beijing sept 2011 final - cern holzapfel beijing sept__ 2011.pdf15 an v e a n overlap (% of tran 12...

9/16/2011

1

Nature, …… red in tooth and claw

oror… green in leaf and flower

J. Gurche, 1991

Communities: do they exist?

Superorganism or coincidence?

Frederick E. Clements (1874-1945) Henry A. Gleason (1882-1975)

Closed community Open communityvs.

9/16/2011

2

Community Views

Clements:• community as superorganism -

functions of various species are connected like the parts of the body

• Gleason: chance associations of species whose adaptations and requirements enable them to live togetherbody.

- component species had coevolved interdependent functioning

- communities are discrete entities with recognizable boundaries

– component species occurred together largely by coincidence

– no distinct boundary where one community meets another

What are communities?

All the populations of different species co-inhabiting a given area.

What delineates an area?

?Are communities just random assemblies?

Regularities can be striking!

Succession after disturbance

from: PhysicalGeography.net

9/16/2011

3

Assembly RulesThe development of “climax” communities (to some extent) suggest that communities assemble by some rules:

The three filters:

Who gets there? DispersalWho gets there? ……….. DispersalWho can live there? …… Adaptation to abiotic conditionsWho can dominate over others? Adaptation to biotic conditions

Facilitation

Community evolution

Dispersal filter Abiotic filter Biotic filter

(Whittaker & Woodwell 1971)

Human transport

Community: an ecological Bigfoot?

How can evolution shape communities (and diversity)?

Answer still open – evolution might be very local and below the species level (metapopulation level)the species level (metapopulation level)

Community evolution: evolutionary adjustments of individuals and populations in regard to each other:

possible but difficult to prove

“Changes in genetic constitution having important consequences at the community level irrespective of taxon”

(Whittaker and Woodwell 1971)

9/16/2011

4

Fusion Ecology: plant communities and global change

FusionEcologyLab

Claus Holzapfel and Lab, Newark

What is “Fusion Ecology”?

Formation of “novel communities” fused by coexistence of non-native and native species

Search for evolution or sorting among new neighbors resulting in easing of conflict

Prediction of future communities and their ecosystem effects

Part of “Novel Ecology”Gradient:

Pristine communitiesNovel communitiesConstructed communities

FusionEcologyLab

9/16/2011

5

What is“Fusion Ecology”?

Urban “Wildlands” Natural Wildlands, e.g. deserts and alpine-tundra

Global Change: Interactions of old and new community members

Natural Wildlands, e.g. deserts and alpine tundra

FusionEcologyLab

Fusion EcologyOlder Evidence

Roy Turkington,UBC, Vancouver

9/16/2011

6

Communities: evidence for community evolution

Older communities show less strong negative interactions

Relative yield of a grass in two species mixtures

from Turkington & Mehrhoff 1990

Lolium

Community stability in dependence of community age: pastures in British Columbia.

7 years

26 years

Axi

s 3

(12.

9% v

aria

tion)

Ordination trajectories of cover values

from: Aarssen & Turkington (1985)

45 years

A

Axis 1 (41.7% variation)

9/16/2011

7

Allopatric site:Lolium absent

1 0

1.5

2.0

2.5

3.0

y w

eigh

t (g)

T L-T-T-L

Design after Connell (1980) 0.0

0.5

1.0

-T-L -T-L -T

Tota

l dry

Sympatric plants are not released from competition as much as the allopatric ones

Sympatric site:Lolium dominant

-T-L T L

from: Turkington & Mehrhoff (1990)

Removal of grass does not improve growth since L and T separated niches

Fusion Ecology

New EvidencePlant communities and global change: invasion and climate change

1. Borders between new and old neighbors

2. Root interactions among new and old neighbors

FusionEcologyLab

3. Desert shrubs and plant invasion

4. Middle Eastern plants and climate change

5. Tibetan plant communities and climate change

9/16/2011

8

Fusion EcologyNew Evidence

1. Borders between new and old neighbors2. Root interactions among new and old neighbors

FusionEcologyLab




Species 1 Species 2

Border Description: the transect technique

FusionEcologyLab

FusionEcologyLab

Melanie Kaeser, Jack Chapman, Ting-Min Wu, Jessica Schnell, Greg Burdulis, Mina Andrews, Jose Moreno, Dennis Solis, Mohammad Misbah, Hadas Parag, Jonathan Lansey, Kunj Patel, Christi Cincotta, Mark June-Wells

