Multivariate Analysis in EcologyI: Unconstrained Ordination
Jari Oksanen
Oulu
January 2016
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 1 / 103
Introduction What is Ordination?
Multivariate Analysis and Ordination
Basic ordination methods to simplify multivariate data into low dimensionalgraphics
Analysis of multivariate dependence and hypotheses
Analyses can be performed in R statistical software using vegan package andallies
Course homepage http://cc.oulu.fi/~jarioksa/opetus/metodi/
Vegan homepage https://github.com/vegandevs/vegan/
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 2 / 103
Introduction What is Ordination?
Outline
1 IntroductionWhat is Ordination?Gradient Analysis
2 Unconstrained OrdinationNMDSEigenvector MethodsPCACAGraphicsEnvironmental VariablesGradient Model and Ordination
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Introduction What is Ordination?
Why Ordination?
Nobody should want to make an ordination, but they are desperate withmultivariate data
Map multidimensional table into low-dimensional display
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Introduction What is Ordination?
Two Ways of Analysing Data
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Introduction Gradient Analysis
Gradient Analysis
Gradient Analysis developed in 1950s in USA, with R. H. Whittaker as themain founding father
Only two or three environmental variables, or Gradients needed to explaincomplicated community patterns
Against classification: Species responses smooth along gradients
Against organism analogies: Species responses individualistic
The basis of modern theory and praxis: Ordination and Gradient modelling ofcommunities
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 6 / 103
Introduction Gradient Analysis
The Gradient ModelR.H.Whittaker (1956) Vegetation of The GreatSmoky Mountains. Ecological Monographs 26, 1–80.
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Introduction Gradient Analysis
Types of Gradients
1 Direct gradients: Influence organims but are not consumed.
Correspond to conditions.
2 Resource gradients: Consumed
Correspond to resources.
Complex gradients. Covarying direct and/or resource gradients: Impossibleto separate effects of single gradients.
Most observed gradients.
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Introduction Gradient Analysis
Landscapes and Gradients
Landscape
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Introduction Gradient Analysis
Species responses
Species have non-linear responses along gradients.Often assumed to be Gaussian. . .
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Introduction Gradient Analysis
Gaussian Response Function
Gaussian Response Function has three in-terpretable parameters that define the ex-pected response µ along the gradient x
Location of the optimum u on thegradient x
Width of the response t in the unitsof gradient x
Height of the response h in theunits of response height µ +
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http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 11 / 103
Introduction Gradient Analysis
Dream of species packing
Species have Gaussian responses and divide the gradient optimally:
Equal heights h.
Equal widths t.
Evenly distributed optima u.
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Introduction Gradient Analysis
Evidence for Gaussian Responses
Whittaker reported a large numberof different response types
Only a small proportion weresymmetric, bell shaped responses
Still became the standard of ourtimes
Comparison of ordination methodsbased on simulation, and many ofthose use Gaussian responses
We need to use simulation becausethen we know the truth that shouldbe found
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Unconstrained Ordination
Ordination
Ordination maps multivariate data onto low dimensional displays: “Most datasets have 2.5 dimensions”
Gradients define vegetation: ordination tries to find the underlying gradients
Basic ordination uses only community composition: Indirect Gradient Analysis
Constrained ordination studies only the variation that can be explained by theavailable environmental variables: Often called Direct Gradient Analysis
Distinct flavours of tools:
Nonmetric MDS the most robust methodPCA duly despisedFlavours of Correspondence Analysis popularCanonical method: Constrained Correspondence Analysis
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 14 / 103
Unconstrained Ordination NMDS
Nonmetric Multidimensional Scaling
Rank-order relation with (1) community dissimilarities and (2) ordinationdistances: No specified form of regression, but the the best shape is foundfrom the data.
Non-linear regression can cope with non-linear species responses of variousshapes: Not dependent on Gaussian model.
Iterative solution: No guarantee of convergence.
Must be solved separately for each number of dimensions: A lowerdimensional solutions is not a subset of a higher, but each case is solvedindividually.
A test winner, and a natural choice. . .
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Unconstrained Ordination NMDS
From Ranks of Dissimilarities to Ordination Distances
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Ranks of observed dissimilarities:A B C D E
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Ordination distances:A B C D E
B 0.301C 0.539 0.303D 0.323 0.133 0.421E 0.615 0.414 0.612 0.307F 0.922 0.636 0.636 0.605 0.416
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Unconstrained Ordination NMDS
MDS is a map
MDS tries to draw a map usingdistance data.
MDS tries to find an underlyingconfiguration fromdissimilarities.
Only the configuration counts:
No origin, but only theconstellations.No axes or natural directions,but only a framework forpoints.
