind the tape of life [and] the same conditions were encountered again, evolution could take a very different course."[6]Simon Conway Morrisdisputes this conclusion, arguing that convergence is a dominant force in evolution, and given that the same environmental and physical constraints are at work, life will inevitably evolve toward an "optimum" body plan, and at some point, evolution is bound to stumble upon intelligence, a trait presently identified with at leastprimates,corvids,andcetaceans.[7]

Distinctions

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Cladistics

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Main article:Cladistics

In cladistics, a homoplasy is a trait shared by two or moretaxafor any reason other than that they share a common ancestry. Taxa which do share ancestry are part of the sameclade;cladistics seeks to arrange them according to their degree of relatedness to describe theirphylogeny.Homoplastic traits caused by convergence are therefore, from the point of view of cladistics, confounding factors which could lead to an incorrect analysis.[8][9][10][11]

Atavism

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Main article:Atavism

In some cases, it is difficult to tell whether a trait has been lost and then re-evolved convergently, or whether a gene has simply been switched off and then re-enabled later. Such a re-emerged trait is called anatavism.From a mathematical standpoint, an unused gene (selectively neutral) has a steadily decreasingprobabilityof retaining potential functionality over time. The time scale of this process varies greatly in different phylogenies; in mammals and birds, there is a reasonable probability of remaining in the genome in a potentially functional state for around 6 million years.[12]

Parallel vs. convergent evolution

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Evolution at anamino acidposition. In each case, the left-hand species changes from having alanine (A) at a specific position in a protein in a hypothetical ancestor, and now has serine (S) there. The right-hand species may undergodivergent,parallel, or convergent evolution at this amino acid position relative to the first species.

When two species are similar in a particular character, evolution is defined as parallel if the ancestors were also similar, and convergent if they were not.[b]Some scientists have argued that there is a continuum between parallel and convergent evolution,[13][14][15][16]while others maintain that despite some overlap, there are still important distinctions between the two.[17][18]

When the ancestral forms are unspecified or unknown, or the range of traits considered is not clearly specified, the distinction between parallel and convergent evolution becomes more subjective. For instance, the striking example of similar placental and marsupial forms is described byRichard DawkinsinThe Blind Watchmakeras a case of convergent evolution, because mammals on each continent had a long evolutionary history prior to the extinction of the dinosaurs under which to accumulate relevant differences.[19]

At molecular level

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Evolutionary convergence ofserineandcysteine proteasetowards the same catalytic triads organisation of acid-base-nucleophile in differentprotease superfamilies.Shown are the triads ofsubtilisin,prolyl oligopeptidase,TEV protease,andpapain.

Proteins

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Protease active sites

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Main article:Catalytic triad

Theenzymologyofproteasesprovides some of the clearest examples of convergent evolution. These examples reflect the intrinsic chemical constraints on enzymes, leading evolution to converge on equivalent solutions independently and repeatedly.[5][20]

Serine and cysteine proteases use different amino acid functional groups (alcohol or thiol) as anucleophile.In order to activate that nucleophile, they orient an acidic and a basic residue in acatalytic triad.The chemical and physical constraints onenzyme catalysishave caused identical triad arrangements to evolve independently more than 20 times in differentenzyme superfamilies.[5]

Threonine proteasesuse the amino acid threonine as their catalyticnucleophile.Unlike cysteine and serine, threonine is asecondary alcohol(i.e. has a methyl group). The methyl group of threonine greatly restricts the possible orientations of triad and substrate, as the methyl clashes with either the enzyme backbone or the histidine base. Consequently, most threonine proteases use an N-terminal threonine in order to avoid suchsteric clashes. Several evolutionarily independentenzyme superfamilieswith differentprotein foldsuse the N-terminal residue as a nucleophile. This commonality ofactive sitebut difference of protein fold indicates that the active site evolved convergently in those families.[5][21]

Cone snail and fish insulin

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Conus geographusproduces a distinct form ofinsulinthat is more similar to fish insulin protein sequences than to insulin from more closely related molluscs, suggesting convergent evolution,[22]though with the possibility ofhorizontal gene transfer.[23]

