Distributions in space

Distributions in spaceBiogeography

Tries to understand large scale distributions of living thinks

Evolutionary Ecology

Tries to understand patterns of species diversity through evolutionary history

Macroecology

Tries to link both disciplines and to explain larges scale ecological patterns and processes in space and time

Communitystructure

Lifehistrory

traitsPhenology

Phylogenetic constraints

Speciesassemblage

rules

Niche History

Character evolution

BiogeographyBiotic interactions

Chance processes

Macroecology integrates biogeographic and evolutionary research in an interdisciplinary way.

It tries to explain community structure from a top down (instead of bottom up) perspective.

Basic tools are spatially explicit models and meta-analysis.

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Spatial scale [m2]

Tem

pora

l sca

le [d

ays]

z

patches

Annualecosystemprocesses

Processesin ecological

time

Annual regionalspecies turnover

Landscapeprocesses

Landscapeprocesses in

evolutionary time

Continentalprocesses in

evolutionary time

Continentalprocesses in

ecological time

Macroecology

Ecological processes

Evol

ution

ary

proc

esse

s

Evolutionary processes

Ecological processes

PredationDisturbanceCompetition

Dispersal MetapopulationsSpatial processes

Speciation ExtinctionGeological processes

FluctuationsLocal species turnover

Dispersal MetapopulationsMetacommunities

Speciation ExtinctionClimatic processes

Theory of Island biogeography

The theory of island biogeography tries to understand species diversity on all sorts of isolated islands from stochastic colonization of islands and random extinction on islands.

Colonization rates depend on island area and isolation. Extinction rates depend on island area only.

The model is species based

Robert MacArthur (1930-1972)

Edward O. Wilson(1929-)

Species richness

Immigration Extinction

Equilibrium species richnessRa

te

Two islands

Species richness

Immigration Extinction

Equilibrium species richness

Rate

One islands

The Galapagos Islands

near

far

small

large

Theory of Island biogeography

Isolation

Spec

ies r

ichn

ess

S = S0e-kI

Area

Spec

ies r

ichn

ess

S = S0Az

The species – area relationshipThe species – isolation relationship

Land plant of Britain from Watson (1859)

y = 433.2x0.10

R2 = 0.98

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Num

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Diamond 1972, PNAS 69: 3199-3203

Avifauna of New Guinea

The increase of species richness with sample size

Parasitoid Hymenoptera on a dry meadow on limestone,

Ulrich 2005Butterfly catches by

Preston 1948

Species richness on

deep sea mounts,

Forges 2000

Increase of land plant families in evolutionary time, Knoll 1986

Increase of herbivores

on bracken, Lawton 1986

𝑆=𝑆0 𝑁 𝑧 𝑙𝑛𝑆=𝑙𝑛𝑆0+𝑧𝑙𝑛𝑁The power function species – sample size relationship

𝑆=𝑆0 𝐴𝑧 𝑆=𝑆0 𝑡𝜏The species – area relationship The species – time relationship

𝑆=𝑆0 𝐴𝑧 𝑡 𝜏

The species – area - time relationship

Collembolan species richness across Europe

𝑆=1.36 𝐴0.43

1. The number of species counts increases with area and time.

2. This relationship often follows a power function

3. The slope z of this function measures how fast species richness increases with increasing area. It is therefore a measure of spatial species turnrover or beta diversity

4. The intercept S0 is a measure of the expected number of species per unit of area. It is therefore a measure of alpha diversity

5. Changes in slope through time point to disturbances like habitat fragmentation or destruction

6. The slope of the species – time relationship is a measure of local species extinction rate.

The species – time relationship

Local species area and species time relationships in a temperate Hymenoptera community studied over a period of eight years.

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A

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B

0.70.80.9

11.11.21.31.4

0 5 10t

Turn

over

C

S = S0Az S = S0tt

S = S0Aztt

The accumulation of species richness in space and time follws a power function model

S = (73.0±1.7)A(0.41±0.01) t(0.094±0.01) The mean extinction rate per year is about 9%

Coeloides pissodis (Braconidae)

Species - area relationship of the world birds at different scales

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Area [Acres]

Num

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cies

small areas: z = 0.43

within a regional pool: z = 0.09

between biotas: z = 0.53

Regional SARs have slopes between 0.1 and 0.3.Local and continental SARs have slopes > 0.25.

Preston 1960, Ecology 41: 611-67

The species – area relationship of plants follows a three step pattern as in birds

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1.E-04 1.E-02 1.E+00 1.E+02 1.E+04 1.E+06 1.E+08 1.E+10 1.E+12

Area [km 2]

Num

ber o

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cies

Local scale: z = 0.25

Regional scale: z = 0.14

Intercontinental scale: z = 0.5Shmida, Wilson 1985, J. Biogeogr. 12: 1-20

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Latitude

Spe

cies

Latitudinal gradients in species richness

New worlds birds

Pacific shelve

mollusks

The peak in species richness is not exactly at the equator

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Mean temperature

Spe

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z

Western Atlantic gastropods

Eastern Pacific gastropods

Ecological hotspots

34 regions worldwide where 75% of the planet’s most threatened mammals, birds, and amphibians survive within habitat covering just 2.3% of the Earth’s surface.

