Lecture 3: Evolutionary Patterns

Diverse
Diverse adaptations
adaptations to
to aa dry
dry environment.
environment. The
The adaptations
adaptations ofof plants
plants in
in aa California
California desert
desert include
include
fleshy,
fleshy, leafless
leafless stems
stems that
that store
store water;
water; dense
dense spines
spines that
that reduce
reduce heat
heat load
load by
by reflecting
reflecting light;
light; small
small
leaves;
leaves; widely
widely spreading
spreading roots
roots that
that catch
catch rare
rare rainwater;
rainwater; and
and brief
brief seasonal
seasonal flowering.
flowering. Each
Each of of these
these
adaptations
adaptations hashas evolve
evolve independently
independently in in many
many distantly
distantly related
related plants.
plants.
 Evolutionary Patterns
Tracing the path of evolution to Homo sapiens from the
universal ancestor of all life
Studies of phylogeny allow
for the discovery of
patterns or common
themes of evolution.
Organismic diversity allows for few
generalized “laws” of biology
(similar to those of physics) but
generalizations can be made about
what kinds of evolutionary changes
have been prevalent and
developing such statements is one
of the main tasks for biological
science.
General patterns of change are the most
important phenomena for evolutionary
biology to explain.
Evolutionary Patterns
Many traditional
classifications have been
constructed to convey both
anagenesis and
cladogenesis (such as birds
being separately classified
into the Aves because of
adaptations for flight).
However such traditional
placements obscure close
relationships (such as birds to
crocodiles and dinosaurs
traditionally classed as Reptilia)
and only demonstrate great
divergences.
Remember, modern phylogenetic and
taxonomic studies seek to discover
Evolution has two major features: the branching of a lineage into two or
more descendant lines, called cladogenesis, and evolutionary change of monophyly.
various characteristics in each of the descendants called anagenesis.
Evolutionary
Patterns Even with true estimates of phylogeny,
certain aspects of taxonomy can be
difficult, for example we often find that
an (often) extinct stem group of species
can give rise to a later crown group
with distinctive derived characters
(which may [usually] or may not still be
extant).

Classifying the crown group to the exclusion of

a stem group creates paraphyly.
A cladogram of some carnivorous dinosaurs showing the
two main types of phylogenetic taxon definitions. (A)
Carnosauria is a stem-based taxon (Allosaurus and all
theropods closer to Allosaurus than
than birds).
birds). (B)
(B) Decisions whether members of a monophyletic
Allosauroidea is a node-based taxon (all descendants of
the most recent ancestor of Allosaurus and Sinraptor). group should be split into several taxa or
Thus Cryolophosaurus
Cryolophosaurus and Monolophosaurus are are both
both
carnosaurs but not allosaurids, while “lumped” into one taxon may be quite arbitrary.
Carcharodontosaurus, Acrocanthosaurus
Acrocanthosaurus and (by
definition) Allosaurus and Sinraptor
Sinraptor are carnosaurs and
allosaurids.
Evolutionary Patterns Some patterns of
evolutionary change can
be inferred from
systematics and one very
important one is that most
features of organisms
have been modified
(evolved) from pre-
existing features (they do
not arise from nothing.

The forelimb skeletons of some tetrapod vertebrates showing structural

Related organisms have
homologies. These vertebrate forelimbs are used for different functions, but
have the same sequence and arrangement of bones. In this illustration,
homologous characters
homologous bones are colored in the same way and are labeled on the
human arm.
which have been inherited
(and sometimes modified)
from an equivalent organ
in the common ancestor.
Evolutionary Patterns Homologous characters
generally have similar
genetic and
developmental
underpinnings despite
the often substantial
divergence they may
undergo among species.

Bear in mind that a

Embryos from different vertebrates show striking similarities (developmental
homologies) early in development. Note that the early embryos shown here all character may show
have pharyngeal pouches and a tail.
homology among
species (e.g., toes) but a
given character state
may not (e.g., number of
toes).
Evolutionary Patterns A character (or character
state) is defined as
homologous in two
species if derived from a
common ancestor, but
the diagnosis of
homologous characters
can be quite difficult.

