Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost)...

53
Michael Kearney [email protected] School of BioSciences, The University of Melbourne Mechanistic Niche Models

Transcript of Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost)...

Page 1: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

Michael Kearney [email protected] of BioSciences, The University of Melbourne

Mechanistic Niche Models

Page 2: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

Part I• What is a mechanistic niche model?• Thermodynamic basis to the niche• The importance of temperature• Heat budgets• Microclimates• Water budgets• Play with NicheMapRPart II• Connecting to the Dynamic Energy Budget• Play with NicheMapR• Inferring climatic constraints• Nutritional constraints

Mechanistic Niche Models

Biophysical Ecology

Page 3: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

fundamental niche

realized niche

temperature

hum

idity

What is a mechanistic niche model?

Page 4: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

What is a mechanistic niche model?

Maxent model

NicheMapR model

Environmental Layers

probability of occurrence

potential reproduction

Correlative Model (process implicit)

Mechanistic Model (process explicit)

Page 5: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

51°C

35°C44°C

Military Dragon, Ctenophorus isolepis

Thermodynamic basis to the niche

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51°C

35°C44°C

Thermodynamic basis to the niche

Credit: Elia Pirtle

BOUNDARY

SYSTEM

SURROUNDINGSin

out

energy in = energy out + energy stored

in

outmass in = mass out + mass stored

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C6H12O6 + 6 O2 → 6 CO2 + 6 H2O heat

pSOLAR

pIR,in

p

MET

pIR,out

pCONV

p

pCOND

EVAP

+

+

+

+

+

fe dingJJX,I JX,PJX,G JX,R JX,S+ + + +=

brathnig

JO2,MET

JCO2,MET

JN,MET

JH,MET

+

+

+

= wa er

JJH,PJH,I +=+ +JH,U JH,S+

p = heat fluxJ = mass fluxX = foodH = waterI = ingestedP = product (faeces)U = urinatedG = growthR = reproductionS = storedO2 = oxygenCO2 = carbon dioxideN = nitrogenous wasteMET = ‘metabolism’EVAP = evaporationSOLAR = solar radiationIR = infrared radiationCONV = convectionCOND = conduction

Kearney et al. Functional Ecology (2013) after Porter and Tracy (1983)

pS+

Thermodynamic basis to the niche

Credit: Elia Pirtle

Page 8: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

Temperature ºC5 15 20 30 50100 45403525

Fitn

ess

survival survivaldeath deathreproduction

optimal

e.g. sprint speedegg developmentfeeding ratedigestion rate

The importance of temperature

Page 9: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

Magnet

Reed Switch

Light Sensor on Satellite

Board

Data logger Radio Transmitter

The ‘waddleometer’

Prof. Mike Bull

The importance of temperature

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60 Lizards’ Tbs in 2009

The importance of temperature

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Tfish = water temperature

Tball ∝ air temperature, wind speed, direct and diffuse solar radiation, infra red radiation, ground temperature, humidity, surface area, shape, thermal mass, solar reflectance …

The importance of temperature

Page 12: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

C6H12O6 + 6 O2 → 6 CO2 + 6 H2O heat

pSOLAR

pIR,in

p

MET

pIR,out

pCONV

p

pCOND

EVAP

+

+

+

+

+

fe ding

JJX,I JX,PJX,G JX,R JX,S+ + + +=br

athnigJO2,MET

JCO2,MET

JN,MET

JH,MET

+

+

+

= wa er

JJH,PJH,I +=+ +JH,U JH,S+

p = heat fluxJ = mass fluxX = foodH = waterI = ingestedP = product (faeces)U = urinatedG = growthR = reproductionS = storedO2 = oxygenCO2 = carbon dioxideN = nitrogenous wasteMET = ‘metabolism’EVAP = evaporationSOLAR = solar radiationIR = infrared radiationCONV = convectionCOND = conduction

Kearney et al. Functional Ecology (2013) after Porter and Tracy (1983)

pS+

Thermodynamic basis to the niche

Credit: Elia Pirtle

Page 13: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

Warren PorterUniversity of Wisconsin,

Madison

Ecological Monographs 39(3), 227-244 (1969)

