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Convergent evolution

January 31, 2023

Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last common ancestor of those groups. The cladistic term for the same phenomenon is homoplasy. The recurrent evolution of flight is a classic example, as flying insects, birds, pterosaurs, and bats have independently evolved the useful capacity of flight. Functionally similar features that have arisen through convergent evolution are analogous, whereas homologous structures or traits have a common origin but can have dissimilar functions. Bird, bat, and pterosaur wings are analogous structures, but their forelimbs are homologous, sharing an ancestral state despite serving different functions.

Two succulent plant genera, Euphorbia and Astrophytum, are only distantly related, but the species within each have converged on a similar body form.

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

Many instances of convergent evolution are known in plants, including the repeated development of C4 photosynthesis, seed dispersal by fleshy fruits adapted to be eaten by animals, and carnivory.

Overview

 
Homology and 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 different phylogeny.[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 similar ecological niches (that is, a distinctive way of life) similar problems can lead to similar solutions.[1][2][3] The British anatomist Richard Owen was the first to identify the fundamental difference between analogies and homologies.[4]

In biochemistry, physical and chemical constraints on mechanisms have caused some active site arrangements such as the catalytic triad to evolve independently in separate enzyme superfamilies.[5]

In his 1989 book Wonderful Life, Stephen Jay Gould argued 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 Morris disputes 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 least primates, corvids, and cetaceans.[7]

Distinctions

Cladistics

Main article: Cladistics

In cladistics, a homoplasy is a trait shared by two or more taxa for any reason other than that they share a common ancestry. Taxa which do share ancestry are part of the same clade; cladistics seeks to arrange them according to their degree of relatedness to describe their phylogeny. 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

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 an atavism. From a mathematical standpoint, an unused gene (selectively neutral) has a steadily decreasing probability of 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

 
Evolution at an amino acid position. 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 undergo divergent, 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] while others maintain that despite some overlap, there are still important distinctions between the two.[15][16]

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 by Richard Dawkins in The Blind Watchmaker as 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.[17]

At molecular level

 
Evolutionary convergence of serine and cysteine protease towards the same catalytic triads organisation of acid-base-nucleophile in different protease superfamilies. Shown are the triads of subtilisin, prolyl oligopeptidase, TEV protease, and papain.

Proteins

Protease active sites

Main article: Catalytic triad

The enzymology of proteases provides 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][18]

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

Threonine proteases use the amino acid threonine as their catalytic nucleophile. Unlike cysteine and serine, threonine is a secondary 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 such steric clashes. Several evolutionarily independent enzyme superfamilies with different protein folds use the N-terminal residue as a nucleophile. This commonality of active site but difference of protein fold indicates that the active site evolved convergently in those families.[5][19]

Cone snail and fish insulin, fish-like bacterial copper/zinc superoxide dismutase

Conus geographus produces a distinct form of insulin that is more similar to fish insulin protein sequences than to insulin from more closely related molluscs, suggesting convergent evolution.[20] Although convergent evolution is not impossible in this example, the possibility of horizontal gene transfer cannot be ignored, and it provides the only reasonable explanation of the fish-like copper/zinc superoxide dismutase of Photobacterium leiognathi.[21]

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

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 of Na+,K+-ATPase (ATPalpha). Variation in ATPalpha has been surveyed in various CTS-adapted species spanning six insect orders.[22][23][24] 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.[24] 30 of these substitutions (40%) occur at just two sites in the protein (positions 111 and 122). CTS-adapted species have also recurrently evolved neo-functionalized duplications of ATPalpha, with convergent tissue-specific expression patterns.[22][24]

Nucleic acids

Convergence occurs at the level of DNA and the amino acid sequences produced by translating structural genes into proteins. Studies have found convergence in amino acid sequences in echolocating bats and the dolphin;[25] among marine mammals;[26] between giant and red pandas;[27] and between the thylacine and canids.[28] Convergence has also been detected in a type of non-coding DNA, cis-regulatory elements, such as in their rates of evolution; this could indicate either positive selection or relaxed purifying selection.[29][30]

In animal morphology

 
Dolphins and ichthyosaurs converged on many adaptations for fast swimming.