9/16/2011

9

Sharp borders between sympatric plants

Fallopia Microstegium

80100

aled

%)

Sympatric Border

Fallopia Microstegium

80100

aled

%)

Sympatric Border

0204060

1 2 3 4 5 6 7

Transect

Cov

er (s

ca0

204060

1 2 3 4 5 6 7

Transect

Cov

er (s

ca

Fallopia Artemisia

100d %

)

Allopatric Border

Fallopia Artemisia

100d %

)

Allopatric Border

from Northrup et al. 2005

0

50

1 2 3 4 5 6 7

Transect

Cov

er (s

cale

d

0

50

1 2 3 4 5 6 7

Transect

Cov

er (s

cale

d

FusionEcologyLab

Species pairs with different origins (allopatric) have significantly larger overlap compared to sympatric pairs

20

25

sect

)

0

5

10

15

an ve ve an an ve

Ove

rlap

(% o

f tra

ns

12 10 9 10 1111

FusionEcologyLab

Asian

-Euro

pean

Asian

-Nati

ve

Europ

ean-N

ative

Asian

Europ

ean

Nativ

e

Type of interaction

9/16/2011

10

Species pairs with narrow overlap (sympatric) Species pairs with wide overlap (allopatric)

50

60

soil)

60

70

soil)

Root densities along border gradients

Sympatric Pairs Allopatric PairsRoot density and Border Overlap

0

10

20

30

40

50

Microst. Border Fallopia

Roo

t den

sity

(cm

/cm

3

50

60

m3 s

oil)

60

70m

3 soi

l)

0

10

20

30

40

50

60

Centaurea Border Fallopia

Roo

t den

sity

(cm

/cm

3 s0

10

20

30

40

50

60

70

0 10 20 30

Width of overlap (% of gradient)

Roo

t den

sity

(cm

/cm

3 soi

l)

FusionEcologyLab

0

10

20

30

40

Artemisia Border Centaurea

Roo

t den

sity

(cm

/cm

0

10

20

30

40

50

Artemisia Border Fallopia

Roo

t den

sity

(cm

/cm Width of overlap (% of gradient)

What plant community is this?

9/16/2011

11

A field guide to novel and not so novel plant communities:

the role of introduction of non-native organisms

Traditional assemblies Novel assembliesTraditional assemblies Novel assemblies

Expat Assemblies

Old native communities

New mixedassemblies

New native Assemblies New mixed

bliNew mixed

Simplified after Prasse & Holzapfel (in prep.)

assembliesAssemblies assembliesassemblies

0.9

1

atio

n

Spontaneous Vegetation: Old to Novel gradient

0.1

0.2

0.3

0.4

0.5

0.6

0.7

0.8

opor

tion

of s

pont

anou

s ve

get

New mixedNew NativeExpatOld Native

?

0

Pristine Rural Suburban Urban

Human impact gradient

Pro Old Native

9/16/2011

12



FusionEcologyLab

g





Directionality of root growthroot box experiments

FusionEcologyLab

William Scharf

9/16/2011

13

Directionality of root growthroot box experiments

Measure of directionality

FusionEcologyLab

away towardsother plant

away/(away+towards)

Directionality of root growth: root box experiments

1.0

0 2

0.3

0.4

0.5

0.6

0.7

0.8

0.9

0

rtio

n of

root

s to

war

ds o

ther

FusionEcologyLab

0.0

0.1

0.2

Artemisia Aster Calamagrostis Solidago

Interacting species

Prop

or

Artemisia – Solidago reduce root growth towards each other

9/16/2011

14

Monarda fistolosanative

Solidago canadensis native

Artemisia vulgarisexotic

Mark June-Wells

Root cores

Plant

FusionEcologyLab

9/16/2011

15

Root spacing between Pairs

200

300

400

500

600

ngth

bet

wee

n pl

ants

(cm

)

Border100>1000

p=0.023

p=0.453

p g

0

100

Sympatric AllopatricSpecies Pairs

Roo

t len

FusionEcologyLab

Community affinity between sympatric species pair breaks down at distances >100 m.