Ferrous iron uptake via protein transporters in land plants and chlorophytes

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Distant homologues of the metal ion transportersZIPinland plantsandchlorophyteshave converged in structure, likely to take up Fe2+efficiently. The IRT1 proteins fromArabidopsis thalianaandricehave extremely different amino acid sequences fromChlamydomonas's IRT1, but their three-dimensional structures are similar, suggesting convergent evolution.[24]

Na+,K+-ATPase and Insect resistance to cardiotonic steroids

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Many examples of convergent evolution exist in insects in terms of developing resistance at a molecular level to toxins. One well-characterized example is the evolution of resistance to cardiotonic steroids (CTSs) via amino acid substitutions at well-defined positions of the α-subunit ofNa+,K+-ATPase(ATPalpha). Variation in ATPalpha has been surveyed in various CTS-adapted species spanning six insect orders.[25][26][27]Among 21 CTS-adapted species, 58 (76%) of 76 amino acid substitutions at sites implicated in CTS resistance occur in parallel in at least two lineages.[27]30 of these substitutions (40%) occur at just two sites in the protein (positions 111 and 122). CTS-adapted species have also recurrently evolvedneo-functionalizedduplications of ATPalpha, with convergent tissue-specific expression patterns.[25][27]

Nucleic acids

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Convergence occurs at the level ofDNAand theamino acidsequences produced bytranslatingstructural genesintoproteins.Studies have found convergence in amino acid sequences in echolocating bats and the dolphin;[28]among marine mammals;[29]between giant and red pandas;[30]and between the thylacine and canids.[31]Convergence has also been detected in a type ofnon-coding DNA,cis-regulatory elements,such as in their rates of evolution; this could indicate eitherpositive selectionor relaxedpurifying selection.[32][33]

In animal morphology

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Dolphinsandichthyosaursconverged on many adaptations for fast swimming.

Bodyplans

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Swimming animals includingfishsuch asherrings,marine mammalssuch asdolphins,andichthyosaurs(of the Mesozoic) all converged on the same streamlined shape.[34][35]A similar shape and swimming adaptations are even present in molluscs, such asPhylliroe.[36]The fusiform bodyshape (a tube tapered at both ends) adopted by many aquatic animals is an adaptation to enable them totravel at high speedin a highdragenvironment.[37]Similar body shapes are found in theearless sealsand theeared seals:they still have four legs, but these are strongly modified for swimming.[38]

The marsupial fauna of Australia and the placental mammals of the Old World have several strikingly similar forms, developed in two clades, isolated from each other.[7]The body, and especially the skull shape, of thethylacine(Tasmanian tiger or Tasmanian wolf) converged with those ofCanidaesuch as the red fox,Vulpes vulpes.[39]

Echolocation

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As a sensory adaptation,echolocationhas evolved separately incetaceans(dolphins and whales) and bats, but from the same genetic mutations.[40]

Electric fishes

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TheGymnotiformesof South America and theMormyridaeof Africa independently evolvedpassive electroreception(around 119 and 110 million years ago, respectively). Around 20 million years after acquiring that ability, both groups evolved activeelectrogenesis,producing weak electric fields to help them detect prey.[41]

Eyes

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Main article:Eye evolution
The camera eyes ofvertebrates(left) andcephalopods(right) developed independently and are wired differently; for instance,optic nerve(3)fibres(2)reach the vertebrateretina(1)from the front, creating ablind spot(4).[42]

One of the best-known examples of convergent evolution is the camera eye ofcephalopods(such as squid and octopus),vertebrates(including mammals) andcnidaria(such as jellyfish).[43]Their last common ancestor had at most a simple photoreceptive spot, but a range of processes led to theprogressive refinement of camera eyes—with one sharp difference: the cephalopod eye is "wired" in the opposite direction, with blood and nerve vessels entering from the back of the retina, rather than the front as in vertebrates. As a result, vertebrates have ablind spot.[7]

Flight

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Further information:Flying and gliding animals § Evolution and ecology of aerial locomotion
Vertebrate wings are partlyhomologous(from forelimbs), but analogous as organs of flight in (1)pterosaurs,(2)bats,(3)birds,evolved separately.

