Post on 21-Sep-2018
Organismos edáficos y comunidades vegetales: determinantes ambientales
de su distribución ante el cambio climático
Juan J. Jiménez Jaén Investigador ARAID
IPE-CSIC
XX Cursillo sobre Flora y Vegetación en el Pirineo
Jaca, 22 de Julio de 2015
Global diversity
After: Wall, web sites
Plants 18%
Bacteria and virus <1%
Fungi 4%
Soil animals 23% (i.e. ~ 360,000)
Other Animals
55%
Total number of described living species: ~ 1.5x106
Importance of soil fauna for global biodiversity
Insects 80%
Arachnida 12%
Annelida 1%
Vertebrata <1% Microinv
2%
Other Arthropoda
5%
Taxonomic composition of the global soil fauna
Soil organisms are packed extremely dense
• 300,000 mites m -2
• 100,000 collembola m -2
• 2x10 6 nematodes m -2
Soils are commonly utmost biodiverse
• @ 25% of 1.5x10 6 described species
• Five - fold the diversity of forest canopy
Handful of forest soil harbours
• Hundreds/thousands of species of soil meso-fauna and tens of macrofauna (Schaefer & Schauermann 1990)
• 4,000 genotypes bacteria and 2,000 species of fungi in 1 g soil (Torsvik et al. 1994; Hawksworth 2001)
Soils: the third biotic frontier
The enigma of soil biodiversity, or why so many species?
Main taxonomic groups of soil organisms on a body-size basis
R2= 0,41 20
40
60
80
100
1m 100m 10mm 1m
Mean body size
Ta
xo
no
mic
defi
cit
Deficit 90% for organisms
<100 um
1024
m
1 2 4 8 16 32 64 128 256 512 1024 1 2 4 8 16 32 64 128 256 512
mm
Nematoda
Acari
Collembola
Diplura
Symphyla
Enchytraeidae
Hymenoptera (Formicoidea)
Diptera
Isopoda
Myriapoda
Arachnida
Coleoptera
Mollusca
Oligochaeta
Microfauna Mesofauna Macrofauna
100 m 2 mm 20 mm
Body size
Isoptera
(After Swift et al. 1979; Decaëns 2010) Pictures © J.J. Jiménez
Global number of described species and estimated number of existing species of the main taxonomic groups of soil organisms (after Decaëns, Jiménez et al. 2006)
0,01
0,1
1
10
100
1,000
10,000 B
acte
ria
Fun
gi
Nem
ato
da
Pro
tozo
a
Aca
ri
Colle
mb
ola
Dip
lura
Sym
ph
yla
En
ch
ytr
ae
idae
Iso
pte
ra
Form
icoid
ea
Dip
tera
Iso
po
da
Ch
ilop
od
a
Derm
ap
tera
Bla
tto
idea
Dip
lop
od
a
Ara
ch
nid
a
Cole
op
tera
Mo
llusca
Pa
uro
poda
Olig
och
ae
ta
Cae
cili
an
Squ
am
ata
Ma
mm
alia
Sp
ecie
s n
um
ber
(x 1
.000)
NE
NE
NE
NE
NE
NE N
E
NE
NE
NE
NE
Described species
Undescribed species
Body size
Soil organisms (Known and estimated figures)
• How many soil
invertebrate species
exist worldwide?
• Right now there is no
soil where we are
able to identify or
even quantify all the
resident invertebrate
species.
www.prairieecosystems.pbworks.
com/Dennis-NaturalistGuide
90% for organisms <100 µm
(Herbert, 2003)
!!! an unsuspected number of cryptic species not
distinguished on a morphological basis.
Filters on soil fauna biodiversity Species pool hypothesis
(Decaëns, Jiménez et al. 2006)
Spatial scale
REGION
PATCH
LANDSCAPE
ECOSYSTEM
GLOBAL
COMMUNITY
Anthropogenic
changes
Global changes,
N2 deposition
Soil erosion
Land use changes
Agricultural practices
Exotic introduction
(invasive species)
Pictures © Pavel Krasensky; J.J. Jiménez; Tree of life web
Climatic conditions
Landscape structure
Spatial competitive
mechanisms
Soil abiotic factors
Land use practices
Biotic interactions
Modelos e hipótesis que explican la gran diversidad de
organismos del suelo y sus relaciones?