For anatomical homology, the

most common criteria for their
diagnosis are correspondence
More structural homologies: orchid flowers are diverse in size and of position and
shape, but are comprised of elements that are similar in structure
and orientation. correspondence of structure
(the parts composing the
structure), but correspondence
of shape or function are NOT
useful.
Evolutionary Patterns
Advances in molecular biology have also
revealed non-anatomical similarities among
organisms, the most prominent being the
genetic code (with very few minor exceptions,
all organisms studied to date, use the same
nucleotide codons to specify the same amino
acids to be incorporated into proteins, even
though an enormous number of alternative
codes are theoretically possible and some
would work as well or better than the real one).

Other examples of genetic or molecular homologies are

proteins modified from ancestral proteins for new
A genetic homology: In almost
every organism, the same
functions, processed pseudogenes, shared
nucleotide triplets, or codons of chromosomal flaws (such as CMT1A repeats in chimps
the genetic code, specify the
same amino acids to be and humans), and of course, many shared functional
incorporated into proteins. This
chart shows a portion of the code.
genes.
Evolutionary Patterns
As noted in earlier lectures,
homoplasy is common and the
independent evolution of a
similar character or character
state in different taxa includes
convergence and evolutionary
reversals.

In convergence, superficially
similar features are formed by
Nonhomologous similarities (homoplasies). This shark and different developmental
Orca both have streamlined shapes, powerful tails, and
short fins or flippers, even though one is a fish and the other pathways.
a mammal. These similarities all make sense in light of their
function and are not homologous.
Evolutionary Patterns
In vertebrates the axons of the
retinal cells run over the retina and In cephalopods, the axons run directly from the
converge into the blind spot base of the retinal cells into the optic ganglion.

The eyes of a vertebrate (A) and a cephalopod (B) are an extraordinary example of convergent
evolution. Despite their many similarities, note several important differences, including
interruption of the retina by the optic nerve in the vertebrate but not the cephalopod. In
vertebrates, the axons (nerve fibers) of the retinal cells run over the surface of the retina, and
converge into the optic nerve, forming the “blind spot”. In cephalopods, the axons run directly
from the base of the retinal cells into the optic ganglion.
Evolutionary Patterns
(ancestral state: 8 plates)
The “c” plates
do not develop.
The “b” plates
fuse with the “a”
plate.
Darwin’s example of
convergence in barnacles
showing different ways in which
the number of shell plates of a
barnacle has been reduced to six
from the ancestral condition of
eight in two descendant genera,
Chthamalus and Balanus
(diagrams show shells in cross
section).
Evolutionary Patterns In parallelisms, it
is thought that
similar
developmental
modifications
evolved
independently
(often in closely
Parallel evolution. The evolution of feeding structures (maxillipeds) from thoracic related
legs in crustaceans is marked by parallel reduction or loss of expression of the genes
Ubx and abdA in the same thoracic segments. The photo of Paranebalia (Lepostraca)
organisms),
shows the ancestral condition: there is a mouthpart (MxII) on head segment (H3), and because they are
legs (each with 2 branches, En and Ex) on thoracic segments T1 and T2. In Mysidium
(Mysida), thoracic segment T3 has a normal leg (En branch in yellow), but the likely to have
appendages on T2 and especially T1 show maxilla-like modifications of the En branch
(green and red). The copepod Mesocyclops has similar modifications of morphology
similar
and gene expression. developmental
mechanisms to
begin with.
Evolutionary Patterns
Reversals complicate phylogeny
inference. (a) Read this tree up from
the root, and notice that a change in
the fifth position of this DNA
sequence creates a shared, derived
character in the descendant
populations. (b) If a reversal changed
the fifth position back to the ancestral
state later in the evolution of this
group, it would make it much more
difficult to infer the correct phylogeny.

An evolutionary reversal constitutes a return from a derived state

character state to a more ancestral state.

The diagram above shows this homoplasy at a molecular level but

morphologically it can occur as well (for example: nearly all frogs lack teeth on
their lower jaws (despite ancestors with teeth on both jaws, but one frog genus,
Amphignathodon “re-evolved” teeth in the lower jaw (which constitutes a
reversal to the ancestral state for this genus).
Evolutionary Patterns
Homoplasious features are often
(but not always) adaptations by
different lineages to similar
environmental conditions.

A correlation between a particular

homoplasious character in
different groups and a feature of
Bird groups in which similar bill shape has evolved
independently as an adaptation for feeding on nectar.
(A) A south American honeycreeper. Family
those organism’s environment or
Thraupidae (Cyanerpes caeruleus). (B) A Hawaiian
honeycreeper, family Fringillidae (Vestiaria
niche is often the best initial
coccinea). (C) A hummingbird, family Trochilidae
(Campylopterus hemileucurus). (D) A sunbird, evidence of the feature’s adaptive
family Nectariniidae (Nectarinia pulchella).
significance.
Evolutionary Patterns

Batesian mimicry: When disturbed, the

hawkmoth larva (left) resembles a
snake (right).