Birdsandbatshavehomologouslimbs because they are both ultimately derived from terrestrialtetrapods,but their flight mechanisms are only analogous, so their wings are examples of functional convergence. The two groups have independently evolved their own means of powered flight. Their wings differ substantially in construction. The bat wing is a membrane stretched across four extremely elongated fingers and the legs. The airfoil of the bird wing is made offeathers,strongly attached to the forearm (the ulna) and the highly fused bones of the wrist and hand (thecarpometacarpus), with only tiny remnants of two fingers remaining, each anchoring a single feather. So, while the wings of bats and birds are functionally convergent, they are not anatomically convergent.[3][44]Birds and bats also share a high concentration ofcerebrosidesin the skin of their wings. This improves skin flexibility, a trait useful for flying animals; other mammals have a far lower concentration.[45]The extinctpterosaursindependently evolved wings from their fore- and hindlimbs, whileinsectshavewingsthat evolved separately from different organs.[46]

Flying squirrelsandsugar glidersare much alike in their body plans, with gliding wings stretched between their limbs, but flying squirrels are placental mammals while sugar gliders are marsupials, widely separated within the mammal lineage from the placentals.[47]

Hummingbird hawk-mothsandhummingbirdshave evolved similar flight and feeding patterns.[48]

Insect mouthparts

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Insect mouthparts show many examples of convergent evolution. The mouthparts of different insect groups consist of a set ofhomologousorgans, specialised for the dietary intake of that insect group. Convergent evolution of many groups of insects led from original biting-chewing mouthparts to different, more specialised, derived function types. These include, for example, theproboscisof flower-visiting insects such asbeesandflower beetles,[49][50][51]or the biting-sucking mouthparts of blood-sucking insects such asfleasandmosquitos.

Opposable thumbs

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Opposable thumbsallowing the grasping of objects are most often associated withprimates,like humans and other apes, monkeys, and lemurs. Opposable thumbs also evolved ingiant pandas,but these are completely different in structure, having six fingers including the thumb, which develops from a wrist bone entirely separately from other fingers.[52]

Primates

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Further information:Human skin color § Genetics of skin color variation
Despite the similar lightening ofskin colourafter movingout of Africa,different genes were involved in European (left) and East Asian (right) lineages.

Convergent evolution in humans includes blue eye colour and light skin colour.[53]When humans migratedout of Africa,they moved to more northern latitudes with less intense sunlight.[53]It was beneficial to them to reduce theirskin pigmentation.[53]It appears certain that there was some lightening of skin colourbeforeEuropean and East Asian lineages diverged, as there are some skin-lightening genetic differences that are common to both groups.[53]However, after the lineages diverged and became genetically isolated, the skin of both groups lightened more, and that additional lightening was due todifferentgenetic changes.[53]

Humans Lemurs
Despite the similarity of appearance, the genetic basis of blue eyes is different in humans andlemurs.

Lemursandhumansare both primates. Ancestral primates had brown eyes, as most primates do today. The genetic basis of blue eyes in humans has been studied in detail and much is known about it. It is not the case that onegene locusis responsible, say with brown dominant to blueeye colour.However, a single locus is responsible for about 80% of the variation. In lemurs, the differences between blue and brown eyes are not completely known, but the same gene locus is not involved.[54]

In plants

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Inmyrmecochory,seeds such as those ofChelidonium majushave a hard coating and an attached oil body, anelaiosome,for dispersal by ants.

The annual life-cycle

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While most plant species areperennial,about 6% follow anannuallife cycle, living for only one growing season.[55]The annual life cycle independently emerged in over 120 plant families of angiosperms.[56][57]The prevalence of annual species increases under hot-dry summer conditions in the four species-rich families of annuals (Asteraceae,Brassicaceae,Fabaceae,andPoaceae), indicating that the annual life cycle is adaptive.[55][58]

Carbon fixation

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C4photosynthesis,one of the three major carbon-fixing biochemical processes, hasarisen independently up to 40 times.[59][60]About 7,600 plant species ofangiospermsuse C4carbon fixation, with manymonocotsincluding 46% of grasses such asmaizeandsugar cane,[61][62]anddicotsincluding several species in theChenopodiaceaeand theAmaranthaceae.[63][64]

Fruits

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Fruitswith a wide variety of structural origins have converged to become edible.Applesarepomeswith fivecarpels;their accessory tissues form the apple's core, surrounded by structures from outside the botanical fruit, thereceptacleorhypanthium.Other edible fruits include other plant tissues;[65]the fleshy part of atomatois the walls of thepericarp.[66]This implies convergent evolution under selective pressure, in this case the competition forseed dispersalby animals through consumption of fleshy fruits.[67]