AG Above- and belowground interactions
(after Kardol and Wardle 2010)
BG Level of
org
aniz
ation
Level of organization = criterion level (Allen and Hoekstra 1990)
sp
ecie
s
com
munity
ecosyste
m
Species interactions
Ecosystem function
Community composition
Trophic
Non-trophic
Environmental heterogeneity (constraints)
Spatial and temporal stability
Diversity, assemblages
Carbon storage,
Water infiltration
Patch
Plot
Landscape
Spatial scale
belowground
aboveground
direct pathway to plants indirect pathway to plants
Trophic based below- and aboveground interactions
root feeding fauna
mycorrhiza
pathogens
(Wardle et al. 2004; Osler and Sommerkorn 2007)
food web
- Integration of biogeochemical and soil food web models
Soil ecosystem engineering – Non-trophic based interactions
Ecosystem engineers (sensu Jones et al. 1994)
OM
Aggregates
(Decaëns et al. 2008)
Major drawbacks: Ecosystem engineering is overlooked
Soil habitat transformation
Indirect modification of resources via
production of physical structures
• In soil: earthworms, ants, termites,
and roots
Pheidole sp1
Atta laevigata
Trachymyrmex sp.
Atta cephalotes
Velocitermes sp.
Nasutitermes sp.
Microcerotermes sp.
Termitinae
Ruptitermes sp.
Spinitermes sp.
Globulares
Martiodrilus sp.
Hyperiodrilus africanus
Millsonia anomala
colonización y biomasa radicular
Granulares
Prosellodrilus sp.
Existing Models on SOM Turnover – relevance for Climate Change predictions
PF
C
HS
SOM
Turnover
Molecular structure
Physical
heterogeneity
Humic
substances
Fire-
derived C
Soil depth
Roots
Permafrost
Soil micro-
organisms
Main elements
State-of-the-art (Schmidt et al. 2011)
Soil fauna totally
ignored!
ESSEM COST Action ES1406: “Soil fauna: key to soil organic matter dynamics and modelling (KEYSOM)”
COST is supported by the EU Framework Programme Horizon 2020
Functional diversity of soil
ecological engineers
Visible and Near Infrarred Spectroscopy (NIRS) signatures of biogenic structures
Specific organic fingerprints
in biostructures
(Jiménez et al., sin publicar)
Soil as a mosaic of
functional domains
Axis I (94%)
-4 -3 -2 -1 0 1 2 -0.6
-0.4
-0.2
0
0.2
0.4
0.6
0.8
Earthworm casts
Bulk soil
Axi
s II
(5
%)
Bulk soil Ant 2 BS
Earthworm BS Ant 1 BS
350 2500 0
0.1
0.2
0.3
0.4
0.5
Ant deposits 1
Ant deposits 2
nanometers
Ab
sorb
ance
val
ue
s
A
B
VNIR spectra
Study site
Surroundings of Ordesa and Monte
Perdido National Park, UNESCO Heritage
and LTER network site.
Alpine grasslands grazed by domestic
cattle during summer.
July-August 2014;
Earthworm species:
Aporrectodea rosea Savigny 1826
Lumbricus friendi Cognetti, 1904
Prosellodrilus pyrenaicus (Cognetti, 1904)
Climate alpine, 5 ºC and 1,720 mm
Study site
Smooth grazing gradient
Stocking pressure High Intermediate Low
Nardion strictae Bromion erecti
(Bueno and Jiménez, 2014)
Plant richness Medium High Low
Plant community Rumicion Bromion Nardion
Dominant plant Chenopodium bonus-henricus, Festuca rubra, Nardus stricta
species Trifolium repens, Agrostis capillaris,
Poa supina Trifolium pratense,
Lotus corniculatus
Lab protocol
EW removal 1d
Soil and ew casts
Incubation
1 – 32 (64) days
3 reps /
3 species
casts
retrieval
sieved at <200 µm
NIR spectral
readings
QualitySpec®
spectrophotometer
C and N determinations with dry combustion
method (Variomax CN Analyzer, Germany) Diffuse reflectance
(Stenberg et al. 2010)
<2 mm sieved soil
+
A =
log(1
/R)
(UnscramblerX 10.3, CAMO software)
Data analysis
Reflectance (R) is converted to absorbance (A)
NIR spectra from 1100
to 2300 nm (10 nm
intervals) further
transformed to
Savitzky-Golay 2nd deriv
(noise reduction).