Mimicry is an interesting condition whereby features of

one species evolve to specifically resemble those of
another unrelated species.

Batesian mimicry is when either a palatable or an innocuous

animal evolved to resemble an unpalatable or dangerous animal
(the model) which teach predators by experience to avoid
attacking the mimic.
Evolutionary Patterns
Mullerian mimicry: both the cuckoo bee
(left), and the yellow jacket (right)
have toxin-releasing stingers and their
cross-mimicry in their appearance
benefits both species because predators
learn more quickly to avoid any prey
with such distinctive markings.

In Müllerian mimicry, two (or more) distasteful or dangerous

species evolved similar characteristics, and a predator that
associates the unpleasant features avoids attacking both species.

There are many mimicry “rings” involving both distantly related palatable
(Batesian mimics) and unpalatable (Müllerian mimics) moths and butterflies
sharing the same color pattern.

Mimicry evolution involves many complications including the spectrum of

intermediates between the Batesian and Müllerian extremes.
Evolutionary Patterns
Archaeopteryx, a bird with modern feathers and a dinosaur-
like skeleton making it a mosaic and because of mosaic
evolution it is not accurate to consider one species more
advanced than another.
The evolution of different characters
at different rates within a lineage is
called mosaic evolution meaning no
species evolves as a whole but
piecemeal (quasi-independently)
though those features that function
together will usually evolve in
concert, justifying the theory of
evolutionary mechanisms, where
evolution is not examined in terms of
entire organisms but rather in terms
of individual features (or the genes
underlying those features).
Characters that are retained with little or no
change over long periods within lineages
are conservative characters.
Evolutionary Patterns
Evolution is often gradual
(proceeds by small
incremental changes) rather
than by huge leaps (called
saltations), but whether this is
always true is unknown but
much debated.

While intermediate forms between

higher taxa are still somewhat
Variation in the shape and length of the bill among sandpipers. The three uncommon in the fossil record
vertical series are drawn to scale to match bill length differences. Note the
gradations in both curvature and length. The phylogeny of these species is
(though some do exist) gradations
not well resolved but the variation demonstrates how very different bills among living species are
could have evolved through incrementally small changes.
commonplace providing great
support for gradual evolution.
Evolutionary Patterns
In Bigoniaceae, terminal leaflets
In
Passifloraceae,
stipules are
of the tripartite leaves are
modified into suckers… One reason a homologous
character may differ so
modified into
tendrils.

greatly among taxa is that its

In
Ranuculaceae, …and tendrils.
form may have evolved as its
leaves are
modified into
tendrils
function changed and form
change is often correlated
with function.
In Rubiaceae,
inflorescence
petioles are
modified into
hooks.
The diagram shows how in many groups
Structures modified for climbing in vines of diferent plant that evolved a vinelike climbing habit, the
groups show that structures can become modified for new
functions, and that different evolutionary paths to the same structures that have been modified into
functional end may be followed in different groups climbing organs include roots, leaves,
leaflets, stipules, and inflorescences.
Evolutionary Patterns
The similarity between species
changes throughout ontogeny
such that their embryos will
look more similar than the
adults.

Features that are common to a more

inclusive taxon often appear before
the specific characters of lower-level
taxa (von Baer’s law).

However, contrary to Haeckel’s

Micrographs show the similarities—and differences—among several
vertebrate embryos at different developmental stages. Each begins
with similar basic structure (though they may acquire this structure at
different stages and sizes, but as the embryos develop, they become
less and less alike.
“biogenetic law” ontogeny does really not
recapitulate phylogeny (to a large extent
because various features develop at
different rates relative to one another and
embryos and juvenile stages have stage-
specific adaptations of their own).
Evolutionary Patterns Development underlies
In this primitive
whale from the
Eocene,
some common patterns
differentiation of
the teeth has
been reduced.
of evolution.

Most “reptiles” such as this extinct

lizard, have homodont (uniform) teeth
that are not individualized.
In modern toothed
The bodies of organisms
cetaceans such as
dolphins, the teeth are consist of modules which are
again homodont and no
longer individualized. distinct units with their own
genetic specifications,
developmental patterns,
locations, and interactions
Typical mammals, such as this elephant shrew,
have heterodont, individualized teeth, differentiated
into incisors, canines, premolars, and molars.

