Convergent evolution is the independent
evolution
Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
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
Recurrent evolution is the repeated evolution of a particular trait, character, or mutation. Most evolution is the result of drift, often interpreted as the random chance of some alleles being passed down to the next generation and others not. Recu ...
of flight is a classic example, as flying
insect
Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s,
bird
Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
s,
pterosaurs
Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 to ...
, and
bat
Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most bi ...
s 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, 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
plant
Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclud ...
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 that is formed from the ovary after flowering.
Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...
s adapted to be eaten by animals, and
carnivory.
Overview
![Analogous & Homologous Structures](https://upload.wikimedia.org/wikipedia/commons/0/08/Analogous_%26_Homologous_Structures.svg)
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 resources and competitors (for ...
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 anatomist and paleontologist. Owen is generally considered to have been an outstanding naturalist with a remarkable gift for interpreting fossils.
Owe ...
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 (binding site) a ...
arrangements such as the catalytic triad 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 paleontologist, evolutionary biologist, and historian of science. He was one of the most influential and widely read authors of popular science of his generation. Gould sp ...
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, a trait presently identified with at least primates
Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
, corvids
Corvidae is a cosmopolitan family of oscine passerine birds that contains the crows, ravens, rooks, jackdaws, jays, magpies, treepies, choughs, and nutcrackers. In colloquial English, they are known as the crow family or corvids. Currently, 133 ...
, and cetaceans
Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel them ...
.[
]
Distinctions
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
A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
; cladistics seeks to arrange them according to their degree of relatedness to describe their phylogeny
A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
. 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 the branch of mathematics concerning numerical descriptions of how likely an Event (probability theory), event is to occur, or how likely it is that a proposition is true. The probability of an event is a number between 0 and ...
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 biologist and author. He is an emeritus fellow of New College, Oxford and was Professor for Public Understanding of Science in the University of Oxford from 1995 to 2008. An ath ...
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
Protease active sites
The enzymology
Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products. A ...
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 ...
. 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
Enzyme catalysis is the increase in the rate of a process by a biological molecule, an "enzyme". 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 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 ...
. 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 clash
Steric effects arise from the spatial arrangement of atoms. When atoms come close together there is a rise in the energy of the molecule. Steric effects are nonbonding interactions that influence the shape ( conformation) and reactivity of ions ...
es.
Several evolutionarily independent enzyme superfamilies with different protein folds use the N-terminal residue as a nucleophile. This commonality of 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 (binding site) a ...
but difference of protein fold indicates that the active site evolved convergently in those families.
Cone snail and fish insulin, fish-like bacterial copper/zinc superoxide dismutase
''Conus geographus
''Conus geographus'', popularly called the geography cone or the geographer cone, is a species of predatory cone snail. It lives in reefs of the tropical Indo-Pacific, and hunts small fish. While all cone snails hunt and kill prey using venom, t ...
'' 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 ''INS'' gene. It is considered to be the main anabolic hormone of the body. It regulates the metabolism o ...
that is more similar to fish insulin protein sequences than to insulin from more closely related molluscs, suggesting convergent evolution. 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
Superoxide dismutase (SOD, ) is an enzyme that alternately catalyzes the dismutation (or partitioning) of the superoxide () radical into ordinary molecular oxygen (O2) and hydrogen peroxide (). Superoxide is produced as a by-product of oxygen me ...
of ''Photobacterium leiognathi
''Photobacterium leiognathi'' is a species of Gram-negative bacteria. The species is bioluminescent and a symbiont of ponyfish
Leiognathidae, the ponyfishes, slipmouths or slimys / slimies, are a small family of fishes in the order Perciformes ...
''.
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 and the amino acid
Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha am ...
sequences produced by translating structural gene
A structural gene is a gene that codes for any RNA or protein product other than a regulatory factor (i.e. regulatory protein). A term derived from the Lac operon, ''lac'' operon, structural genes are typically viewed as those containing sequences ...
s into protein
Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
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, cis-regulatory elements, such as in their rates of evolution; this could indicate either positive selection
In population genetics, directional selection, is a mode of negative natural selection in which an extreme phenotype is favored over other phenotypes, causing the allele frequency to shift over time in the direction of that phenotype. Under dir ...
or relaxed purifying selection.
In animal morphology
Bodyplans
Swimming animals including fish
Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...
such as herring
Herring are forage fish, mostly belonging to the family of Clupeidae.
Herring often move in large schools around fishing banks and near the coast, found particularly in shallow, temperate waters of the North Pacific and North Atlantic Oceans, i ...
s, marine mammals
Marine mammals are aquatic mammals that rely on the ocean and other marine ecosystems for their existence. They include animals such as seals, whales, manatees, sea otters and polar bears. They are an informal group, unified only by their relia ...
such as dolphins, and ichthyosaur
Ichthyosaurs (Ancient Greek for "fish lizard" – and ) are large extinct marine reptiles. Ichthyosaurs belong to the order known as Ichthyosauria or Ichthyopterygia ('fish flippers' – a designation introduced by Sir Richard Owen in 1842, altho ...
s ( of the Mesozoic) all converged on the same streamlined shape. A similar shape and swimming adaptations are even present in molluscs, such as ''Phylliroe
''Phylliroe'' is a genus of average sized (up to ), highly transparent pelagic nudibranchs, marine gastropod molluscs in the order Opisthobranchia, that consists of two known species. It is notable for being an open-ocean hunter that resembl ...
''. 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 seal
The earless seals, phocids or true seals are one of the three main groups of mammals within the seal lineage, Pinnipedia. All true seals are members of the family Phocidae (). They are sometimes called crawling seals to distinguish them from th ...
s 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 ( , or , also ) (''Thylacinus cynocephalus'') is an extinct carnivorous marsupial that was native to the Australian mainland and the islands of Tasmania and New Guinea. The last known live animal was captured in 1930 in Tasman ...
(Tasmanian tiger or Tasmanian wolf) converged with those of Canidae
Canidae (; from Latin, ''canis'', "dog") is a biological family of dog-like carnivorans, colloquially referred to as dogs, and constitutes a clade. A member of this family is also called a canid (). There are three subfamilies found within th ...
such as the red fox, '' Vulpes vulpes''.
marsupial
Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in a po ...
(dolphins and whales) and bats, but from the same genetic mutations.
(around 119 and 110 million years ago, respectively). Around 20 million years after acquiring that ability, both groups evolved active
, producing weak electric fields to help them detect prey.
(such as jellyfish).
Their last common ancestor had at most a simple photoreceptive spot, but a range of processes led to the
— 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
.