Environmental Drivers of Species Diversity and Turnover

Bort (Robert) Edwards, Postdoctoral Fellow, Smithsonian Institution, Washington DC (Vicki Funk)
 Environmental Drivers of Species Diversity and Turnover

Two fundamental questions:

Where do plants grow? (what is their distribution and diversity?)

Why or how do they grow there? (what is their environment?)

 Environmental Drivers of Species Diversity and Turnover The Compositae

Astereae (~2800sp)
Bahieae(~85)
Cardueae (~2500)
Chaenactideae (<30 sp)
Coreopsideae (~550 sp)
Eupatorieae (~2200 sp)

Helenieae (~120 sp)

Heliantheae (~1500 sp)
Madieae (~200 sp)
Millerieae (~400 sp)
Perityleae(~50sp)

Polymnieae (<10 sp)*

Compositae (Asteraceae) Tageteae (~270 sp)

daisies, sunflowers, thistles, tarweeds etc Vernonieae(~1300 sp)
~1 in 10 flowering plants
Environmental Drivers of Species Diversity and Turnover The Compositae

Mandel et al., unpublished

Compositae (Asteraceae)
13 majority Nth American Tribes
Environmental Drivers of Species Diversity and Turnover: Collections Data

13 tribes

3,116 species

>1,954,000 collection records reduced to ~500,000 after cleaning)

Environmental Drivers of Species Diversity and Turnover Collections Data

100km grid cells

1900 (Astereae) - 152 (Perityleae)

Biodiverse: Laffan et al., 2010

Environmental Drivers of Species Diversity and Turnover Diversity
 Environmental Drivers of Species Diversity and Turnover Diversity

Centers of diversity show considerable similarity across tribes

Correspond to areas of W and SW USA, Central and Northern Mexico

Regions recognized as having diverse and mosaic environments

Extremes in environment

But: do differences between tribes represent idiosyncratic environmental preferences?

- What environmental variables are associated

with diversity and turnover?
Environmental Drivers of Species Diversity and Turnover Environment

187 variables

Climate
AdaptWest 1km averages derived from daily
weather data
(Wang et al., 2016)
Soil
SoilGrids 250m v0.5
(Hengl et al., 2017)

Geochemistry
USGS interpolated layers from 4800 point samples
(Smith et al., 2014)

Topography
Digital Elevation Model - served by SoilGrids
(Hengl et al., 2017)
Environmental Drivers of Diversity: Environmental Data
“Variable reduction is a truly dark art” – Voldemort (probably)
 Environmental Drivers of Diversity: Variable Reduction

n=187
Soil var1
Soil var2
Soil var3
Soil var4
Soil var5
Soil var6
Soil var7
Soil var8
Soil var9
Soil var10
Soil var11
Soil var12

n=50
Soil var13
Soil var14
Soil var15
Soil var16
Soil var17
Soil var18 Soil var1
GEOCHEM

Soil var19
Soil var20
Soil var21
Soil var22
Soil var2
Soil var23
Soil var24
Soil var3
Soil var4
n=7-16
Soil var25
Soil var26
Soil var27
Soil var28
Soil var29
Soil var5
Soil var6
Soil var30
Soil var31
Soil var32
Soil var33 Soil var7 Geochem var1
Soil var34
Soil var35
Soil var36
Soil var37
Soil var8 Geochem var2
Soil var9
Soil var38
Soil var39
Soil var40
Clim var1
Soil var10 Geochem var3
Clim var2
Clim var3
Clim var4
Clim var5
Soil var11 Geochem var4
Clim var1
Clim var6
Clim var7
Clim var2 Geochem var5
GDM
CLIMATE

Clim var8
Clim var9
Clim var10
Clim var11
Clim var3 Geochem var6
Clim var12
Clim var13
Clim var14
Clim var4
Clim var5 Sig. n = 5 -1 0
Remove Elimination of Significant
Clim var15
Clim var16
Clim var17
Clim var18
Clim var6
Drivers of Species Clim var1
Clim var19
Correlated Clim var7 Low GDM
Clim var2 Variables &
Clim var20
Clim var21
Topo var1
Turnover (GDM) Explanatory
Clim var22
var

Variables
Clim var23

Clim var3 Sig.

