Table S1 Review of previous studies that compared phylogenetic (and functional) distinctiveness between introduced and native species to
test Darwin’s Naturalization Conundrum in natural plant communities (see Thuiller et al., 2010 for a summary of all studies investigating
phylogenetic patterns of invasions). Results find phylogenetic and/or functional similarity (i.e. clustering, “-“), dissimilarity (i.e.
overdispersion, “+”) of invasive species nested in or compared to the native community, or no significant pattern (“ns”). Statistical models are
coded as follows: response variable ~ explanatory variable1 + explanatory variable2. Test is indicated in parentheses, GLM = generalized
linear model.
Reference
Taxonomic
Scale
Spatial Scale
(grain)
Phylogeny
type
(method)
Supertree
Phylogenetic
distance
metrics
MNNPD
MPD
Traits
Strauss et
al., 2006
Poaceae
(genuslevel)
California
(state and five
ecologically
diverse
reserves)
Cadotte et
al., 2009
Angiosperms
Cadotte et
al., 2010
Davies et
al., 2010
Trait
difference
metrics
χ2
Statistical
model (test)
Null
Hypothesis
Results:
Traits
Phylogenetic
distance
metrics ~
pest vs. nonpest (t-test)
None
ns
Australia,
small scale
(Royal
National Park)
and large scale
(whole
continent)
Supertree
&
Maximum
likelihood
(matK,
rbcL, &
ITS),
missing
species
added as
polytomies
Phylogenetic
signal of
occupancy
(Blomberg’s
K)
None
NA
Compared to
95% CI of the
expected
distribution
under the
null model
Randomize
species
occurrence
matrix
NA
- (large
scale), ns
(small scale)
Angiosperms
Central and
Northern
California, 4
sites, 30-50
replicate plots
(1m2)
Megaphylogeny
(matK,
rbcL, &
ITS)
PD
MNND*
MPD*
None
NA
Compared to
95% CI of the
expected
distribution
under the
null model
Randomize
species
occurrence
matrix (swap
names)
NA
- (within
pots), low
phylogenetic
turnover
Angiosperms
California,
serpentine
ecosystem,
small (16m2)
and large
(10,816m2)
spatial scales
Supertree
MNNPD
None
NA
Compared to
95% CI of the
expected
distribution
under the
null model
Randomize
species
occurrence
matrix
NA
+ (stronger at
small scales)
Area of
origin
Results:
Phylogenetic
(scales)
+
Reference
Taxonomic
Scale
Spatial Scale
(grain)
Schaefer et
al., 2011
Angiosperms
(specieslevel)
Azores
(plots 100m2 1km2, Sao
Miguel
Islands,
Azores
Archipelago)
Carboni et
al., 2012
Angiosperms
(specieslevel)
Mediterranean
coastal dunes
(plots 4m2,
64m2, 35km2 )
Phylogeny
type
(method)
Megaphylogeny
(matK,
rbcL)
Phylogenetic
distance
metrics
PNND
MPD **
Traits
Supertree
DNNS †
MDNS †
Leaf
thickness,
SLA, height,
seed mass,
seed shape
Life form,
seed weight,
plant height,
seed number,
dispersal
type, mode of
introduction,
pollination
system
Trait
difference
metrics
AET
DNNTV
Statistical
model (test)
Null
Hypothesis
Results:
Traits
Phylogenetic
distance
metrics ~
Invasive vs.
non-invasive
introduced
(t-test)
3 models:
Probability of
invasion
success ~
phylogenetic
distance
metrics (+)
AET (+)
DNNTV
(GLM)
none
AET: life
form and
seed weight
explain
more
variation
than PNND
DNNTV:
PNND &
dissimilarity
in life form
Abouheif’s
test
Probability of
community
invasion ~
phylogenetic
distance
metrics + plot
identity
(random
effect) +
invader
identity
(random
effect)
(MCMCglmm)
Randomize
presence
/absence of
invasives
among sites,
2-tailed pvalue
explains
pattern for
each invader
Phylogenetic
signal for
leaf
thickness,
seed mass &
seed shape
Results:
Phylogenetic
(scales)
+ (>1km2))
ns (100m2 )
+ (4m2)
- (64m2,
35km2 )
Reference
Taxonomic
Scale
Spatial Scale
(grain)
Phylogeny
type
(method)
Bayesian
from
barcode
regions
(rbcL &
matK)
Phylogenetic
distance
metrics
PNND
MPD
Traits
Bezeng et
al., 2013
Angiosperms
Robben Island,
South Africa,
50m2 plots
Park &
Potter,
2013
Cardueae,
(specieslevel)
Ordonez,
2014
Vascular
Plants
(genuslevel)
Statistical
model (test)
Null
Hypothesis
Results:
Traits
None
Trait
difference
metrics
NA
Phylogenetic
distance
metrics ~
invasive vs.