Map of Europe from road distances
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Unconstrained Ordination NMDS
Shepard Diagram
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Non−metric fit, R2 = 0.967Linear fit, R2 = 0.835
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 18 / 103
Unconstrained Ordination NMDS
Iterative Optimization
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Unconstrained Ordination NMDS
Recommended procedure
NMDS may be good, but its use needs special care: Not every NMDSautomatically is good
1 Use adequate dissimilarity indices: An adequate index gives a good rank-orderrelation between community dissimilarity and gradient distance.
2 No convergence guaranteed: Start with several random starts and inspectthose with lowest stress.
3 Satisfied only if minimum stress configurations are similar.
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 20 / 103
Unconstrained Ordination NMDS
metaMDS I
> vare.mds <- metaMDS(varespec)
Square root transformationWisconsin double standardizationRun 0 stress 0.184Run 1 stress 0.196Run 2 stress 0.185... procrustes: rmse 0.0494 max resid 0.158Run 3 stress 0.209Run 4 stress 0.215Run 5 stress 0.235Run 6 stress 0.196Run 7 stress 0.234Run 8 stress 0.196Run 9 stress 0.222Run 10 stress 0.185Run 11 stress 0.195Run 12 stress 0.229Run 13 stress 0.184... New best solution... procrustes: rmse 3.6e-05 max resid 0.000139*** Solution reached
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 21 / 103
Unconstrained Ordination NMDS
metaMDS II
> vare.mds
Call:metaMDS(comm = varespec)
global Multidimensional Scaling using monoMDS
Data: wisconsin(sqrt(varespec))Distance: bray
Dimensions: 2Stress: 0.184Stress type 1, weak tiesTwo convergent solutions found after 13 triesScaling: centring, PC rotation, halfchange scalingSpecies: expanded scores based on ‘wisconsin(sqrt(varespec))’
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Unconstrained Ordination NMDS
Plot metaMDS
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> plot(vare.mds)http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 23 / 103
Unconstrained Ordination NMDS
Numbers
Badness of fit measure stress is based on the residuals from the non-linearregression
A proportional measure in the range 0 (perfect) . . . 1 (desperate) related togoodness of fit measure 1 − R2
Random configuration typically ≈ 0.4 and 0 degenerateOften given in percents (but omitting the percent sign: 15 = 0.15, sincecannot be > 1)
Orientation, rotation, scale and origin of the coordinates (scores) areindeterminate: only the constellation matters
Vegan arbitrarily fixes some of these:
Axes are centred, but the origin has no special meaningAxes are rotated so that the first is the longest (technically: rotated toprincipal components)Axes are scaled so that one unit corresponds to halving of similarity from the“replicate similarity”The sign (direction) of the axes still undefined
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Unconstrained Ordination NMDS
Half-change Scaling in NMDS
Replicate similarity: dissimilarityat ordination distance = 0
Maximum dissimilarity = 1:nothing in common
Linear area of ordinationdistance – dissimilarity: 0 . . . 0.8
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Replicate dissimilarity
Half−change
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 25 / 103
Unconstrained Ordination NMDS
What happened in metaMDS?
1 Square root transformation and Wisconsin double standardization
2 Bray–Curtis dissimilarities
3 monoMDS with several random starts and stopping after finding two identicalminimum stress solutions
4 Solution rotated to PCs
5 Solution scaled to half-change units
6 Species scores as weighted averages
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Unconstrained Ordination NMDS
Dissimilarity measures
Use a dissimilarity that describescorrectly gradient separation
Bray–Curtis (Steinhaus), Jaccard,Kulczynski
Wisconsin double standardizationoften helpful
Should use dissimilarities whichreach their maximum (1) when nospecies are shared (like those listedabove)
Indices with no bound maximum areusually bad (Euclidean distance etc.)
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http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 27 / 103
Unconstrained Ordination NMDS
Procrustes rotation
Procrustes rotation to maximal similarity between two configurations:
Translate the origin.Rotate the axes.Deflate or inflate the axis scale.
Single points can move a lot, although the stress is fairly constant: Especiallyin large data sets.
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Unconstrained Ordination NMDS
Procrustes Rotation
> tmp <- wisconsin(sqrt(varespec))> dis <- vegdist(tmp)> vare.mds0 <- monoMDS(dis, trace = 0)> pro <- procrustes(vare.mds, vare.mds0)> pro
Call:procrustes(X = vare.mds, Y = vare.mds0)
Procrustes sum of squares:0.186
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Unconstrained Ordination NMDS
Plot Procrustes Rotations
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Unconstrained Ordination NMDS
Number of dimensions
In NMDS, 2D solution is not a plane in 3D space
Solution must be found separately for each dimensionality
Some people very disturbed: how do they know the correct number
Answer is easy: there is no correct number, although some numbers may beworse than others
“Most data sets have 2.5 dimensions”
Typically you try with 2 and 3
Do you need more dimensions to explain species patterns and environmentaldata?