?alien/allopatric

Testing for Neighborhood Effects

Stre

ngth

of C

ompe

titio

n

Stre

ngth

of C

ompe

titio

n ?

co-evolved/sympatric

Neighborhood EffectLocal Far

S

Time

S

June-Wells & Holzapfel (in prep.)

9/16/2011

16


1. Borders between new and old neighbors2. Root interactions among new and old neighbors

FusionEcologyLab

3. Desert shrubs and plant invasion4. Middle Eastern plants and climate change


Mojave, CA: exotic grasses dominate the annual communities around shrubs

9/16/2011

17

Ecological problems with non-native grasses in the arid and semi-arid West

Bromus rubensin the Mojave Desert

FusionEcologyLab

Bromus tectorumin the Great Basin Desert

Do aliens have stronger negative effects on shrubs than natives?

-0.5

0.0

0.5

on s

hrub

pro

duct

ion

-1.5

-1.0

40 60 80 100

Proportion of aliens (% of density)

r= 0.143< 0.001

2

P

Neg

ativ

e ef

fect

FusioEcoloLab

9/16/2011

18

pastPrior

invasion??

presentInvadedsystem

Aliensystem

?

future

Shrub-annual interaction in California and Israel

Ambrosia dumosa

R2 = 0.2987

40

60

80

100

ality

(% c

over

)

60

80

100

ity (%

cov

er)

Sarcopoterium spinosum

0

20

40

0 50 100 150 200 250

Biomass of annual community (g/m2)

Shr

ub v

ita

R2 = 0.0152

0

20

40

0 50 100 150 200 250

Biomass of annual community (g/m2)

Shr

ub v

itali

FusionEcologyLab

9/16/2011

19


1. Borders between new and old neighbors2 Root interactions among new and old neighbors

FusionEcologyLab





Community effects of Climate Change: a German-Israeli project

Climate versus biotic control of plant coexistence

What determines species distribution? What will happen if species move without

Claus Holzapfel Marcelo Sternberg Dan MalkinsonRutgers University Newark Tel Aviv University, Tel Aviv Haifa University, Haifa

New Jersey Israel Israel

What will happen if species move without their communities?

9/16/2011

20

Prediction of biotic response to climate change

What determines species distribution?

• ‘Climate-envelope’ ApproachSpecies are adapted to a particular climate (or environment)Climate change will shift species distribution accordingly

• Realized vs. Potential NicheSpecies distribution and ecological behavior depends on interaction with other species

‘Climate-Envelope’ Approach…correct on a larger scale

9/16/2011

21

Empirical studies show that in many cases neighbor effects are important:

a single organism vs. an organism in its community context

So, what’s more important:abiotic or biotic factors?

its community context

What determines species performance:Environment (Climate) or Community?

TopographySouth-facing slopes with stony and shallow soil

Mesic Mediterranean - 780 mm

The Climatic Gradient

(Terra rossa to desert lithosol on hard limestone and chalk)

TemperatureMean annual temperature 140C-230C

RainfallMainly winter 5 summer

~ 48

0 kmMediterranean - 540 mm

Mainly winter - 5 summer months with no rainfallRange North-South: 780 to 90 mm

Semiarid – 300 mm

Arid – 90 mm

9/16/2011

22

Mesic Mediterranean

Semi-arid Mediterranean

Avena sterilisFusionEcologyLab

Arid Semi-arid Mediterranean

Crithopsis delileanaFusionEcologyLab

9/16/2011

23

Soil Transplants (Moving the whole Neighborhood)

To make plants move is easy

Avena sterilis : Environment vs community

Environment vs. Community Experiment

AEnvironment vs. community

0.4

0.6

0.8

1

1.2

e-gr

ound

mas

s (g

)

drierhomewetter

Avena

Environment nsCommunity ***E x C nsNeigh mass ns

bb

ab

a

b

a

0

0.2

Semi-arid Medit. Mes.Med.

Sites

Abo

ve Neigh.mass ns

FusionEcologyLab

9/16/2011

24

Crithopsis delileana : Environment vs. community

0 2

Environment vs. Community Experiment

0.05

0.1

0.15

0.2

ove-

grou

nd m

ass

(g)

drierhomewetter

Environment ***

Crithopis

b

a

a b

aba

a

0Arid Semi-arid Medit.