The teeth of reptiles and mammals provide an example of the acquisition and loss of with other modules, and
individualization. The teeth of most reptiles are uniform; teeth become individualized
during the evolution of mammals. Distinct tooth identity was later lost in the
some of these modules lack
evolution of toothed whales. distinct identities and may be
considered aspects of a
single character (called
serially homologous if
arrayed along a body axis
and homonymous if they are
not).
Evolutionary Patterns Heterochrony is broadly
defined as an evolutionary
change in the timing or rate
of developmental events
(though other developmental
mechanisms can produce
similar changes).
Largely global heterochronic
changes (affecting many
characters simultaneously) are
shown by cases in which
developmental time of most
Paedomorphosis in salamanders. (A-B) The tiger salamander, like
somatic features (except
most salamanders undergoes metamorphosis from (A) an aquatic reproductive structures) is altered
salamander to (B) a terrestrial adult. (C) The adult axolotl with gills
and tail fin resembles the larva of its terrestrial relative, and it remains
relative to the initiation of
aquatic throughout its life This is an example of paedomorphosis: the reproduction.
evolution of a more juvenilized morphology for the reproductive
adult.
At left is an example of paedomorphosis,
its opposite (called peramorphosis) is the
evolution of delayed maturity resulting in
larger size, associated with extended
development of “hyper-adult” features.
Evolutionary Patterns
Allometry refers to the
differential growth of different
parts or dimensions of an
organism during its ontogeny
and changes in the allometric
growth of individual characters
(called “local” heterochrony)
appear to have played a very
important role in evolution.

Many evolutionary changes can be

Allometric differences in the length of upper and lower jaws among described as if local heterchronies
three closely related families of fishes. The differences in form can be
accounted for by changes in the rate of jaw growth relative to body
had altered the shape of one or more
growth. characters (as at left)
Evolutionary Patterns In the allometric growth
equation y = bxa, y and
x represent
measurements (e.g.,
height and width), b is
the value of y where it
intercepts the vertical
axis. a is the allometric
coefficient (describing
the relative growth rates
of x and y) so if a = 1
Hypothetical curves showing various allometric growth relationships between two body then growth is
measurements, y and x according to the equation y = bxaa (A) Arithmetic plots. Curves 1
and 2 show isometric growth (a = 1), in which y is a constant multiple (b) of x. Curves 3 isometric, if a > 1
and 4 show positive (a > 1) and negative (a < 1) allometry respectively. (B) Logarithmic
plots of the same curves have a linear slope (log y = log b + a log x. The shape
(positive allometry) then
differences depend on a. Curves 1 and 2 have slopes equal to 1. y increases faster than
x, and if a < 1 (negative
allometry) then y
increases more slowly
than x.
Evolutionary Patterns This diagram at left
demonstrates that
In the
descendant,
feature y
becomes
more
peramorphosis can
exaggerated
relative to x. result from
evolutionary changes
in either rate of
development or the
duration of
development due to a
A diagrammatic representation of some forms of heterochrony, expressed as
logarithmically plotted allometric growth of structures or dimensions y and x. change in α or β.
The x-axis may represent body size, the y-axis a character such as leg length.
The dashed line, with slope 1, is for reference. (A) The blue line shows
ontogenetic change in the ancestor from age α to age β. Growth is positively
allometric (slope > 1). (B) A longer growth period (extension of growth to age β Here x represents body size, y
+ Δβ; purple arrow) results in peramorphosis: an exaggerated structure y in the
descendant. is the size of some feature that
begins to develop at at age α
(onset) and ends at age β
(offset). Here peramorphosis
can result if the duration of
development is extended (a
change from β to β + Δβ).
Evolutionary Patterns
The gigantic antlers of the
extinct Irish elk, which
were larger in relation to
body mass than those of
any other deer, are a
peramorphic feature
associated with the
animal’s extended
development to a larger
size.