Bodyplans

Swimming animals including fish such as herrings, marine mammals such as dolphins, and ichthyosaurs (of the Mesozoic) all converged on the same streamlined shape.[31][32] A similar shape and swimming adaptations are even present in molluscs, such as Phylliroe.[33] The fusiform bodyshape (a tube tapered at both ends) adopted by many aquatic animals is an adaptation to enable them to travel at high speed in a high drag environment.[34] Similar body shapes are found in the earless seals and the eared seals: they still have four legs, but these are strongly modified for swimming.[35]

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 the thylacine (Tasmanian tiger or Tasmanian wolf) converged with those of Canidae such as the red fox, Vulpes vulpes.[36]

Echolocation

As a sensory adaptation, echolocation has evolved separately in cetaceans (dolphins and whales) and bats, but from the same genetic mutations.[37]

Electric fishes

The Gymnotiformes of South America and the Mormyridae of Africa independently evolved passive electroreception (around 119 and 110 million years ago, respectively). Around 20 million years after acquiring that ability, both groups evolved active electrogenesis, producing weak electric fields to help them detect prey.[38]

Eyes

 
The camera eyes of vertebrates (left) and cephalopods (right) developed independently and are wired differently; for instance, optic nerve (3) fibres (2) reach the vertebrate retina (1) from the front, creating a blind spot (4).[39]
Main article: Eye evolution

One of the best-known examples of convergent evolution is the camera eye of cephalopods (such as squid and octopus), vertebrates (including mammals) and cnidaria (such as jellyfish).[40] Their last common ancestor had at most a simple photoreceptive spot, but a range of processes led to the progressive 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, cephalopods lack a blind spot.[7]

Flight

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

Birds and bats have homologous limbs because they are both ultimately derived from terrestrial tetrapods, 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 of feathers, strongly attached to the forearm (the ulna) and the highly fused bones of the wrist and hand (the carpometacarpus), 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][41] Birds and bats also share a high concentration of cerebrosides in the skin of their wings. This improves skin flexibility, a trait useful for flying animals; other mammals have a far lower concentration.[42] The extinct pterosaurs independently evolved wings from their fore- and hindlimbs, while insects have wings that evolved separately from different organs.[43]

Flying squirrels and sugar gliders are 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.[44]

Hummingbird hawk-moths and hummingbirds have evolved similar flight and feeding patterns.[45]

Insect mouthparts

Insect mouthparts show many examples of convergent evolution. The mouthparts of different insect groups consist of a set of homologous organs, 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, the proboscis of flower-visiting insects such as bees and flower beetles,[46][47][48] or the biting-sucking mouthparts of blood-sucking insects such as fleas and mosquitos.

Opposable thumbs

Opposable thumbs allowing the grasping of objects are most often associated with primates, like humans, monkeys, apes, and lemurs. Opposable thumbs also evolved in giant 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.[49]

Primates

Further information: Human skin color § Genetics of skin color variation
 
 
 
  Despite the similar lightening of skin colour after moving out 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.[50] When humans migrated out of Africa, they moved to more northern latitudes with less intense sunlight.[50] It was beneficial to them to reduce their skin pigmentation.[50] It appears certain that there was some lightening of skin colour before European and East Asian lineages diverged, as there are some skin-lightening genetic differences that are common to both groups.[50] However, after the lineages diverged and became genetically isolated, the skin of both groups lightened more, and that additional lightening was due to different genetic changes.[50]

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

Lemurs and humans are 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 one gene locus is responsible, say with brown dominant to blue eye 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.[51]

In plants

 
In myrmecochory, seeds such as those of Chelidonium majus have a hard coating and an attached oil body, an elaiosome, for dispersal by ants.

Carbon fixation

While convergent evolution is often illustrated with animal examples, it has often occurred in plant evolution. For instance, C4 photosynthesis, one of the three major carbon-fixing biochemical processes, has arisen independently up to 40 times.[52][53] About 7,600 plant species of angiosperms use C4 carbon fixation, with many monocots including 46% of grasses such as maize and sugar cane,[54][55] and dicots including several species in the Chenopodiaceae and the Amaranthaceae.[56][57]

Fruits

A good example of convergence in plants is the evolution of edible fruits such as apples. These pomes incorporate (five) carpels and their accessory tissues forming the apple's core, surrounded by structures from outside the botanical fruit, the receptacle or hypanthium. Other edible fruits include other plant tissues;[58] for example, the fleshy part of a tomato is the walls of the pericarp.[59] This implies convergent evolution under selective pressure, in this case the competition for seed dispersal by animals through consumption of fleshy fruits.[60]

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.[61]

