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