Perhaps the most famous example of allometry and peramorphosis is

largest of deer, the extinct “Irish elk” (Megaceros giganteus). Its antlers
were larger relative to body mass, than those of any other deer.
Evolutionary Patterns Paedomorphosis can
be caused by cessation
. of growth at an earlier
age (β - Δβ) in a form
of paedomorphosis
called progenesis or by
reducing the growth of
character y in a form of
paedomorphosis called
neoteny.
Neoteny is the retention of
juvenile morphological traits
in the sexually mature adult,
while progenesis is
increased rate of sexual
development leading to
early maturity in an adult
that remains very small in
size.
Evolutionary Patterns The axolotl (shown
earlier) is an example
of neoteny (it reaches
the same size as its
metamorphosing
relatives) while the tiny
salamanders of the
genus Thorius are
progenetic, having
The space between the paired frontal and
parietal bones is a juvenile feature that is
retained in adult Thorius.
In most salamanders, the frontal and parietal
bones grow together by adulthood. features that are
characteristic of the
Skulls of the progenetic dwarf salamander Thorius and a typical
juveniles of larger
nonprogenetic relative, Pseudoeurycea. The skull of adult Thorius has a
number of juvenile features.
species of
salamanders (appears
as if they have
abbreviated
development).
Evolutionary Patterns
Plants of the genus Philodendron, such as this
Jamaican climber, are lianas. Many lianas, which
include several genera besides Philodendron, have
evolved exposed roots that grow from an aerial stem.
Heterotopy is the evolutionary
change in the position within the
organism at which a phenotypic
character is expressed.
Studies in the distribution of gene
products have revealed many
heterotypic differences among species
in gene expression sites.
Heterotypic differences are common among
species of plants (for example the stem not
leaves of cacti photosynthesize or the growth
of roots along the stems of lianas (some for
holdfasts others for the usual root function).
Evolutionary Patterns Vertebrate bones can
provide an example of
heterotopy in animals,
particularly regarding
phylogenetically new
bones called
sesamoids that
develop in tendons or
other connective
tissues subject to
The right hand of two members of the bear family, a brown bear at left and a
panda at right. A small sesamoid bone in bears has been modified into a false stress.
finger (“thumb”) in the panda which it uses to help manipulate bamboo on
which it feeds. This is an example of heterotopy in animals.
Many dinosaurs had ossified
tendons in their tails and the
giant panda (Ailuropoda
melanoleuca) is famous for its
elongated sesamoid (from the
wrist) that serves as its “thumb”.
Evolutionary Patterns Both paleontological and
phylogenetic studies show
that there have been great
increases in complexity in
the history of life, but it is
often surprising to learn
that morphological
simplification (the reduction
or loss of structures) is a
very common trend witin
clades.
An example of reduction and loss of structures during evolution. The number
of bones in the skull is higher in early fishes (such as the Devonian
Eusthenopteron), from which amniotes were derived, than in early amniotes
(such as the Permian Milleretta). Among later amniotes are placental
mammals such as the dog (Canis) in which the skull sports fewer elements For example, many flowering
still (the reduction in the number in the lower jaw is especially notable).
plants show reduction in the
number of floral elements while
vertebrates show reductions in
various bones (such as toes or
skull bones).
Evolutionary Patterns
Frontal view of the heads of male flies in the dispar subgroup of Zygothrica (Drosophilidae).
Note the gradation in the shape of the head and eyes among the species and within prodispar,
dispar, and exuberuns. These gradations form a phylogenetic series from narrow to broad, as
indicated by the phylogenetic analysis which is based on a number of morphological features.

The term evolutionary trend can refer to a succession of

changes of a character in the same direction, either in a single
lineage or, often, in many lineages independently.
An example is in the fly genus Zygotherica showing a directional trend towards wider
heads (also seen in three other fly clades).
Evolutionary Patterns
During evolutionary
radiations (the divergent
evolution of numerous
related lineages within a
relatively short time) the
lineages are modified for
different ways of life (often
termed an adaptive
radiation).

Here lineage characteristics do

not show sustained directional
Adaptive radiation of Darwin’s finches in the Galápagos Islands and Cocos
Island. The bills of these species are adapted to their diverse feeding habits. trends but rather evolutionary
radiation and may be the most
common pattern for long-term
evolution.
Evolutionary Patterns
Some members of the Hawaiian silversword alliance with different growth forms. (A)
This rosette plant lacks a stem except when flowering (as it is here). (B) is a stemmed
rosette plant and (C) is a small shrub.

Though few species of plants and animals colonized the

Hawaiian Islands, these silversword plants and their
relatives occupy habitats ranging from exposed lava rock
to wet forest and show growth forms that include shrubs,
vines, trees, and creeping mats; but despite these
differences they can form fertile hybrids when crossed.
Evolutionary Patterns
A sample of the diverse head shapes among the Cichlidae of the African Great Lakes.
The morphological differences are associated with differences in diet and feeding mode.