Carnivory

 
Molecular convergence in carnivorous plants

Carnivory has evolved multiple times independently in plants in widely separated groups. In three species studied, Cephalotus follicularis, Nepenthes alata and Sarracenia purpurea, there has been convergence at the molecular level. Carnivorous plants secrete enzymes into the digestive fluid they produce. By studying phosphatase, glycoside hydrolase, glucanase, RNAse and chitinase enzymes as well as a pathogenesis-related protein and a thaumatin-related protein, the authors found many convergent amino acid substitutions. 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 that homologous genes in the non-carnivorous plant Arabidopsis thaliana tend 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.[62]

Methods of inference

 
Angiosperm phylogeny of orders based on classification by the Angiosperm Phylogeny Group. The figure shows the number of inferred independent origins of C3-C4 photosynthesis and C4 photosynthesis in parentheses.

Phylogenetic reconstruction and ancestral state reconstruction proceed 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 of natural selection.[63]

Pattern-based measures

Earlier methods for measuring convergence incorporate ratios of phenotypic and phylogenetic distance by simulating evolution with a Brownian motion model of trait evolution along a phylogeny.[64][65] More recent methods also quantify the strength of convergence.[66] 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.[63]

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.[63]

Process-based measures

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 the Ornstein–Uhlenbeck (OU) process to test different scenarios of selection. Other methods rely on an a priori specification of where shifts in selection have occurred.[67]

See also

  • 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

Notes

  1. ^ However, evolutionary developmental biology has identified deep homology between 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 called pre-adaptation or exaptation.

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Further reading

 
Wikimedia Commons has media related to Convergent evolution.
  • Jonathan B. Losos (2017). Improbable Destinies: Fate, Chance, and the Future of Evolution. Riverhead Books. ISBN 978-0399184925.

