Convergent evolution is the independent evolution of similar features
in species of different lineages.
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
The opposite of convergence is divergent evolution, where related
species evolve different traits.
Convergent evolution is similar to
but different from parallel evolution.
Parallel evolution occurs when
two independent but similar 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
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.
2.3 Parallel vs. convergent evolution
3 At molecular level
3.1 Protease active sites
3.2 Nucleic acids
4 In animal morphology
4.6 Opposable thumbs
5 In plants
5.1 Carbon fixation
6 Methods of inference
6.1 Pattern-based measures
6.2 Process-based measures
7 See also
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. The British anatomist
Richard Owen was the first
to identify the fundamental difference between analogies and
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.
In his 1989 book Wonderful Life,
Stephen Jay Gould
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".
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.
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.
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.
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, 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 in
The Blind Watchmaker
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.
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.
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.
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.
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.
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; 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 or relaxed purifying selection.
In animal morphology
Dolphins and ichthyosaurs converged on many adaptations for fast
Swimming animals including fish such as herrings, marine mammals such
as dolphins, and ichthyosaurs (of the Mesozoic) all converged on the
same streamlined shape. 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
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 (Tasmanian wolf) converged with those of
as the red fox, Vulpes vulpes.
Convergence of marsupial and placental mammals
Red fox skeleton
Skulls of thylacine (left), timber wolf (right)
As a sensory adaptation, echolocation has evolved separately in
cetaceans (dolphins and whales) and bats, but from the same genetic
The camera eyes of vertebrates and cephalopods developed independently
and are wired differently; for instance, optic nerve fibres reach the
vertebrate retina from the front, creating a blind spot.
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). 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. This means that
cephalopods do not have a blind spot.
Vertebrate wings are partly homologous (from forelimbs), but analogous
as organs of flight in (1) pterosaurs, (2) bats, (3) birds, evolved
Further information: Flying and gliding animals §
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 powered flight, evolved independently. 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
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. The extinct pterosaurs independently
evolved wings from their fore- and hindlimbs, while insects have wings
that evolved separately from different organs.
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.
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,
or the biting-sucking mouthparts of blood-sucking insects such as
fleas and mosquitos.
Opposable thumbs allowing the grasping of objects are most often
associated with primates, like humans, monkeys, apes, and lemurs.
Opposable thumbs also evolved in 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.
Human skin color
Human skin color § Genetics of skin color
Despite the similar lightening of skin colour after moving out of
Africa, different genes were involved in European (left) and Chinese
Convergent evolution in humans includes blue eye colour and light skin
colour. When humans migrated out of Africa, they moved to more
northern latitudes with less intense sunlight. It was beneficial to
them to reduce their skin pigmentation. 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. 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.
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.
In myrmecochory, seeds such as those of
Chelidonium majus have a hard
coating and an attached oil body, an elaiosome, for dispersal by ants.
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. About
7,600 plant species of angiosperms use C4 carbon fixation, with many
monocots including 46% of grasses such as maize and sugar
cane, and dicots including several species in the
Chenopodiaceae and the Amaranthaceae.
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; for
example, the fleshy part of a tomato is the walls of the pericarp.
This implies convergent evolution under selective pressure, in this
case the competition for seed dispersal by animals through consumption
of fleshy fruits.
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
Molecular convergence in carnivorous plants
Carnivory has evolved multiple times independently in plants in widely
separated groups. In three species studied,
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
Methods of inference
Angiosperm phylogeny of orders based on classification by the
Phylogeny Group. The figure shows the number of inferred
independent origins of C3-
C4 photosynthesis and
C4 photosynthesis in
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.
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.
More recent methods also quantify the strength of convergence. 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.
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.
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.
Incomplete lineage sorting: the presence of multiple alleles in
ancestral populations might lead to the impression that convergent
evolution has occurred.
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Patterns of evolution
Coloration evidence for natural selection
Paradox of the plankton