Convergent evolution is the independent

evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...

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 Cladistics ( ; from Ancient Greek 'branch') is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is ...

term for the same phenomenon is homoplasy. The recurrent evolution of flight is a classic example, as flying

insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...

bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...

pterosaurs Pterosaurs are an extinct clade of flying reptiles in the Order (biology), order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosau ...

, 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. The opposite of convergence is

divergent evolution Divergent evolution or divergent selection is the accumulation of differences between closely related populations within a species, sometimes leading to speciation. Divergent evolution is typically exhibited when two populations become separate ...

, 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 A tree frog (or treefrog) is any species of frog that spends a major portion of its lifespan in trees, known as an arboreal state. Several lineages of frogs among the Neobatrachia suborder have given rise to treefrogs, although they are not clos ...

. Many instances of convergent evolution are known in

plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they are predominantly Photosynthesis, photosynthetic. This means that they obtain their energy from sunlight, using chloroplasts derived from endosymbiosis with c ...

s, including the repeated development of C₄ photosynthesis,

seed dispersal In spermatophyte plants, seed dispersal is the movement, spread or transport of seeds away from the parent plant. Plants have limited mobility and rely upon a variety of dispersal vectors to transport their seeds, including both abiotic vectors, ...

by fleshy

fruit In botany, a fruit is the seed-bearing structure in flowering plants (angiosperms) that is formed from the ovary after flowering. Fruits are the means by which angiosperms disseminate their seeds. Edible fruits in particular have long propaga ...

s adapted to be eaten by animals, and

carnivory A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose nutrition and energy requirements are met by consumption of animal tissues (mainly mu ...

Overview

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 niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of Resource (biology), resources an ...

s (that is, a distinctive way of life) similar problems can lead to similar solutions. The British anatomist

Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomy, comparative anatomist and paleontology, palaeontologist. Owen is generally considered to have been an outstanding naturalist with a remarkabl ...

was the first to identify the fundamental difference between analogies and homologies. In biochemistry, physical and chemical constraints on mechanisms have caused some

active site In biology and biochemistry, the active site is the region of an enzyme where substrate molecules bind and undergo a chemical reaction. The active site consists of amino acid residues that form temporary bonds with the substrate, the ''binding s ...

arrangements such as the

catalytic triad A catalytic triad is a set of three coordinated amino acid residues that can be found in the active site of some enzymes. Catalytic triads are most commonly found in hydrolase and transferase enzymes (e.g. proteases, amidases, esterases, aminoac ...

to evolve independently in separate enzyme superfamilies. In his 1989 book '' Wonderful Life'',

Stephen Jay Gould Stephen Jay Gould ( ; September 10, 1941 – May 20, 2002) was an American Paleontology, paleontologist, Evolutionary biology, evolutionary biologist, and History of science, historian of science. He was one of the most influential and widely re ...

argued that if one could "rewind the tape of life ndthe same conditions were encountered again, evolution could take a very different course." 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 Intelligence has been defined in many ways: the capacity for abstraction, logic, understanding, self-awareness, learning, emotional knowledge, reasoning, planning, creativity, critical thinking, and problem-solving. It can be described as t ...

, a trait presently identified with at least

primates Primates is an order of mammals, which is further divided into the strepsirrhines, which include lemurs, galagos, and lorisids; and the haplorhines, which include tarsiers and simians ( monkeys and apes). Primates arose 74–63 ...

, corvids, and

cetaceans Cetacea (; , ) is an infraorder of aquatic mammals belonging to the order Artiodactyla that includes whales, dolphins and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively c ...

Distinctions

Cladistics

In cladistics, a homoplasy is a trait shared by two or more

taxa In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...

for any reason other than that they share a common ancestry. Taxa which do share ancestry are part of the same

clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...

; cladistics seeks to arrange them according to their degree of relatedness to describe their

phylogeny A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or Taxon, taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, M ...

. Homoplastic traits caused by convergence are therefore, from the point of view of cladistics, confounding factors which could lead to an incorrect analysis.

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 Probability is a branch of mathematics and statistics concerning events and numerical descriptions of how likely they are to occur. The probability of an event is a number between 0 and 1; the larger the probability, the more likely an e ...

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.

Parallel vs. convergent evolution

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. Some scientists have argued that there is a continuum between parallel and convergent evolution, while others maintain that despite some overlap, there are still important distinctions between the two. 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 Richard Dawkins (born 26 March 1941) is a British evolutionary biology, evolutionary biologist, zoologist, science communicator and author. He is an Oxford fellow, emeritus fellow of New College, Oxford, and was Simonyi Professor for the Publ ...

in ''

The Blind Watchmaker ''The Blind Watchmaker: Why the Evidence of Evolution Reveals a Universe without Design'' is a 1986 book by Richard Dawkins, in which the author presents an explanation of, and argument for, the theory of evolution by means of natural selecti ...

'' 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.

At molecular level

Proteins

Tertiary structures

Many proteins share analogous structural elements that arose independently across different genomes. There are several examples of convergent protein motifs sharing similar arrangements of structural elements. Whole protein structures too have arisen through convergent evolution.

Protease active sites

The

enzymology An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...

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. Serine and cysteine proteases use different amino acid functional groups (alcohol or thiol) as a

nucleophile In chemistry, a nucleophile is a chemical species that forms bonds by donating an electron pair. All molecules and ions with a free pair of electrons or at least one pi bond can act as nucleophiles. Because nucleophiles donate electrons, they are ...

