Extinctions of species

A normal and common process in evolution and

ecology
If over an extended period of time the birth rate of a
species is less than the death rate, then extinction will
eventually occur
A species goes extinct if it is not able to adapt to changes
in its abiotic environment, or compete effectively with
other organisms for resources.
>99% of all species that existed in the Phanerozoic went
extinct
Average duration of species: 12My
Incumbency: extinctions of incumbents promote
subsequent evolutionary radiations for previously minor
groups within the vacated space
 Intrinsic biotic properties of species
Body size, metabolic rate, physiological tolerance,
feeding type
Dispersal ability
Habitat generalists vs. specialists
=>
Size of biogeographic range

End Cretaceous bivalve genera:

Large biogeographic distributions
enhance survival
Extrinsic biotic properties of species

Predator-prey relations (coevolution)

Red Queen Hypothesis (Van Valen 1973)
Based on the reflection by the Red Queen in Lewis
Carroll's "Through the Looking Glass" that "in this
place it takes all the running you can do, to keep in
the same place,"
"For an evolutionary system, continuing development
is needed just in order to maintain its fitness relative
to the systems it is co-evolving with"

Disturbances in food chains (crucial role

of primary producers)
Interspecific competition for resources
 Abiotic factors involved in extinctions
Bursts in volcanic activity (basaltic traps): change in chemical
composition of oceans and atmosphere
Sea-level changes: areal decrease of marine platform, changes
in oceanic circulations, rising anoxia

Tectonics: paleogeographic changes (ocean closure or opening,

mountain building)
Climatic changes (ice-house/green-house)
Asteroid impacts
Diversity and major Extinctions

Estimated percentages of
extinctions (genera):

1: End-Ordovician: 57%
2: End-Devonian :60%
3: End-Permian: 80%
4: End-Triassic : 48%
5: End-Cretaceous :50%
Main victims of the End-Permian extinction

† Fenestrate Bryozoan

† Blastoids † Blastoids and Crinoids Brachiopod

† Rugose Corals
† Trilobite † Rostroconch
 Extinction metrics
Taxonomic and stratigraphic biases

Influence of the taxonomic structure:

an equal number of extinction of species result in a greater
number of genera extinctions at the first than the second
extinction event

Influence of stratigraphic gaps:

backward smearing of last appearances of species that went
extinct at a major extinction
Survival of Lazarus taxa leads to postulate the existence of
refuges
Diversity of marine classes in time

650 My of geologic history

2800 Marine families distributed in 91
Metazoan classes
82 stratigraphic stages
Resolution: 8 My on familial diversity of
classes

How does richness of classes behave in

time?
Search associations of taxa that share
common times of maximum and
minimum diversity

Factor analysis (Q-mode)

Classes treated as variables, and are

sampled in each time interval
 Evolutionary faunas

Result of analysis: only 3 factors are needed to account

for more than 90% of the data
These 3 factors represents the 3 evolutionary faunas of
the Phanerozoic record
Cambrian fauna dominated by Trilobites, inarticulate
brachipods, etc.
Paleozoic fauna dominated by attached, sessile,
epifaunal invertebrates
Modern fauna dominated by mobile organisms
 Magnitude of Extinctions
Overall decline of intensity
Major extinctions separated from one another by intervals of
lower extinction intensity in an approximately cyclic manner

MacLeod 2003 (% extinction at the genus level)

 Magnitude of Extinctions
Mass extinctions vs. Background extinctions?

MacLeod 2003
 Intensitiy of Extinctions
Genus level
Intensities globally decline through time
Paleozoic « background » extinctions are even larger than
some of the Mesozoic or Cenozoic « mass » extinctions

MacLeod 2003

A real feature or a statistical artefact?

Average duration of Pz stages compares to that of Mz and Cz stages
Null hypothesis (random walk through time) rejected (Monte Carlo-based randomization)
 What regulates the long-term
intensitiy of extinctions?
Biotic hypotheses:
1. Improvement of species fitness over time

2. Increase in species extinction resistance conferred by:

- increase in the number of species per clade with
consequent broadened geographic distributions
- better knowledge of Cenozoic faunas (greater
availability of outcrops)
- progressive radiation of clades into marginal
environments where resistance is gained
by tolerance to environmental fluctuations
 What regulates the long-term
declining intensitiy of extinctions?
Abiotic hypothesis:
Secular increase in nutrients delivrery to marine habitats as a
primary cause for decreasing background of intensities
Direct impact on marine phytoplankton which occupies the base
of marine trophic chains and controls the rate at which
inorganic nutrients are made available to marine trophic
systems
First order trends in environmental proxies:
• 34S (marine circulation intensity)
• 87Sr/86Sr (rate of continental weathering and nutrient runoff)
• 13C (rates of photosynthesis and nutrients recycling
efficiency in organic systems)
 Marine phytoplankton:
The base of the trophic pyramid
Nutrient-limited by P, Fe, N, Si
Converts inorganic nutrients into organic matter
Nutrient Sources:
erosion of continents (major source)
submarine volcanism
Nutrient Pathways:
direct input via river discharge
recycled input via upwelling zones and bioturbation (where nutrients freed
by decomposition are brought into contact with planktonic and benthic
marine biotas)

Diatom
Coccoliths
Dinofagellate

Wind-driven coastal upwelling West-african upwelling

 Environmental proxies

Real features or statistical artefacts?