native
(MannWhitney Utest); PNND
~ scale
(island vs.
plot) + status
category
(invasivenative,
invasiveinvasive,
nativenative)
Randomize
species
occurrence
matrix
NA
Results:
Phylogenetic
(scales)
- (both
observed and
random
communities)
California
(state and
floristic
provinces)
Megaphylogeny
(matK,
trnL-F &
ITS)
MNND
MPD
None
NA
Phylogenetic
distance
metrics ~
invasive vs.
non-invasive
introduced
(t-test)
NRI/NTI
(random
assemblages
of non-native
taxa)
NA
-
Global (83
communities
worldwide;
global,
continental,
habitat types)
Supertree
MNNPD ‡
MPD ‡
SLA,
maximum
height, seed
mass
MFD ‡
MNNFD ‡
Phylogenetic
distance
metrics ~
alien vs.
native
(Bonferronicorrected
unequal
sample sizes
t-test)
NRI/NTI
(random
assemblages)
+
-
Reference
Taxonomic
Scale
Spatial Scale
(grain)
Phylogenetic
distance
metrics
PNND
MPD
D (random
vs. Brownian)
Traits
United
Kingdom
(country and
countryside
survey plots)
Phylogeny
type
(method)
Megaphylogeny
(matK,
rbcL; ultrametric)
Lim et al.,
2014
Vascular
Plants
(specieslevel)
Li et al.,
2015
Lososová
et al., 2015
Trait
difference
metrics
Invasive
vs. noninvasive
introduced
(binary
trait) ~
each trait
(GLM)
Statistical
model (test)
Null
Hypothesis
Results:
Traits
Phylogenetic
distance
metrics ~
Invasive vs.
non-invasive
introduced
(GLM country
scale, linear
mixed effect
model plots))
Randomize
all tips (for
D)
Invasives
have
different
abiotic
preferences
(nitrogen
and
moisture)
Seed Plants
Permanentplot study,
New Jersey,
USA (40 year
survey)
Megaphylogeny
(matK,
rbcL &
ITS)
MNNPD
MPD
MPDab
NA
NA
Standardized
effect size
(SES)
Randomize
exotic tips
NA
-
Vascular
Plants
(specieslevel)
Czech
Republic
Supertree
(ultrametric)
MNTD
MPD
None
NA
Standardized
effect size
(SES)
Random
alien
assemblages
NA
-
Life form,
height,
clonality, and
Ellenberg
indicator
values for
light, soil
fertility, soil
pH, salt
tolerance
Results:
Phylogenetic
(scales)
ns (both
scales)
Phylogenetic Distinctiveness Acronyms
+ : over-dispersion (DNH)
- : clustering
ns : not significant
D: phylogenetic signal of invasiveness (Fritz & Purvis, 2010)
DNNS: distance of the invader to its nearest native species
MDNS: mean distance of the invader relative to native species
MNND: distance from each nonnative taxon to its nearest native relative
MNND*: mean nearest neighbor distance (Webb et al., 2002)
MNNPD: mean phylogenetic distance for each taxa to nearest native
MNTD: mean nearest taxon distance
MPD: mean phylogenetic distance of each taxa to the native community
MPDab: abundance-weighted MPD
MPD*: mean pairwise distance (Webb et al., 2002)
NRI: net relatedness index (difference between the observed and expected
MPD)
NTI: nearest taxon index (difference between the observed and expected
MNNPD)
PD: phylogenetic diversity (Faith, 1992)
PNND: phylogenetic nearest neighbor distance
* Calculated for status groups separately
** At different divergence time intervals (SPACoDi)
† Residuals regressed against total plot richness
‡ Standardized by the maximum distance for all compared taxa
Functional Distinctiveness Acronyms
+ : over-dispersion (DNH)
- : clustering
ns : not significant
AET: absolute ecological trait values
DNNTV: difference in trait values between an
introduced and its nearest native relative
MNNFD: mean functional distance to the nearest native
relative for each taxa
MFD: mean functional distance between each taxa and
all native species it co-occurs with
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