Workshop on Population and Speciation Genomics 2020 W.

Salzburger | Adaptive Radiation

Adaptive Radiation
by Prof. Walter Salzburger
Zoological Institute, University of Basel, Vesalgasse 1, 4051 Basel, Switzerland

The diversity of life on Earth is governed, at the MACROEVOLUTIONARY scale, by two antagonistic ————————
MACROEVOLUTION
processes: Evolutionary radiations increase and extinction events decrease the organismal diversity on our Evolution on the
grand scale, that is,
planet through time. Evolutionary radiations are termed adaptive radiations if new lifeforms emerge evolution at the level
rapidly through the extensive ecological diversification of an organismal lineage. of species and above.

Examples of adaptive radiations ECOLOGICAL NICHE

The relational
Adaptive radiation refers to the evolution of ecological and morphological disparity within a rapidly position of a species
or population in an
diversifying lineage. It is the diversification of an ancestral species into an array of new species that ecosystem. It includes
occupy various ECOLOGICAL NICHES and that differ in traits used to exploit those niches. Adaptive the interactions of all
biotic and abiotic
radiation includes the origination of both new species (speciation) and phenotypic disparity. factors that determine
how a species meets
Archetypal examples of adaptive radiations include Darwin’s finches on the Galápagos archipelago; its needs for food and
shelter, how it
silversword plants on Hawaii; anole lizards on the islands of the Caribbean; threespine stickleback fish survives, and how it
in north temperate waters; and cichlid fishes in the East Africa Great Lakes and in various tropical reproduces.
crater lakes (see FIGURE 1). Adaptive radiations are also visible in the FOSSIL record. For example, the
FOSSIL
CAMBRIAN EXPLOSION is considered an adaptive radiation.
Any preserved
evidence or traces of
life from a past
Anolis lizards Cichlid fishes geological era, such
as impressions and
remains of organisms
mouth embedded in rocks or
morphology in marine or lake
sediments, or
body mineralized bones or
shape plant material.

limb
image sources: anolistollis.wordpress.com, Hofer & Salzburger (2005), Grant (1985), Reno (2014).

CAMBRIAN
length EXPLOSION
(also known as
body “Cambrian
size evolutionary
radiation”)
An evolutionary event
in the Cambrian
period, documented
in the fossil record
(e.g., in the Burgess
shale), during which
Darwin’s finches Threespine stickleback fish most of the animal
phyla emerged within
body shape a relatively short
and size period of time.
————————

armor
plates

beak size
and shape

FIGURE 1. Famous examples of ongoing vertebrate adaptive radiations and examples of adaptive traits in the respective radiation.

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Characteristic features of an adaptive radiation ————————

ENVIRONMENT
An adaptive radiation is defined by four characteristic features, which are also used to detect instances The surroundings in
which a species lives
of adaptive radiations (see Schluter 2000): and operates,
including all of the
1. Common ancestry: Members of an adaptive radiation are characterized by their common ancestry, biotic and abiotic
that is, all species of an adaptive radiation go back to a single common ancestor. Common factors that affect the
survival and
ancestry is not the same as monophyly, as not all descendants of the common ancestor need to be development of an
included in the radiation. organism or
population.
————————
2. Phenotype-environment correlation: In an adaptive radiation, there is a fit between the diverse
phenotypes of the descendant species and the divergent ENVIRONMENTS in which they live. For
example, the body and limb size of Anolis lizards matches the twig diameter in their habitat (see
FIGURE 1). BOX 1 provides further example of phenotype-environment correlations.

3. Trait utility: The morphological and/or physiological traits that differ between the descendant
species of an adaptive radiation are “useful” to exploit the respective ecological niche. For
example, the different bills of Darwin’s finches are useful for a particular food type.
4. Rapid speciation: In an adaptive radiation, speciation is typically rapid. Speciation during adaptive
radiation can be allopatric, but is primarily parapatric or sympatric, as adaptive radiations are, in
most cases, confined to a certain geographic area.

BOX 1: Examples of adaptive radiation and the associated phenotype-environment correlations (modifi ed
from Schluter 2000).

Taxon Region Phenotype-environment Trait Performance

correlation

Animals
Darwin's finches Galápagos bill size and shape - seed size and seed handling time; breaking stress;
(Geospiza) hardness crushing force

Crossbills North America bill size - cone strength, cone stage handling time
(Loxia)

Tits Eurasia body size and limb length - substrate; foraging ability; hanging and perching
(Parus) bill shape - habitat ability

Caribbean lizards Greater Antilles body size and hindlimb length - perch sprint speed; jump distance; running
(Anolis) diameter and hight stability

Sunfishes North America bod size and gape - prey size; handling time; crushing force
(Centrarchus) pharyngeal jaw musculature - snail

Stickelback fish Northern body size - prey size; foraging success; growth rate
(Gasterosteus) hemisphere body shape - habitat