Is convergence very slow? Try another number of dimensions
Scree plot or stress against the number of dimensions often suggested butrarely works
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Unconstrained Ordination NMDS
Scree Plot
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Unconstrained Ordination Eigenvector Methods
Simplified mapping: Eigen analysis
NMDS uses non-linear mapping for any dissimilarity measure: This is verydifficult
Things are much simpler if we accept only certain dissimilarity indices andmap them linearly onto ordination
Linear mapping is only a rotation, and can be solved using eigenvectortechniques
Sometimes said that certain methods are model-based (CA), but they alsoemploy a distance
method metric mapping
NMDS any nonlinearMDS any linearPCA Euclidean linearCA Chi-square weighted linear
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Unconstrained Ordination Eigenvector Methods
Why Not PCA?
We admit that PCA is just a rotation, but it is a linear method
PCA works with species space, but we boldly go to gradient space
CA is an optimal scaling method
Sites with similar species composition packed close to each otherSpecies that occur together simultaneously packed close to each other
CA can handle unimodal species responses, even approximate onedimensional species packing model
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Unconstrained Ordination PCA
Species space
Graphical presentations of datamatrix: Species are axes and spanthe space where sites are points
Some species show more of theconfiguration than others
What is the ideal viewing angle tothe species space?
Shows as much as possible of allspecies in just two or threedimensions 0 10 20 30 40 50 60
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http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 35 / 103
Unconstrained Ordination PCA
Rotation in species space
1 Put sites into species space
2 Move the origin to the centroid
3 Rotate the axes so that the first axis(1) is as close to all points aspossible, and (2) explains as muchof the variance as possible
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Unconstrained Ordination PCA
Goodness of Fit
The total variation (Λ) is the sum of squared distances of points from theorigin
Λ can be expressed as the sum of squares (SS) or variance (SS/n orSS/(n − 1))
The points are projected on the axis, and the sum of projected squareddistances is the eigenvalue of the axis (λi )
The eigenvalues are ordered and non-negative λ1 ≥ λ2 ≥ · · · ≥ λp ≥ 0, andsum up to total variance Λ =
∑pi=1 λi
λi/Λ gives the proportion that an axis explains of the total variance, and λ1
explains the largest proportion
Cumulative sum gives the proportion of variance explained by the first axes:often emphasized but rather useless statistic
PCA is often used to reduce data into a few linearly independent componentsthat explain the most of the original variables
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Unconstrained Ordination PCA
Euclidean Metric of PCA
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Unconstrained Ordination PCA
Running PCA I
> (ord <- rda(dune))
Call: rda(X = dune)
Inertia RankTotal 84.1Unconstrained 84.1 19Inertia is variance
Eigenvalues for unconstrained axes:PC1 PC2 PC3 PC4 PC5 PC6 PC7 PC8
24.80 18.15 7.63 7.15 5.70 4.33 3.20 2.78(Showed only 8 of all 19 unconstrained eigenvalues)
> head(summary(ord), 3, 1)
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Unconstrained Ordination PCA
Running PCA II
Call:rda(X = dune)
Partitioning of variance:Inertia Proportion
Total 84.1 1Unconstrained 84.1 1
Eigenvalues, and their contribution to the variance
Importance of components:PC1 PC2 PC3 PC4 PC5 PC6
Eigenvalue 24.795 18.147 7.6291 7.153 5.6950 4.3333Proportion Explained 0.295 0.216 0.0907 0.085 0.0677 0.0515Cumulative Proportion 0.295 0.510 0.6011 0.686 0.7539 0.8054
PC7 PC8 PC9 PC10 PC11 PC12Eigenvalue 3.199 2.7819 2.4820 1.854 1.7471 1.3136Proportion Explained 0.038 0.0331 0.0295 0.022 0.0208 0.0156Cumulative Proportion 0.843 0.8765 0.9060 0.928 0.9488 0.9644
PC13 PC14 PC15 PC16 PC17Eigenvalue 0.9905 0.63779 0.55083 0.35058 0.19956Proportion Explained 0.0118 0.00758 0.00655 0.00417 0.00237
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Unconstrained Ordination PCA
Running PCA III
Cumulative Proportion 0.9762 0.98377 0.99032 0.99448 0.99686PC18 PC19
Eigenvalue 0.14880 0.11575Proportion Explained 0.00177 0.00138Cumulative Proportion 0.99862 1.00000
Scaling 2 for species and site scores* Species are scaled proportional to eigenvalues* Sites are unscaled: weighted dispersion equal on all dimensions* General scaling constant of scores: 6.3229
Species scores
