bias due to autocorrelation where no one expects it: the ... bias due to autocorrelation where no...

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Bias due to autocorrelation where no one expects it: the relationship between community-weighted mean traits and environmental variables David Zelený Masaryk University, Brno, Czech Republic ISEC 2014, Montpellier, France

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  • Bias due to autocorrelation where no one expects it: the relationship between community-weighted mean traits

    and environmental variables

    David Zelený

    Masaryk University, Brno, Czech Republic

    ISEC 2014, Montpellier, France

  • Bias due to autocorrelation where no one expects it: the relationship between community-weighted mean traits

    and environmental variables

    David Zelený

    Masaryk University, Brno, Czech Republic

    ISEC 2014, Montpellier, France

  • Weighted mean of species attributes

    David Zelený: Compositional autocorrelation and weighted mean 3

    L R

    Q’

    species

    samples

    species attributes

    sample attributes

    R ... matrix of sample attributes (e.g. environmental variables) L ... matrix of species composition Q ... matrix of species attributes (e.g. traits, species indicator values) (notation of matrices follow that in the RLQ and fourth-corner analysis)

  • Weighted mean of species attributes

    David Zelený: Compositional autocorrelation and weighted mean 4

    R ... matrix of sample attributes (e.g. environmental variables) L ... matrix of species composition (Ls ... standardized by sample totals) Q ... matrix of species attributes (e.g. traits, species indicator values) M ... matrix of weighted means of species attributes (e.g. CWM)

    Ls R

    Q’

    species

    samples

    species attributes

    sample attributes

    M M = LsQ

    m𝑛 = a𝑛𝑝 × t𝑝

    𝑛

    𝑝=1

    weighted mean of

    species attr.

    ~

  • Weighted mean of species attributes

    David Zelený: Compositional autocorrelation and weighted mean 5

    testing the relationship correlation, regression, ANOVA, parametric or nonparametric versions

    We should expect inflated Type I error rate

    Ls R

    Q’

    species

    samples

    species attributes

    sample attributes

    M M = LsQ

    m𝑛 = a𝑛𝑝 × t𝑝

    𝑛

    𝑝=1

    weighted mean of

    species attr.

  • Relationship between community weighted mean for SLA and cover of overstory

    David Zelený: Compositional autocorrelation and weighted mean 6

    20 40 60 80

    20

    30

    40

    50

    cover of trees and shrubs [%]

    weig

    hte

    d m

    ean o

    f S

    LA

    (herb

    s)

    r2 = 0.123, P < 0.001

    Data: • forest vegetation

    in Vltava valley • wide range of

    vegetation types • SLA for herb

    species • 97 samples, 233

    herb species

  • Weighted mean of species attributes: compositional autocorrelation

    • Samples with the same species composition will have the same value of WM

    • Samples with similar species composition will have similar value of WM

    David Zelený: Compositional autocorrelation and weighted mean 7

    0.4 0.6 0.8 1.0

    01

    02

    03

    04

    0

    dissimilarity (Whitt. index of association)

    CW

    M o

    f S

    LA

  • Weighted mean of species attributes: compositional autocorrelation

    • Samples with the same species composition will have the same value of WM

    • Samples with similar species composition will have similar value of WM

    David Zelený: Compositional autocorrelation and weighted mean 8

    -0.4 -0.2 0.0 0.2 0.4

    -0.4

    -0.2

    0.0

    0.2

    NMDS (Whittakers index of association)

    NMDS1

    NM

    DS

    2

  • Weighted mean of species attributes: compositional autocorrelation

    • Samples with the same species composition will have the same value of WM

    • Samples with similar species composition will have similar value of WM

    David Zelený: Compositional autocorrelation and weighted mean 9

    -0.4 -0.2 0.0 0.2 0.4

    -0.4

    -0.2

    0.0

    0.2

    NMDS1

    NM

    DS

    2CWM of SLA of herb species

    18

    20

    22

    24

    26

    28 30

    32

    34

    36

    38

    40

  • Relationship of two compositionally autocorrelated variables

    David Zelený: Compositional autocorrelation and weighted mean 10

    -0.4 -0.2 0.0 0.2 0.4

    -0.4

    -0.2

    0.0

    0.2

    NMDS1

    NM

    DS

    2

    CWM of SLA of herb species

    18

    20

    22

    24

    26

    28 30

    32

    34

    36

    38

    40

    -0.4 -0.2 0.0 0.2 0.4

    -0.4

    -0.2

    0.0

    0.2

    NMDS1

    NM

    DS

    2

    Cover of overstorey (trees and shrubs) [%]

    45 5

    0

    55

    60

    65

    70

    75

    80

    85

    20 40 60 80

    20

    30

    40

    50

    cover of trees and shrubs [%]

    weig

    hte

    d m

    ean o

    f S

    LA

    (herb

    s)

    r2 = 0.123, P < 0.001

    Issues: • inflated type I error rate • biased estimation of r or r2

  • The fourth-corner approach

    David Zelený: Compositional autocorrelation and weighted mean 11

    L R

    Q’

    species

    samples

    species attributes

    sample attributes

    R ... matrix of sample attributes (e.g. environmental variables) L ... matrix of species composition Q’ ... matrix of sample attributes (e.g. traits, species indicator values) D ... fourth-corner matrix

    D

  • The fourth-corner approach: testing the significance

    David Zelený: Compositional autocorrelation and weighted mean 12

    Legendre & Legendre (2012)

  • Solution for weighted-mean approach: modified permutation test

    • Modified permutation test: – null distribution of test statistic is created by calculating weighted

    mean from permuted species attributes

    – this test has correct type I error rate in case that there is a link between R and L (sample attributes and species composition matrix)

    – but it’s overly conservative in case that there is no R L link

    – possible to calculate in R package weimea (available on r-forge, beta version)

    David Zelený: Compositional autocorrelation and weighted mean 13

  • Solution for weighted-mean approach: modified permutation test

    David Zelený: Compositional autocorrelation and weighted mean 14

  • MoPeT: program (R-based) for calculation of modified permutation test

    David Zelený: Compositional autocorrelation and weighted mean 15

    www.bit.ly/ellenberg

    http://www.bit.ly/ellenberg

  • Conclusions

    • test of the relationship between weighted mean of species attributes and sample attributes has inflated Type I error rate

    • applies to community weighted mean of species functional traits, mean Ellenberg indicator values, diatom index etc.

    • solution: modified permutation test (based on permuting species instead of sample attributes)

    Thank you for your attention!

    David Zelený: Compositional autocorrelation and weighted mean 16