Lake whitefish Northern gill rakers - habitat foraging ability

(Coregonus) hemisphere

Cichlid fishes Southern premaxilla angle and length - diet biting and suction force
(Cichlidae) hemisphere pharyngeal jaw bone - diet

Plants
Coumbines North America flower orientation - pollinators; pollinator visits; pollen removal
(Aquilegia) spur length, color - pollinator

Brittlebush North America pubescence - temperature, moisture transpiration, photosynthesis

(Encelia) regime

Silverswords Hawaii tissue elastic modulus - habitat turgor pressure

(Dubautia)

Orchids Mediterranean flower morphology, color, flowering pollination success

(Ophrys) time - pollinator

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Triggers of adaptive radiation ————————

ECOLOGICAL
An adaptive radiation can occur under a number of different circumstances, which have in common OPPORTUNITY
The environmental
that they create ECOLOGICAL OPPORTUNITY for the subsequent radiation to occur: conditions that
permit the persistence
1. Colonization of a new area: Many adaptive radiations occurred after an ancestral species colonized a of an evolutionary
new area in which the adaptive radiation takes place (e.g., the Galápagos archipelago in case of lineage.
Darwin’s finches, the Hawaiian islands in case of Drosophila or silversword plants, the islands of the
RESOURCES
Caribbean in case of Anolis lizards, and various small and large lakes in case of cichlid fishes). A substance or object
Adaptive radiations are often connected with the emergence of novel (empty) habitats such as in the environment
required by an
islands or lakes. Such newly colonized areas are typically characterized by reduced predation organism for growth,
pressures and competition on the one hand, and un- or underexploited RESOURCES on the other maintenance, and
reproduction.
hand. Resources can be
exploited (consumed)
2. Extinction or replacement of antagonists: An adaptive radiation may occur after competitors become and thus become
unavailable.
extinct or get replaced. That way, ecological niches that were previously occupied by other taxa
become vacated. Here, extinction refers to a rapid (cataclysmic) process, whereas replacement
EVOLUTIONARY
refers to a more gradual process, for example due to environmental change. INNOVATION
Lineage-restricted
3. Adaptive breakthrough: An adaptive radiation may be initiated by the evolution of a new adaptive trait with a
trait that allows a taxon to outcompete other taxa or to exploit previously underutilized resources. qualitatively new
function, such as
EVOLUTIONARY INNOVATIONS are termed “key innovations” if they are responsible for an adaptive feathers in birds or
radiation. Examples of key innovations are the Antifreeze glycoproteins in Antarctic notothenioid the flower of
flowering plants.
fish (‘icefishes’) allowing them to survive in the ice-cold waters off Antarctica or the nectar spurs in
the flowers of columbine plants leading to a strong associated between plant species and pollinator. MASS EXTINCTION
An extinction event
during which more
than three-quarters
Extinction of all species on our
planet disappear in a
The history of life can be viewed as a cumulative story of evolutionary, often adaptive, radiations, geologically short
time interval.
interrupted by extinction events (on top of the more or less gradual processes of speciation and small- ————————
scale extinction). The fossil record documents past radiations and extinctions, but is by no means
complete. This is because the long-term preservation and subsequent discovery of fossils is an
extremely rare event; fossil remains are usually incomplete or damaged; the fossilization process
requires specific settings and conditions, so that fossils are typically restricted to particular
environmental settings (such as caves, sediments, deserts); finally, the fossil record is heavily biased
towards organisms with hard parts such as mollusks or vertebrates, whereas fossils of soft-bodied
organisms are only rarely to be found.
Extinction is a common process in evolution and it is believed that 99% of all species that ever existed
since the dawn of life on Earth about 3.5 billion years ago have meanwhile gone extinct. However,
extinction is normally balanced out by speciation (and radiation). The fossil record (primarily the one
in the marine realm) nevertheless documents five MASS EXTINCTION events (see BOX 2). Some scientists
believe that a sixth such event is currently ongoing.

BOX 2: The ‘big five’ mass extinction events (modifi ed from Barnosky et al. 2011).

Event end point duration % genera lost (estimate) % species lost (estimate)

Ordovician event ca. 443 My ago 3.3 - 1.9 My 57 86

Devonian event ca. 359 My ago 29 - 2 My 35 75

Permian event ca. 251 My ago 2.8 My - 160 Ky 56 96

Triassic event ca. 200 My ago 8.3 My - 600 Ky 47 80

Cretaceous (K-T) event ca. 65 My ago 2.5 My - 1 y 40 76

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References:
Barnosky et al. (2001) Has the Earth’s sixth mass extinction already arrived? Nature 471: 51-57.
Berner D & W Salzburger (2015) The genomics of organismal diversification illuminated by adaptive radiations. Trends in Genetics 31: 491-499.
Ridley M (2004) Evolution. Blackwell.
Schluter D (2000) The ecology of adaptive radiation. Oxford University Press.

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