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Avaliando padrões emergentes e processos ecológicos por meio de simulação de movimentos de aves em paisagens fragmentadas MILTON CEZAR RIBEIRO [email protected] Referata biodiversa INPE, Julho 2010

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Avaliando padrões emergentes e processos ecológicos por meio

de simulação de movimentos de aves em paisagens fragmentadas

MILTON CEZAR [email protected]

Referata biodiversaINPE, Julho 2010

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Assessing simulated bird species responses to landscape structure:spatial explicit IBM

RIBEIRO * FORTIN * METZGER et al., in reviewUofT-CANADA * USP-BRAZIL

MILTON RIBEIRO – [email protected]

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STUDY BACKGROUND:• Habitat Loss and Fragmentation

• Negative effect on Biodiversity

HABITAT MATRIX

Habitat loss WITHOUT Fragmentation

Habitat loss WITH Fragmentation

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STUDY BACKGROUND:• Fragmentation Threshold:

10-30% of habitat (Andrén 1994; Fahrig 2003)

Radford et al (2005)

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STUDY BACKGROUND:

• Brazilian Atlantic Forest : 12% of remaining habitat

• ~90% of habitat for forest species persistence was lost

• How species respond to habitat configuration?

• How habitat qualityinfluences speciesdistribution?

Ribeiro et al 2009

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OBJECTIVES:

• What influences the response patterns of simulated species?

Habitat AMOUNT * Habitat QUALITY * AGGREGATION

at patch level at landscape level

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APPROACH:

• Individual Based Models• Using generic species profiles that we setup for

the Atlantic Forest bird species

• What influences individuals to move/disperse?• Habitat quality• Degree of crowdingness

• Why Individual Based Models?• Generate testable hyphothesis• Check if expected patterns occurs

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APPROACH:

• Individual Based Model

• When individual are dispersing:

• The mortality rate is high on matrix

• Change the movement pattern• Routine maximum distance < explorative

distance

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APPROACH:

• Individual Based Model

• “Cost” to move:

• Depends on position of individual• Interior of patch -> low cost• edge proximity -> medium cost• Matrix -> high cost

• Habitat quality:• High quality -> low cost• Medium quality -> medium cost• Low quality -> high cost

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Simulated landscapes (n=10,000)

• Habitat amount: from 5 to 70%

• Aggregation: from 0.1 (random) to 1.0 (clumped)

• Habitat Quality: from low (0.1) to high (1.0)

METHODS

Continuous variables – Markov Chain Monte Carlo - MCMC

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High Quality Medium Quality Low Quality/Matrix0.7 – 1.0 0.4 – 0.7 0.1 to 0.4

Simulated Landscape: 512x512 – resolution 30m

Continuous variables – Markov Chain Monte Carlo - MCMC

AMOUNT=60% AGGREG=0.85

AMOUNT=42% AGGREG=0.47

AMOUNT=25% AGGREG=0.32

AMOUNT=12% AGGREG=0.21

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Simulated Landscape: 512x512 – resolution 30m

Continuous variables – Markov Chain Monte Carlo - MCMC

AMOUNT=55% AGGREG=0.92

AMOUNT=42% AGGREG=0.47

AMOUNT=25% AGGREG=0.32

AMOUNT=12% AGGREG=0.21

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EDGE SENSITIVE HABITAT-DEPEND. – DON’T USE CORRIDOR

EDGE AND CORRIDOR TOLERANT – USE CORRIDOR

GAP CROSSING OF 240M

Species profiles:

X

GAP CROSSING OF 60M

GAP CROSSING OF 120MEDGE TOLERANT-DON’T USE CORRIDOR

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120 M

GAPCROSSING

PROBABILITY

Gap Crossing - Awade & Metzger 2008 (understory birds)

Method = Play backThamnophilus caerulescens

HABITAT MATRIX

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Response to Edge – Hansbauer et al 2008 (understory birds)

Method = Telemetry

DISTANCE TO EDGE (m)

PreferenceIndex

>200 M

C. caudata

MATRIX HABITAT

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Home-range & use of Non-Forest habitatsHansbauer et al 2008 (understory birds) -Method = Telemetry

C. caudata

MATRIX = agriculture; pasture; rural building and silviculture

500 m

Habitat

Matrix

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Starting

population size =

Habitat amount/HR

Define position

Start simulation

Map database:

Landscape +

auxiliary maps

Species profile: HR,

routine distance,

dispersing distance

landscape

FOR time step=1 to N

Overcrowding on patch?