Seed dispersal by ants (myrmecochory) has evolved independently more than 100 times, and is present in more than 11,000 plant species. It is one of the most dramatic examples of convergent evolution in biology.[68]

Carnivory

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Molecular convergence incarnivorous plants

Carnivoryhas evolved multiple times independently in plants in widely separated groups. In three species studied,Cephalotus follicularis,Nepenthes alataandSarracenia purpurea,there has been convergence at the molecular level. Carnivorous plants secreteenzymesinto the digestive fluid they produce. By studyingphosphatase,glycoside hydrolase,glucanase,RNAseandchitinaseenzymesas well as apathogenesis-related proteinand athaumatin-related protein, the authors found many convergentamino acidsubstitutions. These changes were not at the enzymes' catalytic sites, but rather on the exposed surfaces of the proteins, where they might interact with other components of the cell or the digestive fluid. The authors also found thathomologous genesin the non-carnivorous plantArabidopsis thalianatend to have their expression increased when the plant is stressed, leading the authors to suggest that stress-responsive proteins have often been co-opted[c]in the repeated evolution of carnivory.[69]

Methods of inference

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Angiospermphylogeny of orders based on classification by the Angiosperm Phylogeny Group. The figure shows the number of inferred independent origins of C3-C4photosynthesis andC4photosynthesisin parentheses.

Phylogenetic reconstruction andancestral state reconstructionproceed by assuming that evolution has occurred without convergence. Convergent patterns may, however, appear at higher levels in a phylogenetic reconstruction, and are sometimes explicitly sought by investigators. The methods applied to infer convergent evolution depend on whether pattern-based or process-based convergence is expected. Pattern-based convergence is the broader term, for when two or more lineages independently evolve patterns of similar traits. Process-based convergence is when the convergence is due to similar forces ofnatural selection.[70]

Pattern-based measures

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Earlier methods for measuring convergence incorporate ratios of phenotypic andphylogeneticdistance by simulating evolution with aBrownian motionmodel of trait evolution along a phylogeny.[71][72]More recent methods also quantify the strength of convergence.[73]One drawback to keep in mind is that these methods can confuse long-term stasis with convergence due to phenotypic similarities. Stasis occurs when there is little evolutionary change among taxa.[70]

Distance-based measures assess the degree of similarity between lineages over time. Frequency-based measures assess the number of lineages that have evolved in a particular trait space.[70]

Process-based measures

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Methods to infer process-based convergence fit models of selection to a phylogeny and continuous trait data to determine whether the same selective forces have acted upon lineages. This uses theOrnstein–Uhlenbeck processto test different scenarios of selection. Other methods rely on ana priorispecification of where shifts in selection have occurred.[74]

See also

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  • Incomplete lineage sorting– Characteristic of phylogenetic analysis: the presence of multiple alleles in ancestral populations might lead to the impression that convergent evolution has occurred.
  • Carcinisation– Evolution of crustaceans into crab-like forms
  • Morphology (biology)– Study of external forms and structures of organisms
  • Iterative evolution– The repeated evolution of a specific trait or body plan from the same ancestral lineage at different points in time.
  • Elvis taxon– Misidentification of later taxon superficially resembling earlier extinct taxon
  • Breeding back– A form of selective breeding to recreate the traits of an extinct species, but the genome will differ from the original species.
  • Orthogenesis(contrastable with convergent evolution; involves teleology)
  • Contingency (evolutionary biology)– effect of evolutionary history on outcomes

Notes

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  1. ^However,evolutionary developmental biologyhas identifieddeep homologybetween insect and mammal body plans, to the surprise of many biologists.
  2. ^However, all organisms share a common ancestor more or less recently, so the question of how far back to look in evolutionary time and how similar the ancestors need to be for one to consider parallel evolution to have taken place is not entirely resolved within evolutionary biology.
  3. ^The prior existence of suitable structures has been calledpre-adaptationorexaptation.

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Further reading

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  • Losos, Jonathan B. (2017).Improbable Destinies: Fate, Chance, and the Future of Evolution.Riverhead Books.ISBN978-0399184925.
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