Clim var24
Topo var2
Final Model
Topo var1
Topo var3 Variables
Topo var2
Topo var3
Topo var4
Topo var5 Topo var4 Clim var4
TOPO

Topo var6
Topo var7
Topo var8 Soil var1
Topo var9
Topo var10
Topo var11 Soil var2 Clim var1 GDM
Soil var3
Sig.
Topo var12
Topo var13
Topo var14
Topo var15
Topo var16
Soil var4 Clim var2
Soil var1 Soil var5 Clim var3
Soil var2
Soil var3 Soil var6
Soil var4
Soil var7
Soil var5
Soil var6
Soil var7 Soil var8 Soil var1 GDM
Soil var8
Soil var9 Soil var2 Sig.
SOILS

Soil var9
Soil var10
Soil var11
Soil var12
Soil var13 Soil var3
Drivers of
Soil var14

Soil var4
Soil var15
Soil var16
Soil var17
Soil var18

Diversity
Soil var19
Soil var20
Soil var21
Soil var22
Soil var23

(PCoA)

Significant
Variables
 Environmental Drivers of Diversity PCoA
Results for Astereae separated by variable class

Comparison between three tribes - A large number of variables have strong

explanatory power

- Idiosyncratic by tribe

- Notable commonalities (11-12 tribes):

Geochem: Quartz

Soil: pH

Topo: Annual Radiation

Climate: Annual Precipitation

 0.5
1.0
As

0
0.25
0.75
te
re
Ba ae
hi

Soil
C Car eae
ha d
en ue
C ac ae

Climate
or tid
eo e
p a

Geochem
Eu sid e
pa ea

Topography
to e
H riea
e e
H le n
e l ie
ia ae
nt
h
M eae
by tribe (%)

ad
M ie a
i ll e e
Pe rie
Top 10 sig variables

rit ae
y
Ta lea
g e
Ve et
rn eae
on
ie
ae
Environmental Drivers of Diversity
PCoA
 0.5
1.0
As

0
0.25
0.75
te
re
Ba ae
hi

Soil
C Car eae
ha d
en ue
C ac ae

Climate
or tid
eo e
p a

Geochem
Eu sid e
pa ea

Topography
to e
H riea
e e
H le n
e l ie
ia ae
nt
h
M eae
by tribe (%)

ad
M ie a
i ll e e
Pe rie
Top 10 sig variables

rit ae
y
Ta lea
g e
Ve et
rn eae
on
ie
ae
As
te
re
Environmental Drivers of Diversity

Ba ae
hi
C Car eae
ha d
en ue
C ac ae
or tid
eo e
p a
Eu sid e
pa ea
to e
H r ie a
el e
H en
el ie
ia ae
nt
h
M eae
ad
M iea
By tribe (%)

il l e e
Pe riea
r it e
y
Ta lea
e
Second 10 sig variables

Ve gete
rn ae
on
ie
ae
PCoA
Environmental Drivers of Diversity gwPCoA
Environmental Drivers of Diversity gwPCoA

Harris et al., 2014

Environmental Drivers of Diversity gwPCoA
Environmental Drivers of Diversity gwPCoA
Environmental Drivers of Diversity gwPCoA

Strongly correlated variables are idiosyncratic

With several exceptions

Classes of variable broadly operate at different scales

Geochemical variables are more highly correlated at more local scales

 Environmental Drivers of Turnover Turnover

n=187
Soil var1
Soil var2
Soil var3
Soil var4
Soil var5
Soil var6
Soil var7
Soil var8
Soil var9
Soil var10
Soil var11
Soil var12

n=50
Soil var13
Soil var14
Soil var15
Soil var16
Soil var17
Soil var18 Soil var1
GEOCHEM

Soil var19
Soil var20
Soil var21
Soil var22
Soil var2
Soil var23
Soil var24
Soil var3
Soil var4
n=7-16
Soil var25
Soil var26
Soil var27
Soil var28
Soil var29
Soil var5
Soil var6
Soil var30
Soil var31
Soil var32
Soil var33 Soil var7 Geochem var1
Soil var34
Soil var35
Soil var36
Soil var37
Soil var8 Geochem var2
Soil var9
Soil var38
Soil var39
Soil var40
Clim var1
Soil var10 Geochem var3
Clim var2
Clim var3
Clim var4
Clim var5
Soil var11 Geochem var4
Clim var1
Clim var6
Clim var7
Clim var2 Geochem var5
GDM
CLIMATE

n = 5 -1 0
Clim var8
Clim var9
Clim var10
Clim var11
Clim var3 Geochem var6
Clim var12
Clim var13
Clim var14
Clim var4
Clim var5 Sig.
Remove Elimination of
Clim var15
Clim var16
Clim var17 Clim var6
Drivers of Species Clim var1
Drivers of Species
Clim var18
Clim var19
Correlated Clim var7 Low Clim var2 GDM Final Model
Clim var20
Clim var21
Topo var1
Turnover (GDM) Explanatory Turnover (GDM)
Clim var22
var

Variables
Clim var23

Clim var3 Sig.