PC1 PC2 PC3 PC4 PC5 PC6Achimill -0.6038 0.124 0.00846 0.160 0.4087 0.1279Agrostol 1.3740 -0.964 0.16691 0.266 -0.0877 0.0474Airaprae 0.0234 0.251 -0.19477 -0.326 0.0557 -0.0796....Callcusp 0.5385 0.180 0.17509 0.239 0.2553 0.1692
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Unconstrained Ordination PCA
Running PCA IV
Site scores (weighted sums of species scores)
PC1 PC2 PC3 PC4 PC5 PC61 -0.857 -0.172 2.608 -1.130 0.4507 -2.49112 -1.645 -1.230 0.887 -0.986 2.0346 1.81063 -0.440 -2.383 0.930 -0.460 -1.0278 -0.0518....20 2.341 1.299 0.903 0.718 -0.0757 -0.9691
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Unconstrained Ordination PCA
Row and Column scores
The scores are centred (= their mean is zero) and either normalized (= allhave equal spread) or proportional to eigenvalues (= spread is higher wheneigenvalue is high)
Normalized scores give the regression coefficients between the axis and thevariables: often used for species
Scores proportional to the eigenvalue give the true configuration of points inthe space defined by normalized scores: often used for sites (hence in speciesspace)
Together these scores give a linear least square approximation of the data
Graphical presentation called biplot
However, there are many alternative scaling systems
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Unconstrained Ordination PCA
Default Plot
−2 −1 0 1 2 3
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PC1
PC
2
Achimill
Agrostol
Airaprae
Alopgeni
Anthodor
BellpereBromhordChenalbuCirsarve
ComapaluEleopalu
Elymrepe
Empenigr
Hyporadi
Juncarti
JuncbufoLolipere
Planlanc
Poaprat
Poatriv
RanuflamRumeacet
Sagiproc
SalirepeScorautu
Trifprat
TrifrepeVicilath
Bracruta
Callcusp
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Unconstrained Ordination PCA
Reading the Plot
Origin: all species (variables) at their average values
The distance from the origin for a row (site) implies how much the pointdiffers from the average
The distance from the origin for a column (species, variable) implies howmuch the point increases to that direction
The change is measured in absolute scale: big changes, long distances fromthe origin
Implies a linear model of species response against axes
The angle between two points implies correlations
90◦ means zero correlation, < 90◦ positive correlation, > 90◦ negativecorrelation, 0◦ implies r = 1
Arrow biplots often used instead of point biplot
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Unconstrained Ordination PCA
Arrow Biplot
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PC
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Achimill
Agrostol
Airaprae
Alopgeni
Anthodor
BellpereBromhordChenalbuCirsarve
ComapaluEleopalu
Elymrepe
Empenigr
Hyporadi
Juncarti
JuncbufoLolipere
Planlanc
Poaprat
Poatriv
RanuflamRumeacet
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Unconstrained Ordination PCA
Linear Model
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ecte
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Unconstrained Ordination PCA
Variances and Correlations
Analysis of raw data explains variances: variables with high variance are mostimportant
If the variables are standardized to unit variance before analysisz = (x − x)/sx all variables are equally important and the analysis explainscorrelations among variables
Standardization can be used when we want all variables to have equal weights
Standardization must be used when variables are measured in different scales,such as for environmental measurements
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Unconstrained Ordination PCA
Reducing the Number of Correlated EnvironmentalVariables I
> (pc <- rda(varechem, scale=TRUE))
Call: rda(X = varechem, scale = TRUE)
Inertia RankTotal 14Unconstrained 14 14Inertia is correlations
Eigenvalues for unconstrained axes:PC1 PC2 PC3 PC4 PC5 PC6 PC7 PC8 PC9 PC10 PC11 PC125.19 3.19 1.69 1.07 0.82 0.71 0.44 0.37 0.17 0.15 0.09 0.07PC13 PC140.04 0.02
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Unconstrained Ordination PCA
The Number of Components
PCA is a rotation in species (character) space and retains the originalconfiguration
The number of PC’s is min(N,S), and all together give the original data
First axes are most important and we may ignore the minor axes
We can either use the axes as variables in other models, or use them toidentify major (almost) independent variables
Often we want to retain a certain proportion of the variance, say 50 %
Sometimes we would like to retain “significant” axes
There really is no way of doing this, but some people suggest comparingeigenvalues against broken stick distribution
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Unconstrained Ordination PCA
Broken Stick and Eigenvalues
PC1 PC2 PC3 PC4 PC5 PC6 PC7 PC8 PC9
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Unconstrained Ordination PCA
Two Dimensions, but which?