. To activate that nucleophile, they orient an acidic and a basic residue in a

. The chemical and physical constraints on

enzyme catalysis Enzyme catalysis is the increase in the rate of a process by an "enzyme", a biological molecule. Most enzymes are proteins, and most such processes are chemical reactions. Within the enzyme, generally catalysis occurs at a localized site, calle ...

have caused identical triad arrangements to evolve independently more than 20 times in different enzyme superfamilies. Threonine proteases use the amino acid threonine as their catalytic

. 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

but difference of protein fold indicates that the active site evolved convergently in those families.

Cone snail and fish insulin

'' Conus geographus'' produces a distinct form of

insulin Insulin (, from Latin ''insula'', 'island') is a peptide hormone produced by beta cells of the pancreatic islets encoded in humans by the insulin (''INS)'' gene. It is the main Anabolism, anabolic hormone of the body. It regulates the metabol ...

that is more similar to fish insulin protein sequences than to insulin from more closely related molluscs, suggesting convergent evolution, though with the possibility of

horizontal gene transfer Horizontal gene transfer (HGT) or lateral gene transfer (LGT) is the movement of genetic material between organisms other than by the ("vertical") transmission of DNA from parent to offspring (reproduction). HGT is an important factor in the e ...

Ferrous iron uptake via protein transporters in land plants and chlorophytes

Distant homologues of the metal ion transporters ZIP in

land plants The embryophytes () are a clade of plants, also known as Embryophyta (Plantae ''sensu strictissimo'') () or land plants. They are the most familiar group of photoautotrophs that make up the vegetation on Earth's dry lands and wetlands. Embryophy ...

and chlorophytes have converged in structure, likely to take up Fe²⁺ efficiently. The IRT1 proteins from ''

Arabidopsis thaliana ''Arabidopsis thaliana'', the thale cress, mouse-ear cress or arabidopsis, is a small plant from the mustard family (Brassicaceae), native to Eurasia and Africa. Commonly found along the shoulders of roads and in disturbed land, it is generally ...

'' and

rice Rice is a cereal grain and in its Domestication, domesticated form is the staple food of over half of the world's population, particularly in Asia and Africa. Rice is the seed of the grass species ''Oryza sativa'' (Asian rice)—or, much l ...

have extremely different amino acid sequences from ''

Chlamydomonas ''Chlamydomonas'' ( ) is a genus of green algae consisting of about 150 species of unicellular organism, unicellular flagellates, found in stagnant water and on damp soil, in freshwater, seawater, and even in snow as "snow algae". ''Chlamydom ...

''s IRT1, but their three-dimensional structures are similar, suggesting convergent evolution.

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. 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. 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.

Nucleic acids

Convergence occurs at the level of

DNA Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...

and the

amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although over 500 amino acids exist in nature, by far the most important are the 22 α-amino acids incorporated into proteins. Only these 22 a ...

sequences produced by translating structural genes into

protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residue (biochemistry), residues. Proteins perform a vast array of functions within organisms, including Enzyme catalysis, catalysing metab ...

s. Studies have found convergence in amino acid sequences in echolocating bats and the dolphin; among marine mammals; between giant and red pandas; and between the thylacine and canids. Convergence has also been detected in a type of

non-coding DNA Non-coding DNA (ncDNA) sequences are components of an organism's DNA that do not encode protein sequences. Some non-coding DNA is transcribed into functional non-coding RNA molecules (e.g. transfer RNA, microRNA, piRNA, ribosomal RNA, and reg ...

cis-regulatory element ''Cis''-regulatory elements (CREs) or ''cis''-regulatory modules (CRMs) are regions of non-coding DNA which regulate the transcription of neighboring genes. CREs are vital components of genetic regulatory networks, which in turn control morpho ...

s, such as in their rates of evolution; this could indicate either positive selection or relaxed purifying selection.

In animal morphology

Bodyplans

Swimming animals including

fish A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...

such as

herring Herring are various species of forage fish, belonging to the Order (biology), order Clupeiformes. Herring often move in large Shoaling and schooling, schools around fishing banks and near the coast, found particularly in shallow, temperate wate ...

s, marine mammals such as

dolphins A dolphin is an aquatic mammal in the cetacean clade Odontoceti (toothed whale). Dolphins belong to the families Delphinidae (the oceanic dolphins), Platanistidae (the Indian river dolphins), Iniidae (the New World river dolphins), Pontopori ...

, and ichthyosaurs ( of the Mesozoic) all converged on the same streamlined shape. A similar shape and swimming adaptations are even present in molluscs, such as '' Phylliroe''. 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. 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. 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. The body, and especially the skull shape, of the

thylacine The thylacine (; binomial name ''Thylacinus cynocephalus''), also commonly known as the Tasmanian tiger or Tasmanian wolf, was a carnivorous marsupial that was native to the Mainland Australia, Australian mainland and the islands of Tasmani ...

(Tasmanian tiger or Tasmanian wolf) converged with those of

Canidae Canidae (; from Latin, ''canis'', "dog") is a family (biology), biological family of caniform carnivorans, constituting a clade. A member of this family is also called a canid (). The family includes three subfamily, subfamilies: the Caninae, a ...

such as the red fox, ''

Vulpes vulpes The red fox (''Vulpes vulpes'') is the largest of the true foxes and one of the most widely distributed members of the order Carnivora, being present across the entire Northern Hemisphere including most of North America, Europe and Asia, plus ...

''. marsupial Marsupials are a diverse group of mammals belonging to the infraclass Marsupialia. They are natively found in Australasia, Wallacea, and the Americas. One of marsupials' unique features is their reproductive strategy: the young are born in a r ...