distributions in space
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Distributions in space. Biogeography Tries to understand large scale distributions of living thinks. Evolutionary Ecology Tries to understand patterns of species diversity through evolutionary history. Macroecology - PowerPoint PPT PresentationTRANSCRIPT
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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Area [miles 2 ]
Num
ber o
f spe
cies
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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Num
ber o
f spe
cies
A
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0 50 100 150Area
Num
ber o
f spe
cies
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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1.0E-01 1.0E+01 1.0E+03 1.0E+05 1.0E+07 1.0E+09 1.0E+11 1.0E+13
Area [Acres]
Num
ber o
f spe
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
f spe
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
cies
rich
ness
z
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Mean temperature
Spe
cies
rich
ness
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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S
z=-8540
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S
z=-10250
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1/T
S
z=-10810
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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1/T
S
z=-9160
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400600800
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0.0032 0.0033 0.0034 0.0035 0.0036 0.0037
1/T
S
z=-7170
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25
0.0033 0.0034 0.0035 0.0036 0.0037
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