These cichlid fishes from the Great Lakes of east

Africa have undergone spectacular adaptive
radiations, with the myriad species varying in color,
body form, and tooth and jaw structure (reflecting their
diverse feeding habits), though each lake supports a
monophyletic clade that radiated rapidly.
Evolutionary Patterns
Genome size variation.
The bars indicate the range
of size for particular
clades. Taxa are arranged
from top to bottom in
order of supposedly
increasing organismal
complexity. Within
eucaryotes, there is little
relationship between
genome size and
oraganismal complexity.
This “C-value paradox”
may be a result of great
variation among lineages
in the amount of repetitive
(noncoding) DNA. (1 pg
of DNA is approximately
equivalent to 1 billion
base pairs.)

The expectation is that physiologically and

behaviorally complex organisms would have
more complex and larger genomes and on a
broad scale this holds true.
Lack of correspondence between genome size
and phenotypic complexity in eukaryotes is the
C-value paradox.
Evolutionary Patterns
Number of genes estimated for some eukaryotes with fully sequenced
genomes, arranged on a provisional phylogeny. Multicellular
organisms with tissue organization (plants and animals, represented by
blue branches) have more genes than single-celled organisms (red
branches) or multicellular organisms that lack pronounced tissue
organization (some fungi and slime molds; green branches)

When non-coding, repetitive DNA is eliminated from the

comparisons, the phylogenetic transition from unicellular to
multicellular eukaryotes shows a considerable increase in gene
number, but surprisingly little variation among plants and
animals.
But endosymbiotic microrganisms living within eukaryotic hosts have smaller
genomes than their free-living relatives
Evolutionary Patterns
Orthology and paralogy in gene families are two forms of homology.
When an ancestral gene (α) undergoes duplication, the resulting two
genes (α + β) have a paralogous relationship to one another (blue
arrow). A speciation event after duplication results in divergence of
the ancestral set of two paralogous genes. Within the genomes of the
two diverged species, the α + β still have a paralogous relationship
(blue arrows). However, the copies of α in species 1 and species 2 are
orthologous (red arrows), because the two genes are related to one
another via speciation, not duplication. Likewise, the copies of β in
species 1 and 2 are orthologous.

Gene duplication is a mutational event by which a new gene (β)

arises as a result of a copy of a preexisting gene (α), so that a
single gene locus in an ancestor is represented by two loci in the
descendent 9and they undergo further evolutionary sequence change and
can thus be distinguished.
Evolutionary Patterns
The phylogeny of genes in the globin family in the human genome.
Myoglobin consists of a single protein unit, whereas mammalian
hemoglobins consist of four subunits, two each from the α and β
subfamilies. Each branch point on the tree denotes a gene duplication
event; some of the events are marked with estimates of when
duplication occurred. The origin of hemoglobin and myoglobin from a
common ancestor gene occurred in the ancestor of all vertebrates, but
the αand β hemoglobin subfamilies originated by duplication in an
ancestor of the jawed vertebrates. The duplication of the β hemoglobin
into two genes occurred in the ancestor of placental mammals, since
the Aγγ/Gγγ/ε genes are lacking in monotremes and marsupials. In some
instances, one of the pair of genes formed by duplication became a
nonfunctional pseudogene, symbolized by Ψ.

The phylogenetic relationships among orthologous and

paralogous genes can be determined by standard phylogenetic
methods.
The process may repeat over evolutionary time, generating gene
families such as seen here for hemoglobin.
Evolutionary Patterns
The nonvertebrate chordates (tunicates and
Amphioxus) have a single cluster of Hox
genes, represented by a circle. The jawless
lamprey has four Hox clusters, implying that
the single cluster has undergone two
duplications. the gnathostomes (jawed
vertebrates0 also have four clusters that have
arisen by two duplications. It is not certain,
however, that the gnathostome clusters are
homologous to those in the lamprey, some of
which may have arisen independently. Three
of the four clusters of the ancestral
gnathostome were duplicated in the ancestor
of teleost fishes.

Many genes are duplicated as parts of large chromosomal blocks

(paralogous regions) that often contain hundreds of genes.
Polyploidy (entire genome duplication) occurs commonly in plants
and some animals.
In the example shown here with Hox genes, it is unclear whether the whole
genome or just parts of it, underwent two successive duplications in the
ancestor of jawed vertebrates, but teleosts do appear to show a polyploid
ancestor.