Functional patches

Habitat qualityDistance to edge/patch

change to disperser

movement state

yes

no

ALIVE?

Not move

no

Dispersing?

yes

Follow

disperser rule

Follow routine rule

noyes

Estimate prob.

of death, movement

cost

New position

Summary for landscape

Dispersal rate Mortality rateMovement costMovement variability

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RESPONSE VARIABLES – *at landscape level*DISPERSAL RATE:

• % of individual dispersing after a simulation

MOVEMENT VARIABILITY:

MORTALITY RATE:• % of mortality after a simulation

MOVEMENT COST: 1

(position * habitat quality)

TOTAL LENGTH

Euclidean DISTANCE

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0.0

0.2

0.4

0.6

0.8

1.0

Distance from edge (m)

Sa

fen

ess

-50

0

-24

0

-18

0

-12

0

-90

-60

-30 0

30

60

90

12

0

18

0

24

0

50

0

Edge sensitive

Edge tolerant

Edge and corridor tolerant

Generalist low gap crossing

Generalist medium gap crossing

0.0

00

00

.00

10

0.0

02

00

.00

30

Distance from edge (m)

Mo

rta

lity p

rob

ab

ility

-50

0

-24

0

-18

0

-12

0

-90

-60

-30 0

30

60

90

12

0

18

0

24

0

50

0

Edge sensitive

Edge tolerant

Edge and corridor tolerant

Generalist low gap crossing

Generalist medium gap crossing

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Experimental Design

• Home range size (HR): 10 ha

HR=Surrogate for Carrying capacity, used to define Starting Population Size

• Time step per simulation: 500 movements

• Maximum ROUTINE movement distance: 30 m• maximum EXPLORATIVE movement distance: 150 m

• N replicates: 10,000 runs, for EDGE AND CORRIDOR TOLERANT species profile

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Expected Patterns:

HABITAT AMOUNT

HABITAT QUALITY AGGREGATION

Mortality

Dispersal rateMov. Variability

Movement cost

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Landscape Ecology And Conservation Lab - USP

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IBM simulations – time steps = 500Population size = 100

Edge sensitive Edge and corridor tolerant

Low mobility Medium mobility

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Amount of habitat (%)

Dis

pe

rsa

l ra

te (

%)

0

5

10

15

20

10 20 30 40 50 60

Landscape-response:Each points is one simulation (500 time steps)For one landscape, and for one species profile

RESULTS

Fitted by Generalized Additive Model (GAM)

Statistical Approach: Model Selection (AIC)

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DISPERSAL RATE

Amount of habitat (%)

Dis

pe

rsa

l ra

te (

%)

0

5

10

15

20

10 20 30 40 50 60

Aggregation

Dis

pe

rsa

l ra

te (

%)

0

5

10

15

20

20 40 60 80

Quality of landscape

Dis

pe

rsa

l ra

te (

%)

0

5

10

15

20

0.3 0.4 0.5 0.6

HABITAT AMOUNT AGGREGATION HABITAT QUALITY

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Amount of habitat (%)

Mo

rta

lity

ra

te (

%)

0

2

4

6

8

10 20 30 40 50 60

Agregation

Mo

rta

lity

ra

te (

%)

0

2

4

6

8

20 40 60 80

Quality of landscape

Mo

rta

lity

ra

te (

%)

0

2

4

6

8

0.3 0.4 0.5 0.6

MORTALITY RATE

HABITAT AMOUNT AGGREGATION HABITAT QUALITY

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Amount of habitat (%)

Mo

ve

me

nt co

st

50

100

150

200

250

300

10 20 30 40 50 60

Agregation

Mo

ve

me

nt co

st

50

100

150

200

250

300

20 40 60 80

Quality of landscape

Mo

ve

me

nt co

st

0

50

100

150

200

250

300

0.3 0.4 0.5 0.6

MOVEMENT COST

HABITAT AMOUNT AGGREGATION HABITAT QUALITY

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Amount of habitat (%)