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Unconstrained Ordination PCA
Methods Related to PCA
Metric Scaling a.k.a. Principal Coordinates Analysis
Used dissimilarities instead of raw dataWith Euclidean distances equal to PCA, but can use other dissimilarities
Factor Analysis
A statistical method that makes a difference between systematic componentsand random errorIn PCA we just ignore latter components, but here we really identify the realcomponentsMuch used in human sciences and often referred to in ecology (but usuallymisunderstood)
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Unconstrained Ordination PCA
Confirmatory Factor Analysis
P
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Mg
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http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 54 / 103
Unconstrained Ordination CA
Correspondence Analysis
Minor variant of PCA: Weighted PrincipalComponents with Chi-square metric
All sites should have all species in in the sameproportions as in the whole data
Site and species marginal profiles define theexpected abundances
Null model: Species composition is identical in allsampling units
Chi-square transformation tells how much theobserved proportions fij differ from the expectedproportions eij :
χij =fij − eij√
eij
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Cla.uncCla.cocCla.corCla.graCla.fimCla.criCla.chlCla.bot
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Unconstrained Ordination CA
Chi-squared metric
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Unconstrained Ordination CA
CA Rotation
1 Sites in a species space
2 Relative proportions are axes andpoints have weights
3 Chi-square transformation
4 Weighted rotation
5 De-weighting
CA1
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http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 57 / 103
Unconstrained Ordination CA
Running CA I
> (ord <- cca(dune))
Call: cca(X = dune)
Inertia RankTotal 2.12Unconstrained 2.12 19Inertia is mean squared contingency coefficient
Eigenvalues for unconstrained axes:CA1 CA2 CA3 CA4 CA5 CA6 CA7 CA8
0.536 0.400 0.260 0.176 0.145 0.108 0.092 0.081(Showed only 8 of all 19 unconstrained eigenvalues)
> head(summary(ord), 2, 1)
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 58 / 103
Unconstrained Ordination CA
Running CA II
Call:cca(X = dune)
Partitioning of mean squared contingency coefficient:Inertia Proportion
Total 2.12 1Unconstrained 2.12 1
Eigenvalues, and their contribution to the mean squared contingency coefficient
Importance of components:CA1 CA2 CA3 CA4 CA5 CA6
Eigenvalue 0.536 0.400 0.260 0.1760 0.1448 0.108Proportion Explained 0.253 0.189 0.123 0.0832 0.0684 0.051Cumulative Proportion 0.253 0.443 0.565 0.6486 0.7170 0.768
CA7 CA8 CA9 CA10 CA11Eigenvalue 0.0925 0.0809 0.0733 0.0563 0.0483Proportion Explained 0.0437 0.0382 0.0347 0.0266 0.0228Cumulative Proportion 0.8117 0.8500 0.8847 0.9113 0.9341
CA12 CA13 CA14 CA15 CA16Eigenvalue 0.0412 0.0352 0.02053 0.01491 0.00907Proportion Explained 0.0195 0.0167 0.00971 0.00705 0.00429
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 59 / 103
Unconstrained Ordination CA
Running CA III
Cumulative Proportion 0.9536 0.9702 0.97995 0.98700 0.99129CA17 CA18 CA19
Eigenvalue 0.00794 0.00700 0.00348Proportion Explained 0.00375 0.00331 0.00164Cumulative Proportion 0.99505 0.99836 1.00000
Scaling 2 for species and site scores* Species are scaled proportional to eigenvalues* Sites are unscaled: weighted dispersion equal on all dimensions
Species scores
CA1 CA2 CA3 CA4 CA5 CA6Achimill -0.909 0.0846 -0.586 -0.00892 -0.660 -0.1888Agrostol 0.934 -0.2065 0.282 0.02429 -0.139 -0.0226....Callcusp 1.952 0.5674 -0.859 -0.09897 -0.557 0.2328
Site scores (weighted averages of species scores)
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Unconstrained Ordination CA
Running CA IV
CA1 CA2 CA3 CA4 CA5 CA61 -0.812 -1.083 -0.1448 -2.107 -0.393 -1.83462 -0.633 -0.696 -0.0971 -1.187 -0.977 0.0658....20 1.944 1.069 -0.6660 -0.553 1.596 -1.7029
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Unconstrained Ordination CA
Goodness of Fit of Scores
Inertia is “mean square contingency coefficient”: Chi-squared of a matrixstandardized to unit sum, or Chi-square of x∑
x
Eigenvalues are non-negative and ordered like in PCA, but they are bound tomaximum 1
The origin gives the expected abundances for all species and all sites
The deviant species and deviant sites are far away from the origin
CA is weighted analysis, and the weighted sum of squared scores is theeigenvalue
The species and site scores are (scaled) weighted averages of each other:proximity matters
Rare species have low weights: they are further away from the origin
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Unconstrained Ordination CA
Weighted Average?