Null hypothesis (random walk through time) rejected (Monte Carlo-based randomization)
MacLeod 2003
 A proxy for terrestrial runoff and
nutrient input: 87Sr/86Sr

Alpine-Himalayan
orogeny

Weathering of continental
igneous rocks: higher proportions
of 87Sr transfered by river
discharge

End-Devonian
End-Ordovician End-Permian

MacLeod 2003

First order trend: declining nutrient input

Positive deviations: active mountain-building
Negative deviations: terrestrial biomass decreases the amount Hydrothermal input from oceanic
crust into sea-water: relatively
of transfer from continent to oceans, but increases chemical
high proportions of 86Sr
weathering (soils)
 A proxy for oceanic circulation
and oxygenation: 34S

Present-day thermohaline oceanic

circulation

End-Permian (superanoxia)

MacLeod 2003
Sluggish circulation of Pz. oceans in comparison with Mz. and Cz. oceans
Supported by greater number of black shales events in Pz.
Sluggish circulation and highly stratified water column may have
confined high-nutrient bottom waters to anoxic basins, thus starving the
photic zone and deeper shelves Black shales deposit
 Extinction-intensity patterns and
oceanic circulation
High amplitude of early Pz.
Extinctions->
Early Pz. Environmental
instability

If the four major extinctions from

the Carboniferous on are removed,
a striking decrease in amplitude of
extinction appear.

MacLeod 2003

Stabilisation of background extinction intensity signal, combined with progressive decline,

reflect damping of extinction probalities by improved nutrient delivery resulting from a long
term and progressive improvement of marine circulation.
 The long-term decrease of
extinction-intensity:
A dynamical interaction between
tectonics and evolution of biomass?
Roles of tectonics:
Configuration of oceanic basins (affects climate patterns of oceanic circulation)
Continental positions (affect climates)
Mountain building (affects climate of and rates of weathering)
Mid-oceanic spreading ridges (provide nutrients via hydrothermalism)

Nutrient delivery to photic zone and shallow water marine habitats:

Long term decrease of terrestrial input (87Sr/86Sr)
Long-term increase in improvement of oceanic circulation and oxygenation (34S)->
increased upwellings

Roles of organic evolution:

Development of strategies for storage of nutrients in soils and photic zone and
improved recycling through longer food-chains
In return, large biomass reservoirs influence climate and atmospheric composition
 A proxy for productivity and nutrient
recycling: 13C
The first order trend:

Early Pz. « light » (negative)

values -> low productivity or
oxydation of organic carbon
reservoirs, low nutrient
recycling

Mz. And Cz. « heavier »

(positive) values -> increased
rates of marine and terrestrial
Radiation of higher photosynthesis; increased
land plants oxygenation of marine carbon
reservoirs, high nutrient
recycling

MacLeod 2003

The secular increase of 13C values as an explanation for the nutrient input paradox:
storage of nutrient into a more usable form (biomass) and more efficient recycling by
specialization of phytoplankton, expansion of deposit feeding niches, and lengthening of
food chains
Ecological linkage between marine
phytoplankton and evolutionary
faunas?
 Ecological linkage with phytoplankton:
The Echinoid exemple

Regular echinoids
grazers, epifaunal

The first successful invasion of the deposit-feeding adaptive zone

coincides with: Irregular echinoids
phytoplankton bloom deposit feeders,
increased circulation rates and productivity infaunal
BUT drop down of nutrients transfer from continent
 Flood basalts and extinctions:
a near-perfect correlation

Deccan Traps (K/T Bdry) >2000m, >512000Km3

World map of main traps and hot spots

Effects:
change compositions of
atmosphere and oceans
(CO2, SO2)
disruption of biological
pump, climate, and
oceanic circulation

Age-correlation
between traps and
extinctions
Mantle plumes:
the deep source of
flood basalts Courtillot 1999
Environmental effects of flood
basalts
Sea-level, flood basalts and impacts
Additional useful videos to this course

Animal Dawn - Research Tuesday - Presentation

https://www.youtube.com/watch?v=39vlt1N5o5A

Permian - Triassic Mayhem: Earth's Largest Mass Extinction

https://www.youtube.com/watch?v=J8rynQqg9x8

EGU2010: Mass extinctions, volcanism, impacts and global

environmental change (Press Conference)
https://www.youtube.com/watch?v=H8rc8vkDijs