Sites

Abo Community ns

E x C nsNeigh.mass *

FusionEcologyLab

)1(*)))(1(4( 2 KNee NffccrdtdN

Dan’s Model

dt

Logistic growth curve

plusc = effects of affinity to community

ƒe = adaptation to given environment

FusionEcologyLab

9/16/2011

25

1high


Dan’s ModelLogistic growth curve

0.4

0.6

0.8

Gro

wth

rat

e (r

)Afinity to

communitylow

medium

(c)

0

0.2

0 0.1 0.2 0.3 0.4 0.5

Degree of adaptation to environment (fe)(ƒe)


Dan’s Model

)1(*)))(1(4)1(( 21 KNeeg NffccrdtdN

Logistic growth curve (plus additions)

c = effects of affinity to community

ƒe = adaptation to given environment

g = de facto community

9/16/2011

26

g - the ‘de facto community’ (=degree of similarity to home community)

0.81

0.5

)1(*)))(1(4)1(( 21 KNeeg NffccrdtdN

0

0.2

0.4

0.6

0.8

1S

1 S3 S

5 S7 S

9-1-0.8-0.6-0.4-0.200.20.40.6

00.

20.

4

0.6

0.8

1S

1 S3 S

5 S7 S

9

-4.5-4-3.5-3-2.5-2-1.5-1-0.50

0.1

0.5

0.9

S1 S

3 S5 S

7 S9

-10

-8

-6

-4

-2

0

2

= affinity to environment

g

r

(ƒe)

environment

Affinity to community c

low medium high

Fugitive Local hero

Discussion

“Local Hero” “The Fugitive”

Avena Crithopis

High community affinityHigh levels of co adaptation

Low community affinity L ki f d t tiHigh levels of co-adaptation

with native neighbors?

Mechanisms?

Root interaction, mycorrhizal interactions, …

Lacking of co-adaptation with native neighbors?

Species of temporal variable environments (disturbance, climate fluctuations)

9/16/2011

27

“We have reasons tobelieve that species inthe state of nature arelimited in their rangesby the competition ofother organic beingsother organic beingsquite as much, or morethan, by adaptations toparticular climates"

Charles Darwin, 1859, Origin: Chapter V, Laws of Variation: Acclimatization.



3 Desert shrubs and plant invasion3. Desert shrubs and plant invasion



9/16/2011

28

Effects of climate change on high altitude vegetation:

the role of abiotic and biotic control

With

Yangjian Zhang, Xiao Ming, Hadas Parag

9/16/2011

29

Naqu31o30 N4500m

Damxung30o29 N4300m

CommunityLhasa29o40 N3700m

Community experiment2011: Latitudinal gradient

4800m

Community experiment2011: Altitudinal gradient

4800m

4550mDamxung

4300mNaqu

Lhasa

9/16/2011

30

Tibet population experiment - 2012•Move soil and plants down slope : proxi for warming

Naqu31o 30

Damxung 30o29

N

(b)

•Test target plant performance at home now (control) vs. warmer with old & new community

•Community without and without target

•Microbial community with Biolog plates (samples to be send to NJ)

•All plots will be disturbed and one local

Lhasa29o40

s

pwill be kept intact

•(a) Elevation gradient in Damxung (as in community experiment) and (b) latitudinal gradient (Naqu to Lhasa)

(a)

9/16/2011

31

Fusion EcologyMakes any Sense?

Do these pattern suggest evolution within communities?

Are there alternative explanations?

e.g. niche differentiation prior to contact (species sorting)

FusionEcologyLab

If evolution took place in situ:

How important are such processes?

What mechanisms are responsible?

Community assembly is challenged by rapid global change:

invasion and climate change

Facilitation

Community evolution

FusionEcologyLab

Dispersal filter Abiotic filter Biotic filter

9/16/2011

32

Community Evolution: an ecological Bigfoot?

Evolutionary adjustments of individuals and populations in regard to each other are possible but difficult to prove.

However, communities seem to be more than Random assembliesHowever, communities seem to be more than Random assembliesThere are no Superorganisms either.

FundingNJ Department of Environmental Protection

Meadowlands Commission

- Deutsche Forschungs-gemeinschaft- Andrew Melon Foundation- US Department of Defense

FusionEcologyLab

German BMFT

Chinese Academy of Sciences

9/16/2011

33

Fusion EcologyThanks

FusionEcologyLab

beijing sept 2011 final - cern holzapfel beijing sept__ 2011.pdf15 an v e a n overlap (% of tran 12...

Documents