Mo

ve

me

nt va

ria

bility

5

10

15

20

10 20 30 40 50 60

Agregation

Mo

ve

me

nt va

ria

bility

5

10

15

20

20 40 60 80

Quality of Landscape

Mo

ve

me

nt va

ria

bility

5

10

15

20

0.3 0.4 0.5 0.6

MOVEMENT VARIABILITY

HABITAT AMOUNT AGGREGATION HABITAT QUALITY

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Response variables Selected Models DAICc wAICc (wAIC)

Movement variability Amount of habitat+Aggregation 0.00 0.89 0.89

Dispersal rate Quality of habitat patches 0.00 0.52 0.80

Quality of landscape 2.48 0.15

Amount of habitat 2.83 0.13

Mortality rate Quality of habitat patches 0.00 0.24 0.69

Quality of landscape 0.15 0.23

Amount of habitat 0.18 0.22

Movement cost Amount of habitat+Aggregation 0.00 0.93 0.93

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Expected Patterns:

HABITAT AMOUNT

HABITAT QUALITY AGGREGATION

Mortality

Dispersal rateMov. Variability

Movement cost

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Observed Patterns:

HABITAT AMOUNT

HABITAT QUALITY AGGREGATION

Mov. VariabilityMovement cost

MortalityDispersal rate Mortality

Dispersal rateMov. Variability

Movement cost

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Future Work

• Other species profiles including those with gap-crossing capability of 60 and 120 m?(functional connectivity)

• Daily-maximum distances >30 m?

• When dispersing-maximum distances >150 m

• Larger HR sizes (>50 ha)? (Top-predators)

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Frame-work advantages• A completely continuous modelling

• Habitat amount, quality and aggregation• Individual position completely free (not pixel-

based)• Movement distances (daily or when dispersing):

continuous following a uniform distribution• Home-range size: fixed our variables• Perceptual range position, habitat quality and

mortality • Simulate different species profile, varying all

parameters that allow to adjust to species that can respond to different scales

• Fully spatial explicit response pattern, including density-dependence behaviour

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Untangling the effects of landscape spatial

structure, habitat quality, and species

traits on ecological processes

RIBEIRO * MARTENSEN* FORTIN * METZGER - in prepUofT-CANADA * USP-BRAZIL

MILTON RIBEIRO – [email protected]

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OUTLINE:• Study background and research motivation• Confounding Effects• BioDIM (Biologically scaled Dispersal Model)• Response patterns• Sensitivity analysis• Results across and within species• Summary

• Future developments for BioDIM

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STUDY BACKGROUND:• Biodiversity and ecossystem variability

• How to explain?• How much we can explain?

Background * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

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STUDY BACKGROUND:• Biodiversity and ecossystem variability

• How to explain?• How much we can explain?

Background * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

System variability

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STUDY BACKGROUND:• Biodiversity and ecossystem variability

• How to explain?• How much we can explain?

Background * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

System variability

Processes(responses)

Patterns(the effects)

DispersalMatingReproductionGene flowMortalityForest sucessionBiomass production

Habitat lossFragmentationPatch size distrib.IsolationHabitat qualityConnectivityLand-cover dynamic

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STUDY BACKGROUND:• Biodiversity and ecossystem variability

• How to explain?• How much we can explain?

Background * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

Processes(responses)

Patterns(the effects)

DispersalMatingReproductionGene flowMortalityForest sucessionBiomass production

Habitat lossFragmentationPatch size distrib.IsolationHabitat qualityConnectivityLand-cover dynamic

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Background * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

Full system variability

Pattern 1

Pattern 2

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Background * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

Pattern 1

Pattern 2

Processes(responses)

Patterns(the effects)

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STUDY BACKGROUND:• Habitat Loss and Fragmentation

• Negative effect on Biodiversity• Habitat quality are important for:

• Species persistence• Population density• Carrying capacity• Reproductive success

• Landscape structure• Influences species movement• Patch accessibility• Dispersal success• Meta-community maintenance

• Species traits• How individuals interact with landscape features• Edge sensitivity / Gap crossing capatity

Background * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

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OBJECTIVES:• Estimate the strength of influence for the three

groups of factors on species response patterns:

Landscape structureHabitat QualitySpecies traits

Background * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

Studies bottle neck:• Assess the influencing factors independently

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APPROACH:

