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
* 1 Overview
* 2 Distinctions
* 2.3 Parallel vs. convergent evolution
* 3 At molecular level
* 3.1 Protease active sites
* 3.2 Nucleic acids
* 4 In animal morphology
* 4.1 Bodyplans
* 4.2 Echolocation
* 4.3 Eyes
* 4.4 Flight
* 4.6 Opposable thumbs
* 5 In plants
* 5.1 Carbon fixation
* 5.2 Fruits
* 6 Methods of inference
* 6.1 Pattern-based convergence measures
* 6.2 Process-based convergence measures
* 7 See also
* 8 Notes
* 9 References
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 . Further
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 homologies.
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 argued that if
one could "rewind the tape of life 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.
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. 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 _ 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 amino acids. In 2013 the
first genome-wide study of convergence was published. Comparisons of
the genomes of echolocating bats and the dolphin identified numerous
convergent amino acid substitutions in genes implicated in hearing and
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
Canidae such as
the red fox, _
Vulpes vulpes _.
* Convergence of marsupial and placental mammals
Skulls of thylacine (left), timber wolf (right)
Red fox skeleton
As a sensory adaptation, echolocation has evolved separately in
cetaceans (dolphins and whales) and bats, but from the same genetic
The camera eye of vertebrates and cephalopods developed
independently and are wired differently. Main article: Eye
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 separately. Further information: Flying and gliding
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 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
convergent. Similarly, the extinct pterosaur also shows an
independent evolution of vertebrate fore- and hindlimbs to wing. An
even more distantly related group, the 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
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 § Genetics of skin color
Despite the similar lightening of skin color 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 color . However, a single locus is responsible
for about 80% of the variation. In lemurs, the difference(s) 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
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.
The emergence of 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.
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 in the repeated
evolution of carnivory.
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 .
PATTERN-BASED CONVERGENCE 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. 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.
PROCESS-BASED CONVERGENCE 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.
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
Paradox of the plankton
Paradox of the plankton
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