Biophysical Ecology

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Solar(gained)

Infra-red(gained)

Conduction(gained/lost)

Convection(gained/lost)

Evaporation(lost)

Metabolism(gained)

Infra-red(lost)

+ +

+ + +

=

(Heat) Energy Balance of a Lizard

0][]273[ 4 =−−+− abcba TThTQ σ

air temperaturewind speed V,

organism size Dinfra-red and solar radiation

gained

infra-red radiationlost

convection

5.0

5.0

49.3DVhc =

Biophysical Ecology – Heat Budgets

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What would the body temperature be if …?

Diameter = 0.015 m

Wind speed = 2.0 m/s

0][49.3]273[ 5.0

5.04 =−−+− abba TT

DVTQ σ

Air temperature = 20 ºC

0]20[015.00.249.3]273[700 5.0

5.04 =−−+− bb TTσ

Radiation = 700 W/m2

If we know the environmental conditions, we can find the body temperature which satisfies the energy balance equation

Tb = 26 ºC

Biophysical Ecology – Heat Budgets

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weather station

air temperature layer

?

Biophysical Ecology – Microclimates

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56545250484644424038363432

http://img1.jurko.net/

Biophysical Ecology – Microclimates

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60 Lizards’ Tbs in 2009

Biophysical Ecology – Heat Budgets

Page 19: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

60 Lizards’ Tbs in 2009

Biophysical Ecology – Heat Budgets

Page 20: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

available range

body temperature

Environmental Layers

Thermodynamic basis to the niche

target (optimal) range

survivable range

selected environments through time

available range

hum

idity

Biophysical Ecology

Page 21: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

C6H12O6 + 6 O2 → 6 CO2 + 6 H2O heat

pSOLAR

pIR,in

p

MET

pIR,out

pCONV

p

pCOND

EVAP

+

+

+

+

+

fe ding

JJX,I JX,PJX,G JX,R JX,S+ + + +=br

athnigJO2,MET

JCO2,MET

JN,MET

JH,MET

+

+

+

= wa er

JJH,PJH,I +=+ +JH,U JH,S+

p = heat fluxJ = mass fluxX = foodH = waterI = ingestedP = product (faeces)U = urinatedG = growthR = reproductionS = storedO2 = oxygenCO2 = carbon dioxideN = nitrogenous wasteMET = ‘metabolism’EVAP = evaporationSOLAR = solar radiationIR = infrared radiationCONV = convectionCOND = conduction

Kearney et al. Functional Ecology (2013) after Porter and Tracy (1983)

pS+

Thermodynamic basis to the niche

Credit: Elia Pirtle

Page 22: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

Qevap,cut = Aevaphd (Vd,skin - Vd,air) λ

area wet, m2

mass transfer coefficient, m s-1

vapor density,kg m-3

latent heat of vaporisation,

J kg-1

Greer 1989 Biology and Evolution of Australian Lizards

Aevap

Water budget - evaporation

Qevap,resp = λ(mout,resp – min,resp)

mout

min

Page 23: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

C6H12O6 + 6 O2 → 6 CO2 + 6 H2O heat

pSOLAR

pIR,in

p

MET

pIR,out

pCONV

p

pCOND

EVAP

+

+

+

+

+

fe ding

JJX,I JX,PJX,G JX,R JX,S+ + + +=br

athnigJO2,MET

JCO2,MET

JN,MET

JH,MET

+

+

+

= wa er

JJH,PJH,I +=+ +JH,U JH,S+

p = heat fluxJ = mass fluxX = foodH = waterI = ingestedP = product (faeces)U = urinatedG = growthR = reproductionS = storedO2 = oxygenCO2 = carbon dioxideN = nitrogenous wasteMET = ‘metabolism’EVAP = evaporationSOLAR = solar radiationIR = infrared radiationCONV = convectionCOND = conduction