Biodiversity is most sensitive to minimum temperatures and the temperature range

The latitudinal distribution of temperatures

The general patternHillebrand (2004, Am. Nat. 163: 192-211 ) conducted a meta-analysis for about 581

published latitudinal gradients

Regional

Local

Scale

High

Low

Global richness

High

Low

Body size

High

Low

Tropic level

New world

Old world

Longitude

Terrestrial, marine

Freshwater

Realm

Latitude

Spec

ies r

ichn

ess

• Nearly all taxa show a latitudinal gradient

• Body size and realm are major predictors of the strange of the latitudinal gradient

• The ubiquity of the pattern makes a simple mechanistic explanation more probable than taxon or life history type specific

Counterexamples

These taxa are most species rich in the northern Hemisphere

Soybean aphid, Photo by David VoegtlinThe sawfly Arge coccinea, Photo by Tom Murray

The ichneumonid Arotes sp., Photo by Tom Murray

The aquatic macrophyte Hydrilla verticilliata, Photo by FAO

The geographical distribution of body size

Trichoplax adhaerens

Loxodonta africana

Balaenoptera musculus

Neotrombicula autumnalis

Goliathus regius

Tinkerbella nana

Biogeographic distributions of invertebrate body sizes (Makarieva et al. 2005)

Makarieva, Gorshkov, Li 2005, Oikos 111: 425-436.

World distribution of largest land vertebrates

Mammals:Phytophages in tropical regionsPredators at higher latitudes

Birds:In tropical regions

Reptiles and Amphibians:In tropical regions

Largest species in

Kleiber’s rule

Hemmingson classic plot of metabolic rate against body size.

Each regression line has a slope of 3/4

𝑀∝𝑊 3 /40

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Slope

Obs

erva

tions

z Peters 1983

The speed of organismal metabolism is related to species body size by a power function.

Simple geometry tells that

The rule of Max Kleiber

The basic equation of metabolic theory

Adding the concentration of an assumed limiting resource Rmin gives

E = Activation energyBoltzmann factor: 8.314 Jmol-1K-1 = 0.0000862eVK-1

T = absolute temperatureW = body mass

𝑀∝𝑣

𝑀∝𝑊 3 /4𝑒− 𝐸𝑘𝑇

𝑀∝𝑊 3 /4

𝑀∝𝑅𝑚𝑖𝑛𝑊3 /4𝑒

− 𝐸𝑘𝑇

Brown et al. 2004, Ecology 85: 1771-1789

The Arrhenius equation of kinetic theory 𝑣 ∝𝑒

− 𝐸𝑘𝑇

The rate of DNA evolution predicted from metabolic theory

3/ 4 E / kT 1/ 4 E / kTMM W e W eW

Body size specific metabolic rate M/W should scale to the quarter power to body weight

and exponentially to temperature.

Now assume that most mutations are neutral and occur randomly. That is we assume that the neutral theory of

population genetics (Kimura 1983)

DNA substitution rate a should be proportional to M/W

1/ 4 E / kTM / W W ea a

• Body weight corrected DNA substitution rates (evolution rates) should be a linear function of 1/T with slope –E/k = -7541.

• Higher environmental temperatures should lead to higher substitution rates (faster evolution).

• Body weight corrected DNA substitution rates (evolution rates) should decrease with body weight.

• Large bodied species should have lower substitution rates (slower evolution).

Population size

Extin

ction

rate

Speciation ratePopulation size

Body

size

Body size

Extin

ction

rate

Speciation rate

c

𝛼∝𝑆∝𝑒−7500 /𝑇

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1/T

S

S=e

z=-10005

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North American

trees

Costa Rican trees along an elevational gradient

North American

amphibians

Ecuadorian amphibians

Fish species richness Prosobranchia species richness Ectoparasites of marine teleosts

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z=-9160

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400600800

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S

z=-7170

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1/T

S

z=-8510

The energy equivalence rule

𝑀∝𝑊 3 /4 𝐷∝𝑊 − 𝑧

𝐵=𝑀𝐷∝𝑊 3 /4− 𝑧

If z = ¾Energy equivalence rule

Damuth’s rule

𝐵=𝑀𝐷=𝑐𝑜𝑛𝑠𝑡Hoste Thesis 2013

Soil animals of Kampinowski National Park

Local and regional species richness

• Species richness on bracken is higher at richer sites

• At species poorer sites there seem to be many empty niches

• Local habitats are not saturated with species

Bracken occurs whole over the world

Species numbers of phytophages on bracken differ

Is this difference an effect of competitive exclusion or do empty niches exist?

John H.Lawton

The common brush tail Possum Trichosurus vulpecula is at its

introduced sites often free of natural parasites. There are empty niches

Pteridium aquilinum

Cynipid gall wasps in Norh America (Cornell 1985) Lacutstrine fish in North America (Gaston 2000)

Relationship between local species richness and the regional species pool size for 14 vegetation types in Estonia (Pärtel et al. 1996)

Dry grasslands Moist grasslands

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Local and regional species richness

Numbers of species incidences among sitesThe spatial distribution of abundance

Karelian plant species (Linkola 1916)

Core species

Satellite species

Intermediate species

Core (resident, permanent) species are often• of regionally higher abundance• good competitors• Pronounced species interactions• have stable species interactions• have low abundance fluctuations• are K-selected species

Satellite (transient, tourist) species are often• of regionally lower abundance• worse competitors• Weak species interactions• have unstable species interactions• have higher abundance fluctuations• are r-selected species

Numbers of species incidences in timeThe temporal distribution of abundance

Importance of ecological interactions

British Channel fish species (Magurran, Henderson 2003)

Abundance rank order

Abundance rank order

Verberk et al. 2010, J. Anim. Ecol. 79: 589

Local abundance and regional distribution in pond macroinvertebrates

Habitat generalists

Habitat specialists

Habitat generalists

High colonisation

ability

Low extinction

Wide regional distribution

Larger local populations

Habitat specialists

Low colonisation

ability

Higher extinction

Narrow regional distribution

Smaller local populations

Feedback loop between local

abundance and regional

occupancy (distribution)

Habitat specialists have often locally higher abundances than habitat generalists.

Local abundance is often positively correlated to regional distribution