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Describing the habitat of Atlantic salmon parr (Salmo salar) using the common spatial dependence between fish abundance and environmental variables: validation and application of a new multi-scale method Guillaume Guénard – [email protected] Pierre Legendre Daniel Boisclair Département de sciences biologiques Université de Montréal

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Page 1: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Describing the habitat of Atlantic salmon parr(Salmo salar) using the common spatial dependence between fish abundance andenvironmental variables: validation and application of a new multi-scale method

Guillaume Guénard – [email protected]

Pierre LegendreDaniel Boisclair

Département de sciences biologiquesUniversité de Montréal

Page 2: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Atlantic salmon

• Anadromous migratory• Juveniles (parr) are stream dwelling• Most wild stocks are at risk in North

America

Page 3: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Atlantic salmon

• Anadromous migratory• Juveniles (parr) are stream dwelling• Most wild stocks are at risk in North

America

Deterioration of habitat (spawning and feeding)

Page 4: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

2 Objectives

1. Develop a method to integrate spatial information about environmental descriptors to improve habitat models

2. Build models that describe the habitat used by Atlantic salmon parr

Page 5: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Modeling fish abundance using multiple linear regression

Abundance = Descriptors · b

Observations:Sampling in space

Availability of information: GPS location systems, etc.

What about using this information for modelingpurposes?

Page 6: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

PCNM variables

1. Truncated matrix of distances among sites2. Correct the matrix (Euclidean)3. Number of PCNM variables = n – 14. Sine-shaped spatial variables5. Orthogonal to one another6. Each PCNM variable has a mean of 0

)1()1(2

−⋅+≈

niLn

Page 7: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

-0.10

-0.05

0.00

0.05

0.10

PC

NM

var

iabl

e va

lue

-0.10

-0.05

0.00

0.05

0.10

-0.10

-0.05

0.00

0.05

0.10

Position (km)

0 1 2 3 4 5 6-0.10

-0.05

0.00

0.05

0.10

Scale

6200 m

775 m

123 m

62 m

Page 8: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Basic idea

If both the fish abundance and a habitat descriptor show high correlation to a given PCNM variable:

1. They are likely to have a common spatial dependence at the scale described by this PCNM variable.

2. This descriptor is suitable for predicting fish abundance at this spatial scale.

Page 9: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Method

• Least-square codependence analysis withrespect to PCNM variables (matrix W)

][]'[][]'[][']['

,, xxxxyyyyxxyy

iiii

iixy −−⋅−−

−−= WWC W�

Cy,x,W is the vector of scale-dependant correlations between the fish abundances (y) and the habitat descriptor (x).

� : Hadmard product

Page 10: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Method

• Calculate a pivotal statistic to test the significance of each coefficient of vector C using permutations

( )( ) ( )�� −−−⋅−−−

−−−−=

ikkiskki

iisi

kkisisxy

xxxxyyyy

xxyypnt sk

2,,

2

,,,

2

]['][]['][

][']['1

ww

www

Page 11: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Permutation testing

Two permutation approaches are possible

• Permute y and x together (pairwise) with respect to W.

H0: variables y and x are correlated but their correlation is not spatially dependent.

• Permute y and x independently.H0: variables y and x are neither correlated nor

spatially dependent.

Page 12: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Corrections for multiple testing

1. Non-sequential Šidák correction of p-values

kpp )1(1' −−=

1)1(1' −−−−= stepkpp

2. Sequential Šidák correction of p-values

Page 13: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Stepwise testing procedure1. Compute all Cy,x,W coefficients.2. Sort them in decreasing order of absolute

values.3. Test the first or subsequent coefficient(s)

for significance.4. Use a correction for multiple testing in the

test of subsequent coefficients.

5. Repeat from step 3 as long as coefficients are significant. Stop when a non-significant coefficient is found.

Page 14: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Simulation results

Page 15: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Estimation of type I error rate of the test, using independent permutationsand sequential correction, for 3 commonly-used significance levels (α)

Number of observations along transect

0 20 40 60 80 100 120

Type

I er

ror r

ate

0.001

0.01

0.1

1

α = 0.1α = 0.05α = 0.01

χ2 = 7.8

Page 16: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Results of all simulations using independant pairs of randomvariables

7.8SequentialIndependent

16.9Non-sequentialIndependent

14.4SequentialPairwise

17.7Non-sequentialPairwise

χ²11CorrectionPermutation

Page 17: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Results

Simulating type II error rate, using pairs ofvariables with know common spatial dependence…

…remains to be done!