(Savitzky and Golay 1964)
PCAs and Partial Least Square Regression and NIRS library use
Data analysis
Noise reduction
Unscrambler software
d = 5
s1_1
s1_16
s1_2
s1_32
s1_4
s1_8
A. rosea
RMSECV: 52.6%
d = 5
s2_1
s2_16
s2_2
s2_32
s2_4
s2_8
L. friendi
RMSECV: 57.3%
NIR spectra of ageing casts
d = 2
s3_1
s3_16
s3_2
s3_32 s3_4
s3_8
P. pyrenaicus
RMSECV: 49.8%
NIR spectra of ageing casts
d = 5
s1_1
s2_1
s3_1
Not identified:
44.4% (4 casts) !!
Projection of field signals
A. rosea
L. friendi
P. pyrenaicus 11.1% identified (1 cast)
33.3% identified (3 casts)
11.1% identified (1 cast)
Sem
i-v
aria
nce
Lag distance (m)
0 5 10 15 20
0.005
0
0.015
0.010
0.020
0.030
0.025
0.035
335224
251268447161
178
0.0106067 Nug(0) + 0.0220092 Sph(8.11863)1
20
30
40
50
60
70
45
0 4515 30
15
30
C
concentration
(g kg-1)
Spatial distribution of selected soil variables
(Jiménez et al. 2011)
Lag distance (m)
0 5 10 15 20
Sem
i-v
aria
nce
1.46103 Nug(0) + 4.41099 Sph(721.71)
1
178
161447
335
224
251
268
0
1
0.2
0.4
0.6
0.8
1.2
1.4
1.6
1.8
0
20
40
60
80
100
120
140
160
180
0 4515 30
45
15
30
Soil penetration
resistance (kg cm-3)
Fine roots Length (m)
Easting (X)
Nort
hin
g (
Y)
Coarse roots Length (m)
Easting (X)
Nort
hin
g (
Y)
Fine roots Weight (kg)
Easting (X)
Nort
hin
g (
Y)
Coarse roots Weight (kg)
Easting (X)
Nort
hin
g (
Y)
Spatial distribution of resources and organisms
Regular grid of 10x10 points
Soil sampling (Metal cylinders):
1 Size-class aggregates
2 Root length and biomass (fine and
coarse)
3 C, N and P determinations
4 Bulk density, hydraulic conductivity
and soil compaction
Vegetation – soil invertebrate
interactions
(Jiménez et al. 2014)
Spatial distribution of resources (fine and coarse roots)
(m sample-1) (g dry wt sample-1)
Cross-correlogram for roots
and soil nutrient- and
physical-related variables:
FiRL with SOC and P
CoRL with N, SOC and
P
FiRW with SOC and P
and litter
CoRW with SOC, P
FiRL with Aggregates,
Cond and Moisture
CoRL with Aggregates
and Cond
FiRW with Aggregates
and BD
CoRW with Aggregates
Spatial exclusion
Glossodrilus
Aymara Andiodrilus
Genus 1
Andiorrhinus
Ocnerodrilidae Martiodrilus
Litter
N0-5
N5-10
P0-5
P5-10
C0-5
C5-10
C:N 0-5
C:N5-10
FiRL
CoRL
FiRW
CoRW
PR5 PR10 PR20
0.053-
0.125
0.125-0.25 0.25-0.5
0.5-1
1-2
2-5
5-10
>10
BD
Comp
Cond
Hum(%)
-0.43
0.36 -0.36 0.3
F I (64.12%)
F II (1
7.7
4%
)
p<0.0001
Co-Inertia analysis +
Partial Mantel test
0 5 10 15 20 25 30 35 40 45 0
5
10
15
20
25
30
35
40
45
0
5
10
15
20
25
30
35
40
45
Aymara
Ocnerodrilidae
Overlaid contour and classed post maps (surfer) of
SADIE clustering indices for counts of two species
(Jiménez et al. 2011)
Ew effect
Nutrient resources
En
do
gei
cs
ass
emb
lage
Eart
hw
orm
com
mu
nit
y
Intersampling
distance = 10 m
Ep
igeic +
an
ecic
assem
bla
ge
An
dio
dri
lus,
Aym
ara
an
d n
ew
gen
us
1 a
ssem
bla
ge
Martio
drilu
s,
Glo
ssodrilu
sa
nd
new
gen
us 2
assem
bla
ge
(Jiménez et al. 2014)
Variation partitioning in a spatial context
10 20 30 40 50 60
0.0
02
0
.00
6
0.0
10
0
.01
4
Distance (m)
Sem
ivar
ian
ce
PCNM1
PCNM3
PCNM5
PCNM8
PCNM12
PCNM33
PCNM51
Principal Component of Neighbouring Matrices (Dray et al. 2006) and
Variation partitioning (Borcard et al. 1992; Peres-Neto et al. 2006)
Multi-scale spatial relationships
(Jiménez et al. 2014)
Environmental contribution to observed spatial pattern
(>30 m) (10-20 m) (<10 m)
CLIMATE CHANGE – WHAT IS PREDICTED?