• BioDIM (Individual Based Models)Biologically scaled Dispersal Model

• Using generic species profiles that we setup for the Atlantic Forest bird species

Background * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

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Species profiles:

Background * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

A

B

C

D

E

F

Binary landscape

EdgeTolerant

Gap crossing60 m

Edge sensitive

Edge and corridor tolerant

Gap crossing 120 m

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RESPONSE VARIABLES – *at landscape level*

DISPERSAL RATE: % of individual dispersing after a simulation

MOVEMENT VARIABILITY:

MORTALITY RATE: % of mortality after a simulation

ENCOUNTER RATE: number of enconters Male/Female

MOVEMENT COST: 1

(position * habitat quality)

TOTAL LENGTH

Euclidean DISTANCE

Background * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

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Background * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

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EXPERIMENTAL DESIGN:

Landscape structure:PLAND: Amount of habitat (5% to 70%)AGGREG: Aggregation of habitat (0.1 to 1.0)

Habitat quality:HQBIN: include quality on model: Yes/No?HQLAND: Habitat quality at landscape levelHQFRAG: Habitat quality at patch level

Species traits:SPECSENSIT: Species sensitivity to spatial heterogeneity

(i.e. Species profiles)HORANG: home-range size, varying continuously from 5 to 30 ha;ROUTDIST: maximum routine distance (30 to 90 m; uniform distribution)EXPLDIST: maximum explorative distance (1.0 and 3.0 times the

ROUTDIST, i.e. 90 and 270 m; uniform distribution)

10 000 replicates / 500 time steps

Background * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

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SENSITIVITY ANALYSIS (Saltelli et al. 2008):

Standardized Regression Coefficient:SRC = -1 to +1 & +Std. Error

sensitivity package; Pujol 2008

Background * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

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SENSITIVITY ANALYSIS (Saltelli et al. 2008):

Standardized Regression Coefficient:SRC = -1 to +1 & +Std. Error

sensitivity package; Pujol 2008

Background * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

Positive effect Negative effect

+++

Or

- - -

++

Or

- -

+

Or

-

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RESULTSBackground * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

Positive effect

Negative effect

Including all species

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RESULTSBackground * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

Response variables

Explanatory

variables All species

Edge-

sensitive

Edge-

tolerant

Edge- and

corridor-

tolerant

Low gap-

crossing

Medium

gap-crossing Observation

Dispersal rate SPECSENSIT + + +

PLAND - - - - - - - - - - - always present

AGGREG - - - - - - - - - - - - + strong/trend/always present

HQBIN 0 0 0 0 0 +

HQLAND 0 0 + - 0 - - partial

HQFRAG 0 0 - 0 0 +

HORANG 0 - - 0 + + trend/partial

ROUTDIST + + + + + + + + 0 trend/partial

EXPLDIST 0 0 0 - - - partial

Movement variability SPECSENSIT 0

PLAND + - + + 0 0 + + partial

AGGREG + 0 0 + + + partial

HQBIN - - - - - 0 partial

HQLAND + + + 0 0 0 0

HQFRAG - - - - - 0 partial

HORANG 0 - - 0 0 0

ROUTDIST 0 - 0 - 0 - partial

EXPLDIST + + + + + + + 0 trend/partial

SPECSENSIT 0 0 0 0 0 0

Movement Cost SPECSENSIT ++

PLAND - - - - - - - - - - - - - - - - - - strong/always present

AGGREG 0 - - - + + trend/always present

HQBIN + + 0 0 + + partial

HQLAND - - 0 - - - - partial

HQFRAG + + + 0 + + + partial

HORANG + + + + + + + + + + + + always present

ROUTDIST 0 0 0 0 0 0

EXPLDIST - 0 - - - - Partial

Within species

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RESULTSBackground * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