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For quantitative data: plots wheresspecies is abundant are heavier and theweighted average is closer to them
Sampling units (SU) are close to speciesthat occur on them
CA is a weighted average method: ittries to put SUs close to the speciesthat occur in them, and all SUs withsimilar species composition close to eachother: Unimodal response
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Unconstrained Ordination CA
Default Plot and Effect of Scaling
−2 −1 0 1 2 3 4
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Achimill
Agrostol
Airaprae
Alopgeni
Anthodor
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Eleopalu
Elymrepe
Empenigr
Hyporadi
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Juncbufo
Lolipere
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Poatriv
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Hyporadi
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Unconstrained Ordination CA
Weighted Averages
Species scores are [proportional to] weighted averages of site scores, andsimultaneously
Site scores are [proportional to] weighted averages of species scores
Either one (but not both) of these can be a direct weighted average of other
If sites scores are weighted averages of species scores, site point is in themiddle of points of species that occurs in the site
The location of the point is meaningful whereas in PCA the main things weredistance and direction from the origin (but these, too, matter)
Can approximate unimodal response model and therefore CA is much betterfor community ordination than PCA
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Unconstrained Ordination CA
Linear and Unimodal Models
PCA implies linear relations between axes and species abundances
CA packs species and approximates a unimodal model
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Unconstrained Ordination CA
Optimal Scaling
The locations of species optima (tops)should be widespread: spread ismeasured as SSB
The species responses should be narrow:width is measured as SSw
The total variance is their sumSST = SSB + SSw
High SSB means that species havedifferent optima, and low SSw meansthat species have narrow tolerance
Scaling is optimal if most of variance isbetween species and SSB is high
The criterion of variance is theeigenvalue maximized in CA:λ = SSB/SST
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Unconstrained Ordination CA
Goodness of Fit Statistics: Repetition
NMDS: stress of nonlinear transformation from observed dissimilarities toordination distances
In range 0 . . . 1 (0 . . . 100 %), but in practice 0.4 for random configuration0.1 is good, and 0.2 is not bad, 0 is suspect
PCA: sum of eigenvalues is variance (or SS)
Upper limit is total variance, large is good
CA: sum of all eigenvalues is (scaled) Chi-square
Single eigenvalue maximum 1high is good, but λ < 0.2 may not be badEigenvalues λ > 0.7 are suspect: disjunct or very heterogeneous data
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Unconstrained Ordination CA
Nonlinear and Linear Mapping
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Unconstrained Ordination CA
Nonlinear and Linear Mapping: A Difficult Case
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Unconstrained Ordination Graphics
Anatomy of a Plot
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Dicrpoly
Hylosple
PleuschrPolypili
PolyjuniPolycomm
Pohlnuta Ptilcili
Barbhatc
Cladarbu
Cladrang
Cladstel
Cladunci
Cladcocc
CladcornCladgrac
CladfimbCladcris
Cladchlo
Cladbotr
Cladamau
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Flavniva
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Dicrfusc
Dicrpoly
Hylosple
PleuschrPolypili
PolyjuniPolycomm
Pohlnuta Ptilcili
Barbhatc
Cladarbu
Cladrang
Cladstel
Cladunci
Cladcocc
CladcornCladgracCladfimb
Cladcris
Cladchlo
Cladbotr
Cladamau
Cladsp
Cetreric
Cetrisla
Flavniva
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Callvulg
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RhodtomeVaccmyrt
Vaccviti
Pinusylv
Descflex
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Vacculig
Diphcomp
Dicrsp
Dicrfusc
Dicrpoly
Hylosple
PleuschrPolypili
PolyjuniPolycomm
Pohlnuta Ptilcili
Barbhatc
Cladarbu
Cladrang
Cladstel
Cladunci
Cladcocc
CladcornCladgracCladfimb
Cladcris
Cladchlo
Cladbotr
Cladamau
Cladsp
Cetreric
Cetrisla
Flavniva
Nepharct
Stersp
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Icmaeric
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Descflex
Cladphyl
BetupubeFlavniva
Dicrpoly
PolycommStersp
Cladchlo
Cladbotr
Vacculig
Pinusylv
Polypili
Dicrfusc
Dicrsp
Ptilcili
Cladcocc
Cladrang
Pohlnuta
Pleuschr
Polyjuni
Cetreric
Empenigr
Cladsp
Cladunci
Peltapht
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Callvulg
Empenigr
RhodtomeVaccmyrt
Vaccviti
Pinusylv
Descflex
Betupube
Vacculig
Diphcomp
Dicrsp
Dicrfusc
Dicrpoly
Hylosple
PleuschrPolypili
PolyjuniPolycomm
Pohlnuta Ptilcili
Barbhatc
Cladarbu
Cladrang
Cladstel
Cladunci
Cladcocc
CladcornCladgracCladfimb
Cladcris
Cladchlo
Cladbotr
Cladamau
Cladsp
Cetreric
Cetrisla
Flavniva
Nepharct
Stersp
Peltapht
Icmaeric
Cladcerv
Claddefo
Cladphyl
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 71 / 103
Unconstrained Ordination Graphics
Plotting functions
All vegan ordination functions have a plot function, and ordiplot can beused for other functions as well
For full control, use first plot(x, type="n") and then add configurablepoints or text
Congested plots can displayed with orditorp or edited with orditkplot
Lattice graphics can be made with ordixyplot, ordicloud or ordisplom
Dynamic, spinnable 3D plots can be made with ordirgl function in thevegan3d package
Items can be added to the plots with ordiarrows, ordihull, ordispider,ordihull, ordiellipse, ordisegments, or ordigrid
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 72 / 103
Unconstrained Ordination Environmental Variables
Ordination and Environment
We take granted that vegetation is controlled by environment, so
1 Two sites close to each other in ordination have similar vegetation
2 If two sites have similar vegetation, they have similar environment
3 Two sites far away from each other in ordination have dissimilar vegetation,and perhaps
4 If two sites have different vegetation, they have different environment
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 73 / 103
Unconstrained Ordination Environmental Variables
Fitted Vectors
Direction of fitted vector shows thegradient of the environmental variable,length shows its importance.