January 31, 2023
convergent, evolution, independent, evolution, similar, features, species, different, periods, epochs, time, creates, analogous, structures, that, have, similar, form, function, were, present, last, common, ancestor, those, groups, cladistic, term, same, pheno. Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time Convergent evolution creates analogous structures that have similar form or function but were not present in the last common ancestor of those groups The cladistic term for the same phenomenon is homoplasy The recurrent evolution of flight is a classic example as flying insects birds pterosaurs and bats have independently evolved the useful capacity of flight Functionally similar features that have arisen through convergent evolution are analogous whereas homologous structures or traits have a common origin but can have dissimilar functions Bird bat and pterosaur wings are analogous structures but their forelimbs are homologous sharing an ancestral state despite serving different functions Two succulent plant genera Euphorbia and Astrophytum are only distantly related but the species within each have converged on a similar body form The opposite of convergence is divergent evolution where related species evolve different traits Convergent evolution is similar to parallel evolution which occurs when two independent species evolve in the same direction and thus independently acquire similar characteristics for instance gliding frogs have evolved in parallel from multiple types of tree frog Many instances of convergent evolution are known in plants including the repeated development of C4 photosynthesis seed dispersal by fleshy fruits adapted to be eaten by animals and carnivory Contents 1 Overview 2 Distinctions 2 1 Cladistics 2 2 Atavism 2 3 Parallel vs convergent evolution 3 At molecular level 3 1 Proteins 3 1 1 Protease active sites 3 1 2 Cone snail and fish insulin fish like bacterial copper zinc superoxide dismutase 3 1 3 Na K ATPase and Insect resistance to cardiotonic steroids 3 2 Nucleic acids 4 In animal morphology 4 1 Bodyplans 4 2 Echolocation 4 3 Electric fishes 4 4 Eyes 4 5 Flight 4 6 Insect mouthparts 4 7 Opposable thumbs 4 8 Primates 5 In plants 5 1 Carbon fixation 5 2 Fruits 5 3 Carnivory 6 Methods of inference 6 1 Pattern based measures 6 2 Process based measures 7 See also 8 Notes 9 References 10 Further readingOverview Edit Homology and 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 different phylogeny 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 similar ecological niches that is a distinctive way of life similar problems can lead to similar solutions 1 2 3 The British anatomist Richard Owen was the first to identify the fundamental difference between analogies and homologies 4 In biochemistry physical and chemical constraints on mechanisms have caused some active site arrangements such as the catalytic triad to evolve independently in separate enzyme superfamilies 5 In his 1989 book Wonderful Life Stephen Jay Gould argued 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 Morris disputes 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 least primates corvids and cetaceans 7 Distinctions EditCladistics Edit Main article Cladistics In cladistics a homoplasy is a trait shared by two or more taxa for any reason other than that they share a common ancestry Taxa which do share ancestry are part of the same clade cladistics seeks to arrange them according to their degree of relatedness to describe their phylogeny 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 Edit 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 an atavism From a mathematical standpoint an unused gene selectively neutral has a steadily decreasing probability of 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 Edit Evolution at an amino acid position 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 undergo divergent 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 while others maintain that despite some overlap there are still important distinctions between the two 15 16 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 by Richard Dawkins in The Blind Watchmaker as 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 17 At molecular level Edit Evolutionary convergence of serine and cysteine protease towards the same catalytic triads organisation of acid base nucleophile in different protease superfamilies Shown are the triads of subtilisin prolyl oligopeptidase TEV protease and papain Proteins Edit Protease active sites Edit Main article Catalytic triad The enzymology of proteases provides 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 18 Serine and cysteine proteases use different amino acid functional groups alcohol or thiol as a nucleophile In order to activate that nucleophile they orient an acidic and a basic residue in a catalytic triad The chemical and physical constraints on enzyme catalysis have caused identical triad arrangements to evolve independently more than 20 times in different enzyme superfamilies 5 Threonine proteases use the amino acid threonine as their catalytic nucleophile Unlike cysteine and serine threonine is a secondary 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 such steric clashes Several evolutionarily independent enzyme superfamilies with different protein folds use the N terminal residue as a nucleophile This commonality of active site but difference of protein fold indicates that the active site evolved convergently in those families 5 19 Cone snail and fish insulin fish like bacterial copper zinc superoxide dismutase Edit Conus geographus produces a distinct form of insulin that is more similar to fish insulin protein sequences than to insulin from more closely related molluscs suggesting convergent evolution 20 Although convergent evolution is not impossible in this example the possibility of horizontal gene transfer cannot be ignored and it provides the only reasonable explanation of the fish like copper zinc superoxide dismutase of Photobacterium leiognathi 21 Na K ATPase and Insect resistance to cardiotonic steroids Edit 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 a subunit of Na K ATPase ATPalpha Variation in ATPalpha has been surveyed in various CTS adapted species spanning six insect orders 22 23 24 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 24 30 of these substitutions 40 occur at just two sites in the protein positions 111 and 122 CTS adapted species have also recurrently evolved neo functionalized duplications of ATPalpha with convergent tissue specific expression patterns 22 24 Nucleic acids Edit Convergence occurs at the level of DNA and the amino acid sequences produced by translating structural genes into proteins Studies have found convergence in amino acid sequences in echolocating bats and the dolphin 25 among marine mammals 26 between giant and red pandas 27 and between the thylacine and canids 28 Convergence has also been detected in a type of non coding DNA cis regulatory elements such as in their rates of evolution this could indicate either positive selection or relaxed purifying selection 29 30 In animal morphology Edit Dolphins and ichthyosaurs converged on many adaptations for fast swimming Bodyplans Edit Swimming animals including fish such as herrings marine mammals such as dolphins and ichthyosaurs of the Mesozoic all converged on the same streamlined shape 31 32 A similar shape and swimming adaptations are even present in molluscs such as Phylliroe 33 The fusiform bodyshape a tube tapered at both ends adopted by many aquatic animals is an adaptation to enable them to