Kearney et al. Functional Ecology (2013) after Porter and Tracy (1983)

pS+

Thermodynamic basis to the niche

Credit: Elia Pirtle

Page 24: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

target (optimal) range

survivable range

selected environments through time

available range

hum

idity

Biophysical Ecology

body temperature

Environmental Layers

water balance

Thermodynamic basis to the niche

Page 25: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

Part I• What is a mechanistic niche model?• Thermodynamic basis to the niche• The importance of temperature• Heat budgets• Microclimates• Water budgets• Play with NicheMapR

Mechanistic Niche Models

Biophysical Ecology

Page 26: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

NicheMapR – a general system for mechanistic niche modelling

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NicheMapR Google Group

https://groups.google.com/forum/#!forum/nichemapr

@NicheMapR

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NicheMapR Google Group @NicheMapR

https://github.com/mrke/NicheMapR

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Niche Mapper

NicheMapR

Page 30: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

MICROCLIMATE

SOLRAD

SFODE IOSOLR

GAMMA

SINEC VSINE

DSUB

OSUB

EVALXZSOILPROPS

WETAIR

MICRO/MICROSEGMT

SNOWLAYER

HUMID

INFIL

DRYAIR

DCHXY

EVAP

DEXPITAB

VAPPRS

f) derived environmental variables d) integration routines

e) soil moisture and snow

b) solar radiation routines

c) hourly interpolation of daily inputs

a) driving subroutine, links to R via function microrun

Microclimate Program Structure

Niche Mapper NicheMapR

Page 31: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

-100 0 100

-50

050

microrun.R

rungads

micro_global.R

Fortran microclimate model library

Fortran GADS libraryGADS Tables

cCampNormTbl9_1

global_climate.nc

ccccc

metout

soil

soilmoist

soilpot

humid

soil.mon.ltm.v2.nc

cUser Input (location,

terrain, shade, soil type, …)

b)

a) Call to Fortran

c) User arguments

d) Climate and soil data setse) Global Aerosol Data Set

f) output

-150 -100 -50 0 50 100 150

-50

050

New et al. 2002

CPC soil moistureNicheMapR

Koepke et al 1997

Page 32: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

microrun.R

rungads

micro_aust.R

Fortran microclimate model library

Fortran GADS libraryGADS Tables

AWAP tmax, tmin, solr vpr, rr

ccccc

metout

soil

soilmoist

soilpot

humid

SLGA bulk dens, sand/silt/clay

cUser Input (location,

terrain, shade, soil type, …)

b)