For every arrow, there is an equally longarrow into opposite direction:Decreasing direction of the gradient.
Implies a linear model: Project sampleplots onto the vector for expected value.
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Partsize Currvel
slope
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widthInstab
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http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 74 / 103
Unconstrained Ordination Environmental Variables
Interpretation of Arrow
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http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 75 / 103
Unconstrained Ordination Environmental Variables
Alternatives to Vectors
Fitted vectors natural in constrained ordination, since these have linearconstraints.
Distant sites are different, but may be different in various ways:Environmental variables may have a non-linear relation to ordination.
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http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 76 / 103
Unconstrained Ordination Environmental Variables
Fitting Environmental Vectors I
> (ef <- envfit(vare.mds, varechem, permu = 999))
***VECTORS
NMDS1 NMDS2 r2 Pr(>r)N -0.050 -0.999 0.21 0.098 .P 0.687 0.727 0.18 0.135K 0.827 0.562 0.17 0.147Ca 0.750 0.661 0.28 0.029 *Mg 0.697 0.717 0.35 0.015 *S 0.276 0.961 0.18 0.143Al -0.838 0.546 0.52 0.002 **Fe -0.862 0.507 0.40 0.013 *Mn 0.802 -0.597 0.53 0.001 ***Zn 0.665 0.747 0.18 0.146Mo -0.849 0.529 0.05 0.581Baresoil 0.872 -0.490 0.25 0.035 *Humdepth 0.926 -0.377 0.56 0.001 ***pH -0.799 0.601 0.26 0.042 *---Signif. codes: 0 ‘***’ 0.001 ‘**’ 0.01 ‘*’ 0.05 ‘.’ 0.1 ‘ ’ 1Permutation: freeNumber of permutations: 999
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 77 / 103
Unconstrained Ordination Environmental Variables
Plotting Environmental VectorsLimit p < 0.1
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http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 78 / 103
Unconstrained Ordination Environmental Variables
Fitting Environmental surfaces
> ef <- envfit(vare.mds ~ Al + Ca, varechem)> plot(vare.mds, display = "sites")> plot(ef)> tmp <- with(varechem, ordisurf(vare.mds, Al, add = TRUE))> tmp <- with(varechem, ordisurf(vare.mds, Ca, add = TRUE, col = "green4"))
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 79 / 103
Unconstrained Ordination Environmental Variables
Plotting Environmental Surfaces
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http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 80 / 103
Unconstrained Ordination Environmental Variables
Factor Fitting I
> dune.ca <- cca(dune)> ef <- envfit(dune.ca ~ A1 + Management, data=dune.env, perm=999)> ef
***VECTORS
CA1 CA2 r2 Pr(>r)A1 0.9980 0.0606 0.31 0.052 .---Signif. codes: 0 ‘***’ 0.001 ‘**’ 0.01 ‘*’ 0.05 ‘.’ 0.1 ‘ ’ 1Permutation: freeNumber of permutations: 999
***FACTORS:
Centroids:CA1 CA2
ManagementBF -0.73 -0.14ManagementHF -0.39 -0.30ManagementNM 0.65 1.44ManagementSF 0.34 -0.68
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 81 / 103
Unconstrained Ordination Environmental Variables
Factor Fitting II
Goodness of fit:r2 Pr(>r)
Management 0.44 0.003 **---Signif. codes: 0 ‘***’ 0.001 ‘**’ 0.01 ‘*’ 0.05 ‘.’ 0.1 ‘ ’ 1Permutation: freeNumber of permutations: 999
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 82 / 103
Unconstrained Ordination Environmental Variables
Plotting Fitted Factors
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http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 83 / 103
Unconstrained Ordination Environmental Variables
Environmental Interpretation
Environmental variables need not be parallel to ordination axes.