Intergovernmental Panel on Climate Change (IPCC 2007) 4th Assessment Report •Elevated atmospheric CO2 (eCO2)
• From 390 ppm in Jan 2011 to over 550 ppm by 2050 • Antarctic ice cores show that current levels are
unprecedented over the past 650 kyr •Increased temperatures, especially over land and at most high northern latitudes (1.8 – 4 ˚C) •The planet has already warmed up by c. 0.75 ˚C in the 20th century
(IPCC 2007)
FUTURE TEMPERATURES : 30 & 100 YEARS
B1 global environmental sustainability
A1B rapid economic growth (balanced emphasis on all energy sources)
A2 regionally oriented economic development (a more divided world)
FUTURE RAINFALL: 2090-99 RELATIVE TO 1980-99
Increased rainfall in high latitudes and east Africa
Decreased rainfall in most subtropical regions
Increased rainfall during rainy seasons in the tropics
Altitudinal gradients of (a) species richness of ants communities in the Smoky Mountains (after Kusnezov, 1957) and French earthworms (after Dahmouche, 2007; Bouché, 1972), and (b) taxonomic richness of several macro-invertebrate groups in Sarawak mountains (after Collins, 1980).
Altitude (m) N
um
be
r o
f fa
mil
ies
0
2
4
6
8
10
12
0 500 1000 1500 2000 2500
Isoptera
Coleoptera
Diptera
b
0
10
20
30
40
50
60
0 500 1000 1500 2000 2500
Sp
ec
ies
ric
hn
es
s
Lumbricidae
Formicoidea
a
0
40
80
120
160
70 60 50 40 30 20 10 0 10 20 30 40
Sp
ec
ies
ric
hn
es
s
Degrees of latitude
50
North South Isoptera Formicoidea
Acari
Sp
ec
ies
ric
hn
es
s
0
100
200
300
400
500
0 10 20 30 40 50 60 70 80
North or south
a
b
Latitudinal gradients of species richness for (a) oribatid mites (after Maraun et al., 2007) and (b) termites (after Lavelle & Spain, 2001) and ants (after Lavelle & Spain, 2001; Kusnezov, 1957).
Abundance gradients General rule: Increased species richness towards the equator for terrestrial ecosystems and increased diversity towards the pole in marine ecosystems
Axis I = 37.8%
Ax
is I
= 1
3.5
%
P<0.001
10N_0
10N_10 10N_30 10N_5
10OTC_0
10OTC_30
N_0
N_10
N_30
N_5
OTC_0
OTC_30
N addition + Temp increase
Ecoplacas BiologTM
???
Combined effects of climate change related factors
on soil organisms (bacteria) – CTP Project
Plateau de Beille (Francia)
Facts on soil organisms in response to climate change
• Important role in C cycling; they control the C sequestration process and influence
greenhouse gas emissions; COST Action 2015-2019.
• Few studies to date so lack of generalizations, but single and combined effects are
expected; responses context dependent.
• Soil living species (relatively constant temperatures) are less sensitive to changes in
T than above-ground species, which live under more fluctuating T regimes
• Changes in geographical distribution of most invertebrates; even to remain in areas
to which they are well adapted.
• Biotic responses include persistence in situ, range shifts to more tolerable climes or,
failing these, extinction;
• Likely increase in the abundance and diversity of invertebrate pests;
Cock, M.J.W., Biesmajer, J.C., Canon, R.J.C., Gerard, P.J., Gillespie, D., Jiménez, J.J., Lavelle, P., Raina, S.K. 2015.
“Invertebrate genetic resources for food and agriculture and climate change”. In: FAO (Ed.) Coping with climate change.
Rome.
Soil organisms will have to adapt, move or extinct