Topic: Analogous structure
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.
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
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 [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.
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;
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.
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
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
Proteins
Protease active sites
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.
Cone snail and fish insulin
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.
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 6 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 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
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. 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 (Tasmanian tiger or Tasmanian wolf) converged
with those of Canidae such as the red fox, Vulpes vulpes.
Convergence of marsupial and placental mammals
Echolocation
As a sensory adaptation, echolocation has evolved separately in cetaceans (dolphins and
whales) and bats, but from the same genetic mutations.
Eyes
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. As a result, cephalopods
lack a blind spot.
Flight
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. 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.Hummingbird hawk-moths and hummingbirds have evolved similar flight and
feeding patterns.
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, 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.
Primates
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.
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 plants
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. 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.
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; 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 biology.
Carnivory
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
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 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 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.
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 non-crab-like crustaceans into crab-like forms
Notes
References
Further reading
Jonathan B. Losos (2017). Improbable Destinies: Fate, Chance, and the Future of Evolution.
Riverhead Books. ISBN 978-0399184925.