a) Call to Fortran

c) User arguments

d) Climate, terrain and soil data setse) Global Aerosol Data Set

f) output

NicheMapR

Koepke et al 1997

McVicarwind

GEODATA 9 Second DEM

Page 33: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

50 100 150 200 250 300 350

05

1015

20

Day of Year

Hou

r of D

ay

The Heat and Water Budget Model

ccccc

metout

soil

soilmoist

soilpot

humid

output

ectotherm.Rccc

environ

enbal

masbal

SURROUNDINGS

BOUNDARY

wa er

brathnig

fe ding

Page 34: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

The Heat and Water Budget Model

c) Steadystateheat

budget

THERMOREG

f) thermoregulation

GEOM

d) heat (and water) budget components

METRESPSEVAP

SOLARABOVEGRND

BELOWGRND

BURROWIN

ECTOTHERM

b) driving subroutinelinks to R via function ectorun

SELDEP

CONDRADOUTCONVRADIN

SHADEADJUST

WETAIRDRYAIR

VAPPRSZBRAC ZBRENT

FUN

WATER

ccccc

shadmetshadsoil

shadmoistshadpot

shadhumid

ccccc

metoutsoil

soilmoistsoilpothumid

cccc

yearsout1yearsout2yearsout3

yearsout…n

ccccc

environenbal

masbaldeboutyearout

traits

morphologybehaviourphysiologyDEB pars

TRAPHRPHOTOPERIOD

e) Fluid properties

a) functional traits and microclimate datah) microclimate input

The Heat and Water Budget Model

Page 35: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

C6H12O6 + 6 O2 → 6 CO2 + 6 H2O heat

pSOLAR

pIR,in

p

MET

pIR,out

pCONV

p

pCOND

EVAP

+

+

+

+

+

fe ding

JJX,I JX,PJX,G JX,R JX,S+ + + +=br

athnigJO2,MET

JCO2,MET

JN,MET

JH,MET

+

+

+

= wa er

JJH,PJH,I +=+ +JH,U JH,S+

p = heat fluxJ = mass fluxX = foodH = waterI = ingestedP = product (faeces)U = urinatedG = growthR = reproductionS = storedO2 = oxygenCO2 = carbon dioxideN = nitrogenous wasteMET = ‘metabolism’EVAP = evaporationSOLAR = solar radiationIR = infrared radiationCONV = convectionCOND = conduction

Kearney et al. Functional Ecology (2013) after Porter and Tracy (1983)

pS+

Thermodynamic basis to the niche

Credit: Elia Pirtle

Page 36: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

TNZ

Thermo-Neutral

zone

5 15 20 30 50100 45403525Temperature ºC

Requ

irem

ents

Upper criticaltemperature (UCT)

Lower criticaltemperature (LCT)

Tcore = 36 ºC

Biophysical Ecology – Heat Budgets

Page 37: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

0

1

2

3

4

5

-10 0 10 20 30 40chamber temperature

met

abol

ic ra

te (W

)

23.1 31.1

C

woodrat

weasel

Brown J.H. & Lasiewski R.C. (1972). Ecology, 53, 939-943.

Biophysical Ecology – Heat Budgets

Page 38: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

consider an elliptical animal

Predicting Endotherm Energy and Water Requirements

Page 39: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

flesh

furwind

air

heat energy balance of an endotherm

body resistance furresistance

environmental resistance

Porter & Kearney (2009) PNAS 106, 19666-19672.

C

R

Predicting Endotherm Energy and Water Requirements

Page 40: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

0

1

2

3

4

5

-10 0 10 20 30 40chamber temperature

met

abol

ic ra

te (W

)

0

1

2

3

4

5

-10 0 10 20 30 40chamber temperature

met

abol

ic ra

te (W

)

23.1 31.1

C

22.5 31.7woodrat

weasel

Porter & Kearney (2009) PNAS 106, 19666-19672.

Predicting Endotherm Energy and Water Requirements

Page 41: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

NicheMapR – a general system for mechanistic niche modelling

Page 42: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

Part I• What is a mechanistic niche model?• Thermodynamic basis to the niche• The importance of temperature• Heat budgets• Microclimates• Water budgets• Play with NicheMapRPart II• Connecting to the Dynamic Energy Budget• Play with NicheMapR• Inferring climatic constraints• Nutritional constraints

Mechanistic Niche Models

Biophysical Ecology

Page 43: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

C6H12O6 + 6 O2 → 6 CO2 + 6 H2O heat

pSOLAR

pIR,in

p

MET

pIR,out

pCONV

p

pCOND

EVAP

+

+

+

+

+

fe ding

JJX,I JX,PJX,G JX,R JX,S+ + + +=br

athnigJO2,MET

JCO2,MET

JN,MET

JH,MET

+

+

+

= wa er

JJH,PJH,I +=+ +JH,U JH,S+

p = heat fluxJ = mass fluxX = foodH = waterI = ingestedP = product (faeces)U = urinatedG = growthR = reproductionS = storedO2 = oxygenCO2 = carbon dioxideN = nitrogenous wasteMET = ‘metabolism’EVAP = evaporationSOLAR = solar radiationIR = infrared radiationCONV = convectionCOND = conduction

Kearney et al. Functional Ecology (2013) after Porter and Tracy (1983)

pS+

Thermodynamic basis to the niche

Credit: Elia Pirtle

Page 44: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

C6H12O6 + 6 O2 → 6 CO2 + 6 H2O heat

pSOLAR

pIR,in

p

MET

pIR,out

pCONV

p

pCOND

EVAP

+

+

+

+

+

fe ding

JJX,I JX,PJX,G JX,R JX,S+ + + +=br

athnigJO2,MET

JCO2,MET

JN,MET

JH,MET

+

+

+

= wa er

JJH,PJH,I +=+ +JH,U JH,S+

p = heat fluxJ = mass fluxX = foodH = waterI = ingestedP = product (faeces)U = urinatedG = growthR = reproductionS = storedO2 = oxygenCO2 = carbon dioxideN = nitrogenous wasteMET = ‘metabolism’EVAP = evaporationSOLAR = solar radiationIR = infrared radiationCONV = convectionCOND = conduction