Page 18: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Predicting Salmon parr relative abundance

���

���

−⋅−−⋅−−

=]['

][]'[][]'[ˆ

,

,,,,,

kkis

xykkikkiiisks xx

xxxxyyyysk

wC

wy w

�=s

ksk ,ˆˆ yy �=k

kyy ˆˆ

Page 19: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

0 1 250 2 500 3 750 5 000625Meters

Application to habitat modeling

Sainte-Marguerite River, Québec

Study area

Page 20: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Sampling

1. Salmon parr counted by snorkling2. Mean water velocity and depth3. Substrate composition: 6 grain size classes4. Size of sampling unit in the river: 20 m

Page 21: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Results

Page 22: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Codependence spectrum between fish abundanceand mean water depth

Scale (m)

100 1000

Cab

unda

nce,

mea

n de

pth

-0.10

-0.05

0.00

0.05

0.10

Page 23: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Codependence spectrum between fish abundanceand m ean water velocity

Scale (m)

100 1000

Cab

unda

nce,

mea

n ve

loci

ty

-0.10

-0.05

0.00

0.05

0.10

Page 24: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Codependence spectrum between fish abundanceand percentage of fine substrates (0.5 - 4 m m )

Scale (m)

100 1000

Cab

unda

nce,

% fi

ne s

ubst

rate

s

-0.10

-0.05

0.00

0.05

0.10

Page 25: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Codependence spectrum between fish abundanceand percentage of gravels (4 - 32 m m )

Scale (m)

100 1000

Cab

unda

nce,

% g

rave

ls

-0.10

-0.05

0.00

0.05

0.10

Page 26: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Codependence spectrum between fish abundanceand percentage of pebbles (32 - 64 m m)

Scale (m)

100 1000

Cab

unda

nce,

% p

ebbl

es

-0.10

-0.05

0.00

0.05

0.10

Page 27: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Codependence spectrum between fish abundanceand percentage of small boulders (0.25 - 1m )

Scale (m)

100 1000

Cab

unda

nce,

% s

mal

l bou

lder

s

-0.10

-0.05

0.00

0.05

0.10

Page 28: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

O bserved and raw predicted fish abundance using m eanwater velocity, depth, substra te com position

Position a long river stretch (km )

0 1 2 3 4 5 6

Abu

ndan

ce (f

ish.

20m

-1 s

trech

)

0

2

4

6

8

10

12

14O bserved abundancePredicted abundance

Page 29: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Observed vs. predicted local fish abundance

Observed local abundance (fish.20m-1)

0 2 4 6 8 10 12 14

Pre

dict

ed lo

cal a

bunc

ance

(fis

h.20

m-1

)

0

2

4

6

8

10

12

14 R² using codependence = 26.4%

R² using multiple regression = 9.5%

Page 30: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Conclusions

1. The spatial distribution of an environmental descriptor also containsinformation.

2. This information can be used to improve prediction of Atlantic salmon parr distribution.

3. Further simulations have to be done to determine type II error rate and the statistical power of the method.

Page 31: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

The End

Page 32: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Supplementary explanations

� �

� �

= =

= =

−⋅−

−⋅⋅−⋅=

n

i

n

iii

n

i

n

iijiiji

xy

xxyy

xxyy

j

1 1

22

1 1,,

,,

)()(

)()( wwC w

( )( )���

���

�−⋅−⋅−= −

= =� � 1

,,,1 1

2,

2, ][')()(ˆ kkisxy

n

i

n

ikkiisks xxxxyy

skwCwy w �

�=ks

ks,

,ˆˆ yy

Page 33: Describing the habitat of Atlantic salmon parrbiol09.biol.umontreal.ca/ESA_SS/Guenard_et_al_talk.pdf · Observed vs. predicted local fish abundance Observed local abundance (fish.20m-1)

Supplementary explanations

][]'[][]'[

]['][',, xxxxyyyy

xxyy

iiii

ijijwxy j −−⋅−−

−−=

wwC

]['

][]'[][]'[]['

,

,,,,,

kkis

xykkikkiiiisks xx

xxxxyyyyyy sk

−⋅−−⋅−−

=−=w

Cwb w

���

���

−⋅−−⋅−−

==]['

][]'[][]'[ˆ

,

,,,,,,

kkis

wxykkikkiiiskssks xx

xxxxyyyysk

wC

wbwy

�=s

ksk ,ˆˆ yy �=k

kyy ˆˆ