Response variables

Explanatory

variables All species

Edge

sensitive

Edge

tolerant

Edge and

corridor

tolerant

Low gap

crossing

Medium gap

crossing observation

Encounters rate SPECSENSIT + +

PLAND + + + + + + + + + + always present

AGGREG + + 0 + + + + + + + + partial

HQBIN 0 0 - 0 0 0

HQLAND 0 0 0 + 0 0

HQFRAG 0 0 - - 0 0

HORANG - - - - - - - - - - - - - - - - - - strong/always present

ROUTDIST 0 + 0 0 0 0

EXPLDIST 0 0 0 0 0 0

Mortality SPECSENSIT +++

PLAND - - - - - - - - - - - - - strong/always present

AGGREG - - - - - - - - - - - - - - - - strong/always present

HQBIN 0 0 + + 0 0

HQLAND - - 0 - - - 0 partial

HQFRAG + + 0 + + 0 partial

HORANG 0 0 0 0 0 0

ROUTDIST + + + + 0 + partial

EXPLDIST 0 0 0 0 0 0

Within species

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SUMMARYBackground * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

• Species sensitivity was the primary factor to explain DISPERSAL rate and MORTALITY

• Amount of habitat: primary factor for MOVEMENT COST

• Home-range size: primary factor for ENCOUNTER Rate

• Movement variability: no strong factor was detected

Including all species

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SUMMARYBackground * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

• Amount of habitat and aggregation were the secondary factor for DISPERSAL rate and MORTALITY

• Home-range size and Species sensitivity: secondary for MOVEMENT COST

• Species sensitivity : secondary factor for ENCOUNTER Rate ; amount and aggregation of habitat are of tertiary influence on ENCOUNTER RATE

• Habitat quality not influenced any response variables

Including all species

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ImplicationsBackground * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

• Important to define “conservation target”• Species? Taxon?

• Landscape spatial heterogeneity was always important, but its strength of influence varied greatly depending on “process of interest” (i.e. response pattern) and species sensitivity

• Although habitat quality are pointed out as important to population’ s maintenance, this type of information are hard to obtain, and are generally measured at patch scale, not at landscape scale

Within Species

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Future WorkFuture studiesBackground * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

• Ecological Thresholds • BioDIM & Landscape genetic• BioDIM & 3D• BioDIM & Continuous surface• Apply BioDIM for sampling issues:

landscape genetic

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Future WorkEcological thresholdsBackground * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

HABITAT AMOUNT

MO

VEM

ENT

CO

ST

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Future WorkEcological thresholdsBackground * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

HABITAT AGGREGATION

DIS

PER

SAL

RAT

E

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HABITAT AGGREGATION

DIS

PER

SAL

RAT

E

HABITAT AMOUNT

HA

BIT

AT A

GG

REG

ATI

ON

DISPERSAL RATE

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population

GENETICSUB-MODEL

Individuals

LOCI structure

Proximity perception (PROX m)

Sex (F/M) rate

Gene exchange rate

FOR individual =1 to

popsize

Is Female

Exit genetic module

No

yes

Dist < Prox mno

FOR other individual

yes

Other Is Male

yes

no

Add 1 to # meetings

Change LOCI struct for

potential offspring

Example of LOCI STRUCStart: [0,1,1,0,1,0,1] [1,1] [0,1,0,1,1] [0,0]

End: [0,1,0,0,1,1,1] [1,0] [1,1,0,1,1] [1,0]

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Population Size is the sameMovement pattern is the same

Routine daily distance = lower Routine daily distance = higher

Sampling issues on landscape genetic studies

Future studiesBackground * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

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Continuous landscapes

Future studiesBackground * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

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3D scaled landscapes: a challengeBackground * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

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3D scaled landscapes: a challengeBackground * Effects * BioDIM * Responses * Sensitivity * Results * Summary * Future

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680

700

720

740

760

780

800

-200 0 200 400 600 800 1000 1200 1400

Perfil1 Perfil2 Perfil3

Perfil4 Perfil5

Distance between paired fragments - m

Altitude (meters)

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700

710

720

730

740

750

760

770

780

790

-200 0 200 400 600 800 1000 1200 1400

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730

740

750

760

770

780

790

-200 0 200 400 600 800 1000 1200 1400

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Perfil3

735

740

745

750

755

760

765

770

775

780

-200 0 200 400 600 800 1000 1200 1400

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Acknowledgment:

Brownyn Rayfield, Josie Hughes, Ilona Naujokaitis-Lewis and Danilo Boscolo for fruitful discussion about ecological modelling and birds behaviour at Atlantic Forest

• National Council for Scientific and Technological Development (CNPq)• The State of São Paulo Research Foundation (FAPESP)• Petrobras Company

Landscape Ecology And Conservation Lab - USP