Kearney et al. Functional Ecology (2013) after Porter and Tracy (1983)

pS+

Thermodynamic basis to the niche

Credit: Elia Pirtle

Page 45: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

target (optimal) range

survivable range

selected environments through time

available range

hum

idity

Biophysical Ecology

Dynamic Energy/Mass Budgetgrowth

reproductivemaintenance

reservemetabolicworkgonads

metabolicrate

structuresomatic

maintenance

maturity maintenance

κ

1-κ

body temp/metabolic rate

Environmental Layers

water balance

Connecting to the Dynamic Energy Budget

Page 46: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

A year in the life of the Fence Lizard in UtahConnecting to the Dynamic Energy Budget

Page 47: Mechanistic Niche Models - Sciencesconf.org€¦ · Infra-red (gained) Conduction (gained/lost) Convection (gained/lost) Evaporation (lost) Metabolism (gained) Infra-red (lost) +

0

100

200

300

400

500

600

0 5 10 15

wet

mas

s (g

)

time (years)

growth (mass) and reproduction

CO2 H2O O2 N waste

CO2H2OO2N waste

FoodStructureReserveFaeces

=

Inferring climatic constraints

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Inferring climatic constraints

Kearney et al.. 2018. Field tests of a general ectotherm niche model …. Ecological Monographs 88:672–693.

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target (optimal) range

survivable range

selected environments through time

available range

water balance

hum

idity

Biophysical Ecology

Dynamic Energy/Mass Budgetgrowth

reproductivemaintenance

reservemetabolicworkgonads

metabolicrate

structuresomatic

maintenance

maturity maintenance

κ

1-κ

body temp/metabolic rate

Kearney and Porter TREE 2006Kearney et al. PTRS 2010

target (optimal) range

survivable range

available food 1

available food 2

selected foods through time

Geometric Framework

food ingested

carb

ohyd

rate

Environmental Layers

Nutritional constraints

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Food

Faeces

Food

Structure

Reserve

Faeces

Nutritional constraints

iso221

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food ingested

pa

faeces

assimilation SUs

carbohydratereserve

κ (1-κ)pC

somaticmaintenance

maintenance SU

structure

growth SU

pG pS

somaticmaintenance

maintenance SU

structure

growth SU

pG pS

reproduction SU

food ingested

proteinreserve

carbohydratereserve

pA*(1-κP)

pa

κM κM (1-κM)(1-κM)pC pC

structure

pG pGpS pS

growth SU

pR pR

maintenance SU

somaticmaintenance

maturation/reproduction

rejection rejection

return flux

retu

rn fl

ux

faeces

assimilation SUs

somaticmaintenance

maintenance SU

pA*κP

maintenance SU

maturitymaintenance

maturitymaintenance

maintenance SU

pM pM

retu

rn fl

ux

κE

κ E

reserve

Kearney et al. PTRS 2010

pRpM

repro SUmaturity maint SU

maturity maintenance repro

Nutritional constraints

iso221

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Simpson et al. (2010). Trends in Ecology & Evolution, 25, 53-60.

Nutritional Landscape Agent Behaviour

high P:C

low P:C

• Preference behaviour/targets will change with ontogeny

Nutritional constraints

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CLIMATE

MICROCLIMATE

HEAT BUDGET WATER BUDGET

METABOLISM

LIFE HISTORY/PHENOLOGY

VITAL RATES BIOTIC EFFECTS

DEMOGRAPHY

DISTRIBUTION

ABUNDANCE

MECHANISTIC NICHE MODEL

Environment

Indi

vidu

al

PopulationCommunity

predation

competitionresources

Mechanistic Niche Models