Clim var24
Topo var1 Topo var2
Topo var3 Variables
Topo var2
Topo var3
Topo var4
Topo var5 Topo var4 Clim var4
TOPO

Topo var6
Topo var7
Topo var8 Soil var1
Topo var9
Topo var10
Topo var11 Soil var2 Clim var1 GDM
Soil var3
Sig.
Topo var12
Topo var13
Topo var14
Topo var15
Topo var16
Soil var4 Clim var2
Soil var1 Soil var5 Clim var3
Soil var2
Soil var3 Soil var6
Soil var4
Soil var7
Soil var5
Soil var6
Soil var7 Soil var8 Soil var1 GDM
Soil var8
Soil var9 Soil var2 Sig.
SOILS

Soil var9
Soil var10
Soil var11
Soil var12
Soil var13 Soil var3
Drivers of
Soil var14

Soil var4
Soil var15
Soil var16
Soil var17
Soil var18

Diversity
Soil var19
Soil var20
Soil var21
Soil var22
Soil var23

(PCoA)
 Environmental Drivers of Turnover Turnover

n=187
Soil var1
Soil var2
Soil var3
Soil var4
Soil var5
Soil var6
Soil var7
Soil var8
Soil var9
Soil var10
Soil var11
Soil var12

n=50
Soil var13
Soil var14
Soil var15
Soil var16
Soil var17
Soil var18 Soil var1
GEOCHEM

Soil var19
Soil var20
Soil var21
Soil var22
Soil var2
Soil var23
Soil var24
Soil var3
Soil var4
n=7-16
Soil var25
Soil var26
Soil var27
Soil var28
Soil var29
Soil var5
Soil var6
Soil var30
Soil var31
Soil var32
Soil var33 Soil var7 Geochem var1
Soil var34
Soil var35
Soil var36
Soil var37
Soil var8 Geochem var2
Soil var9
Soil var38
Soil var39
Soil var40
Clim var1
Soil var10 Geochem var3
Clim var2
Clim var3
Clim var4
Clim var5
Soil var11 Geochem var4
Clim var1
Clim var6
Clim var7
Clim var2 Geochem var5
GDM
CLIMATE

n = 5 -1 0
Clim var8
Clim var9
Clim var10
Clim var11
Clim var3 Geochem var6
Clim var12
Clim var13
Clim var14
Clim var4
Clim var5 Sig.
Remove Elimination of
Clim var15
Clim var16
Clim var17 Clim var6
Drivers of Species Clim var1
Drivers of Species
Clim var18
Clim var19
Correlated Clim var7 Low Clim var2 GDM Final Model
Clim var20
Clim var21
Topo var1
Turnover (GDM) Explanatory Turnover (GDM)
Clim var22
var

Variables
Clim var23

Clim var3 Sig.

Clim var24
Topo var1 Topo var2
Topo var3 Variables
Topo var2
Topo var3
Topo var4
Topo var5 Topo var4 Clim var4
TOPO

Topo var6
Topo var7
Topo var8 Soil var1
Topo var9
Topo var10
Topo var11 Soil var2 Clim var1 GDM
Soil var3
Sig.
Topo var12
Topo var13
Topo var14
Topo var15
Topo var16
Soil var4 Clim var2
Soil var1 Soil var5 Clim var3
Soil var2
Soil var3 Soil var6
Soil var4
Soil var7
Soil var5
Soil var6
Soil var7 Soil var8 Soil var1 GDM
Soil var8
Soil var9 Soil var2 Sig.
SOILS

Soil var9
Soil var10
Soil var11
Soil var12
Soil var13 Soil var3
Drivers of
Soil var14

Soil var4
Soil var15
Soil var16
Soil var17
Soil var18

Diversity
Soil var19
Soil var20
Soil var21
Soil var22
Soil var23

(PCoA)
 Environmental Drivers of Turnover GDM

Generalised Dissimilarity Model

- A non-linear model that fits dissimilarity (in diversity) between sites as a function
of the environmental variable values at those sites.

- Contribution of each variable to the model

- Significance of variables through permutation

- Response of turnover to environmental gradients

- Extrapolate species turnover to unsampled sites

(where environment is known)

GDM: Farrier et al., 2007

 Environmental Drivers of Diversity Turnover

Astereae Bahieae Cardueae Chaenactideae Coreopsideae

Helenieae

Eupatorieae

Heliantheae Madieae Millerieae

Tageteae
Perityleae

With geography and collections per site included as model terms

 Environmental Drivers of Diversity Turnover

Significant Turnover Variables

Proportion of Significant GDM Variables
1.00

0.75

Topography
0.50 Soil
Geochem
Climate

0.25

0.00
ae ae ae ae ae ae ae ae ae ae ae ae ae
e re hie ue tide ide rie nie the die rie tyle ete nie
t a rd c s t o le n a lle ri g o
As B Ca na epo pa He elia M Mi Pe Ta ern
e r E u H V
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 Environmental Drivers of Diversity GDM