travel at high speed in a high drag environment 34 Similar body shapes are found in the earless seals and the eared seals they still have four legs but these are strongly modified for swimming 35 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 the thylacine Tasmanian tiger or Tasmanian wolf converged with those of Canidae such as the red fox Vulpes vulpes 36 Convergence of marsupial and placental mammals Red fox skeleton Skulls of thylacine left timber wolf right Thylacine skeletonEcholocation Edit As a sensory adaptation echolocation has evolved separately in cetaceans dolphins and whales and bats but from the same genetic mutations 37 Electric fishes Edit The Gymnotiformes of South America and the Mormyridae of Africa independently evolved passive electroreception around 119 and 110 million years ago respectively Around 20 million years after acquiring that ability both groups evolved active electrogenesis producing weak electric fields to help them detect prey 38 Eyes Edit The camera eyes of vertebrates left and cephalopods right developed independently and are wired differently for instance optic nerve 3 fibres 2 reach the vertebrate retina 1 from the front creating a blind spot 4 39 Main article Eye evolution One of the best known examples of convergent evolution is the camera eye of cephalopods such as squid and octopus vertebrates including mammals and cnidaria such as jellyfish 40 Their last common ancestor had at most a simple photoreceptive spot but a range of processes led to the progressive 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 cephalopods lack a blind spot 7 Flight Edit Vertebrate wings are partly homologous from forelimbs but analogous as organs of flight in 1 pterosaurs 2 bats 3 birds evolved separately Further information Flying and gliding animals Evolution and ecology of aerial locomotion Birds and bats have homologous limbs because they are both ultimately derived from terrestrial tetrapods 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 of feathers strongly attached to the forearm the ulna and the highly fused bones of the wrist and hand the carpometacarpus 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 41 Birds and bats also share a high concentration of cerebrosides in the skin of their wings This improves skin flexibility a trait useful for flying animals other mammals have a far lower concentration 42 The extinct pterosaurs independently evolved wings from their fore and hindlimbs while insects have wings that evolved separately from different organs 43 Flying squirrels and sugar gliders are 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 44 Hummingbird hawk moths and hummingbirds have evolved similar flight and feeding patterns 45 Insect mouthparts Edit Insect mouthparts show many examples of convergent evolution The mouthparts of different insect groups consist of a set of homologous organs 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 the proboscis of flower visiting insects such as bees and flower beetles 46 47 48 or the biting sucking mouthparts of blood sucking insects such as fleas and mosquitos Opposable thumbs Edit Opposable thumbs allowing the grasping of objects are most often associated with primates like humans monkeys apes and lemurs Opposable thumbs also evolved in giant 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 49 Primates Edit Further information Human skin color Genetics of skin color variation Despite the similar lightening of skin colour after moving out 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 50 When humans migrated out of Africa they moved to more northern latitudes with less intense sunlight 50 It was beneficial to them to reduce their skin pigmentation 50 It appears certain that there was some lightening of skin colour before European and East Asian lineages diverged as there are some skin lightening genetic differences that are common to both groups 50 However after the lineages diverged and became genetically isolated the skin of both groups lightened more and that additional lightening was due to different genetic changes 50 Humans Lemurs Despite the similarity of appearance the genetic basis of blue eyes is different in humans and lemurs Lemurs and humans are 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 one gene locus is responsible say with brown dominant to blue eye 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 51 In plants Edit In myrmecochory seeds such as those of Chelidonium majus have a hard coating and an attached oil body an elaiosome for dispersal by ants Carbon fixation Edit While convergent evolution is often illustrated with animal examples it has often occurred in plant evolution For instance C4 photosynthesis one of the three major carbon fixing biochemical processes has arisen independently up to 40 times 52 53 About 7 600 plant species of angiosperms use C4 carbon fixation with many monocots including 46 of grasses such as maize and sugar cane 54 55 and dicots including several species in the Chenopodiaceae and the Amaranthaceae 56 57 Fruits Edit A good example of convergence in plants is the evolution of edible fruits such as apples These pomes incorporate five carpels and their accessory tissues forming the apple s core surrounded by structures from outside the botanical fruit the receptacle or hypanthium Other edible fruits include other plant tissues 58 for example the fleshy part of a tomato is the walls of the pericarp 59 This implies convergent evolution under selective pressure in this case the competition for seed dispersal by animals through consumption of fleshy fruits 60 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 61 Carnivory Edit Molecular convergence in carnivorous plants Carnivory has evolved multiple times independently in plants in widely separated groups In three species studied Cephalotus follicularis Nepenthes alata and Sarracenia purpurea there has been convergence at the molecular level Carnivorous plants secrete enzymes into the digestive fluid they produce By studying phosphatase glycoside hydrolase glucanase RNAse and chitinase enzymes as well as a pathogenesis related protein and a thaumatin related protein the authors found many convergent amino acid substitutions 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 that homologous genes in the non carnivorous plant Arabidopsis thaliana tend 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 62 Methods of inference Edit Angiosperm phylogeny of orders based on classification by the Angiosperm Phylogeny Group The figure shows the number of inferred independent origins of C3 C4 photosynthesis and C4 photosynthesis in parentheses Phylogenetic reconstruction and ancestral state reconstruction proceed 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 of natural selection 63 Pattern based measures Edit Earlier methods for measuring convergence incorporate ratios of phenotypic and phylogenetic distance by simulating evolution with a Brownian motion model of trait evolution along a phylogeny 64 65 More recent methods also quantify the strength of convergence 66 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 63 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 63 Process based measures Edit 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 the Ornstein Uhlenbeck OU process to test different scenarios of selection Other methods rely on an a priori specification of where shifts in selection have occurred 67 See also EditIncomplete 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 formsNotes Edit However evolutionary 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