Axes cannot be taken as gradients, but gradients are oblique to axes: Youcannot tear off an axis from an ordination.
Never calculate a correlation between an axis and an environmental variable.
Environmental variables need not be linearly correlated with the ordination,but locations in ordination can be exceptional.
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 84 / 103
Unconstrained Ordination Gradient Model and Ordination
Gradient Model and Ordination
Single gradients appear as curves inlinear ordination methods
PCA horseshoe: curve bends inwardand gives wrong ordering of pointson axis 1
CA arch: axis 1 retains the correctordering of sites despite the curve
Environmental interpretation byvector fitting or surface bound to bebiased
Axes cannot be interpreted as“gradients”
Species packing gradient
PCA CA
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 85 / 103
Unconstrained Ordination Gradient Model and Ordination
The birth of the curve
There is a curve in the species space and PCA shows it correctly
CA deals better wit unimodal responses, but the second optimal scaling axisis folded first axis
Gradient space Species space
CA1
CA2
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 86 / 103
Unconstrained Ordination Gradient Model and Ordination
Solutions to the Curvature
Detrended Correspondence Analysis (DCA)
CA axis retains the correct ordering: keep that, but instead of orthogonal axes,use detrended axesProgramme DECORANA additionally rescales axes to sd units approximating tparameter of the Gaussian modelDistorts space, introduces new artefacts and probably should be avoided
Nonmetric Multidimensional Scaling (NMDS) should be able to cope withmoderately long gradients
Constrained ordination may linearize the responses
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 87 / 103
Unconstrained Ordination Gradient Model and Ordination
Running Detrended Correspondence Analysis
> (ord <- decorana(dune))
Call:decorana(veg = dune)
Detrended correspondence analysis with 26 segments.Rescaling of axes with 4 iterations.
DCA1 DCA2 DCA3 DCA4Eigenvalues 0.512 0.304 0.1213 0.1427Decorana values 0.536 0.287 0.0814 0.0481Axis lengths 3.700 3.117 1.3005 1.4789
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 88 / 103
Unconstrained Ordination Gradient Model and Ordination
Default plot
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Eleopalu
Elymrepe
Empenigr
Hyporadi
Juncarti
Juncbufo
Lolipere
Planlanc
PoapratPoatriv
Ranuflam
Rumeacet
Sagiproc
Salirepe
Scorautu
Trifprat
Trifrepe
VicilathBracruta
Callcusp
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 89 / 103
Unconstrained Ordination Gradient Model and Ordination
Community Pattern Simulation
Truth
4 x
2.5t
PCA PCoA
CA DCA NMDS
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 90 / 103
Unconstrained Ordination Gradient Model and Ordination
Short Gradients: Is There a Niche for PCA?
Folklore: PCA with short gradients (≤ 2t).
Not based on research, but simulation finds PCAuniformly worse than CA: With short gradientsabout as good as CA, but usually worse.
There should be no species optimum withingradient: Shortness alone not sufficient.
PCA best used for really linear cases(environment) or for reduction of variables intoprincipal components (but see FA).
Noise dominates over signal in homogeneousdata.
PCA
2 x
1t
CA
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 91 / 103
Unconstrained Ordination Gradient Model and Ordination
Long Gradients: DCA or NMDS
Curvature with long gradients: Need either DCA orNMDS.
NMDS is a test winner: More robust than DCA.
DCA more popular.
DCA may produce new artefacts, since it twists thespace.
CA
DCA
MDS
8x3t
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 92 / 103
Unconstrained Ordination Gradient Model and Ordination
Extended Dissimilarities and Step-across
How different are sites that havenothing in common?
Use step-across points to estimatetheir distance
Flexible shortest path or theirapproximations, extendeddissimilarities
Extended dissimilarity: use onlyone-site steps, do not updatedissimilarities below a threshold
No shared species since rare specieswere not observed: Swantransformation estimates theprobability of finding an unobservedspecies
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2 4 6 8
0.3
0.4
0.5
0.6
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0.8
0.9
1.0
Gradient Distance
Com
mun
ity D
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mila
rity
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2 4 6 8
0.5
1.0
1.5
2.0
2.5
Gradient Distance
Ext
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d D
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mila
rity
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 93 / 103
Unconstrained Ordination Gradient Model and Ordination
Strong and Weak Ties
Maximum dissimilarities (no sharedspecies) are tied
Strong tie treatment tries to keeptied values together and putsmaximum dissimilarites to a circle
Weak tie treatment allows breakingties and straightens the axes: nowthe default in vegan, whereas earlierwas impossible
strong weak
stepacross swan
8 x 1.5 sd8 × 1.5 sd units, Gaussian binary response
http://cc.oulu.fi/ jarioksa/ (Oulu) Multivariate Analysis in Ecology January 2016 94 / 103