Significant Turnover Variables Significant Diversity Variables

Proportion of Significant GDM Variables
1.00 1.00

0.75 0.75

Topography
0.50 Soil 0.50
Geochem
Climate

0.25 0.25

0.00 0.00
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 Environmental Drivers of Diversity Summary

SUMMARY:

Diversity
- Some common putative drivers of diversity across Asteraceae tribes:
quartz, pH, precipitation, elevation

- Many idiosyncratic suggesting tribe-specific adaptations

- Classes of variable are associated with diversity at different scales:

- Climate = broad
- Geochem = narrow
Turnover
- Putative drivers of turnover across tribes are even more idiosyncratic

- Few common factors

-Variables strongly associated with turnover are not necessarily those

that are associated with diversity
 https://powellcenter.usgs.gov/

Vicki Funk Elisabeth Bui Chase Mason

Marty Goldhaber Ian Pearse Travis Nauman Jen Cartwright

Pamela Soltis Brian Anacker Joe Miller Jim Thompson

@BortEdwards www.bortedwards.weebley.com
Geo-Eco-Evo Community Clusters

Species diversity and distributions vary across the landscape in structured ways
Diversification Rate The Compositae (again)

Mandel et al., unpublished

Compositae (Asteraceae)
14 majority Nth American Tribes
Evolution of environmental tolerances in a phylogenetic framework
Methods – Diversification Analysis

- Tip-rate correlation tests using the

Diversification Rate metric (Jetz et al., 2012)

- First regressed diversification rate

against single environmental
variables

- To control for false positives we generated

1000 random permutations of the data and
repeated the analysis process for each
simulated dataset.
 Coreopsideae
7 best models (R2 = 0.36 - 0.37)

Included variables:
Total 14 Å Clays [C horizon]
Zinc [C horizon]
Total Plagioclase [C horizon]
Total Plagioclase [A horizon]
Sodium [A horizon]
Sodium [C horizon]
Sodium [Top 5 cm]
Total Feldspar [C horizon]
Aluminum [A horizon]
Threshold R2 = 0.09
 Eupatorieae
4 best models (R2 = 0.12 - 0.13)

Included variables:
Phosphorus [A horizon]
Mean Annual Temp.
Mean Winter Temp. (Dec-Feb)
Mean Temp. Coldest Month
Extreme Min. Temp. over 30 yrs

Threshold R2 = 0.03
 Helenieae
4 best models (R2 = 0.13 - 0.15)

Included variables:
Calcite [A horizon]
Total Plagioclase [C horizon]
Total Plagioclase [A horizon]
Total Carbonates [A horizon]
Total Feldspar [C horizon]

Threshold R2 = 0.10
 Heliantheae
1 best model (R2 = 0.18)

Included variables:
Total Zeolites [C horizon]
Growing Degree Days >18°C

Threshold R2 = 0.04
 Madieae
8 best models (R2 = 0.06 - 0.08)

Included variables:
Cation Exchange Capacity [0cm]
Cation Exchange Capacity [5cm]
Silt Content [0cm]
Phosphorus [A horizon]
Phosphorus [C horizon]
Phosphorus [Top 5cm]
Total 10 Å Clays [A horizon]

Threshold R2 = 0.05
 Millerieae
3 best models (R2 = 0.56 - 0.57)

Included variables:
Potassium [C horizon]
Total K Feldspar [A horizon]
Lithium [C horizon]
Total Clays [A horizon]
Total Plagioclase [A horizon]
Total Plagioclase [C horizon]

Threshold R2 = 0.24
 Tageteae
1 best model (R2 = 0.41)

Included variables:
Total K Feldspar [C horizon]
Aluminum [C horizon]

Threshold R2 = 0.11
 What factors drive diversification rates
across North American Asteraceae?
• Abiotic environmental factors can explain >50% of variation in
diversification rate, and up to 30% above randomly simulated
data.

• This varies substantially among tribes.

• Different subset of variables that correlates with total diversity

<5% 15% >30%

Madieae Helenieae Eupatorieae Heliantheae Coreopsideae Tageteae Millerieae

 What factors drive diversification across
North American Asteraceae?

• As a variable class, geochemical and mineralogical

factors are consistently more correlated with
diversification rate than climate, topography, or
soil profile/texture.

Phosphorus, Potassium, Aluminum

Plagioclases, Feldspars, Clays

Al Clays
Geo-Eco-Evo Extreme Environments
Geo-Eco-Evo Extreme Environments