community ecology i. introduction ii. multispecies interactions with a trophic level iii....

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Community Ecology I. Introduction II. Multispecies Interactions with a Trophic Level III. Multispecies Interactions across Trophic Levels IV. Succession V. Biodiversity: Patterns and Processes A.The Species-Area Relationship 1. The pattern 2. The Theory of Island Biogeography 3. Why is this important? Fragmentation 4. The SLOSS debate

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Community Ecology

I. Introduction

II. Multispecies Interactions with a Trophic Level

III. Multispecies Interactions across Trophic Levels

IV. Succession

V. Biodiversity: Patterns and Processes

A.The Species-Area Relationship

1. The pattern

2. The Theory of Island Biogeography

3. Why is this important? Fragmentation

4. The SLOSS debate

- The SLOSS Debate

- So, to preserve biodiversity (and the ecosystem services it provides to humanity), conservationists began to consider the best strategy for maximizing the preservation of diversity...should we preserve several small areas, or a single large one?

• Large > Small

• Minimize Edge

• Clumped

• CorridorsWilson and Willis (1975)

- Why is this important?

- Conserving Diversity: the SLOSS debate

Num

ber o

f Spe

cies

Area in Square Meters

- Why is this important?

- Conserving Diversity: the SLOSS debate

Simberloff and Gotelli (1983)

BUT! Can we maintain all the species if they live on different islands?

A B

C D

E F

G H

A B

C D

E F G H

All species preserved while accommodating the species area effect!

- BUT! Can we maintain all the species if they live on different islands?

- Probably not, because communities are NESTED.

A B

C D

E F

G H

A B

C D

E F G H

- BUT! Can we maintain all the species if they live on different islands?

- Probably not, because communities are NESTED.

Nested Subset Structure: Species on species-poor islands are also found on species-rich islands.

A B

C D

E F

G H

A B

C D

E F G H

- BUT! Can we maintain all the species if they live on different islands?

- Probably not, because communities are NESTED.

Nested Subset Structure: Species on species-poor islands are also found on specie-rich islands.

A B

C D

E F

G H

A B

C D

E F G H

NOT NESTED

A A

A B

A B C A

NESTED

Community Ecology

I. Introduction

II. Multispecies Interactions with a Trophic Level

III. Multispecies Interactions across Trophic Levels

IV. Succession

V. Biodiversity: Patterns and Processes

A.The Species-Area Relationship

1. The pattern

2. The Theory of Island Biogeography

3. Why is this important? Fragmentation

4. The SLOSS debate

5. Nestedness

NESTED-SUBSET STRUCTURE:(Darlington 1957, Patterson and Atmar 1986)

- Why is this important?

- Conserving Diversity: the SLOSS debate - "Nestedness" (Darlington (1957); Patterson and Atmar (1986)

Communities are ‘nested’ if the Communities are ‘nested’ if the species in depauperate species in depauperate assemblages are also found in assemblages are also found in progressively more species rich progressively more species rich communitiescommunities

A B C D E F G H I J K L M N O P Q R S T U V W X Y ZA B C D E F G H I J K L M N O P Q R S T U V W X Y Z2626

A B C D E F G H I J K L M N O P Q R S T U V W X A B C D E F G H I J K L M N O P Q R S T U V W X 2424

A B C D E F G H I J K L M N O A B C D E F G H I J K L M N O ++ Q R S T U V W X Q R S T U V W X 2323

A B C D E F G H I J K L M N O P Q R S T U A B C D E F G H I J K L M N O P Q R S T U ++2121

A B C D E F G H I J K L M A B C D E F G H I J K L M ++ O P Q R S T O P Q R S T ++1919

A B C D E A B C D E ++ G H I J K G H I J K + ++ + N O P N O P ++ 1313

A B C D E A B C D E ++ G G ++ I J K I J K + ++ + N O N O ++ Q Q ++ V V 1414

A B C D E A B C D E + ++ + H I J K L H I J K L ++ N O N O ++ ++ 1212

A B C D E F G H A B C D E F G H + ++ +K L K L + ++ + O P O P ++ 1111

A B C D E F G A B C D E F G ++ I J K I J K + + + + ++ + + + + 1010

A B C D E F G H I J A B C D E F G H I J + + + + ++ + + + + 1010

A B C D E F G A B C D E F G + + ++ + + ++ M M ++ R R 9 9

A B C D E F A B C D E F ++ H H + ++ + L L + ++ + P P 9 9

A B C A B C ++ E E ++ I J I J ++ N N 7 7

A B C D E F I A B C D E F I ++ 77

A B C D E F A B C D E F ++ M M 7 7

A B C D E A B C D E ++ G G ++ M M 7 7

A B C D E F H A B C D E F H ++ 77

A B C D E F H A B C D E F H ++ 77

A B C D E FA B C D E F ++ 66

A B C D A B C D + + F LF L 6 6

A B C D E A B C D E ++ 55

A B C A B C + ++ + F L F L 5 5

A B A B ++ D E D E 4 4

A B A B + + F F 3 3

A B A B ++ 22

CC 1 1

CC 1 1

A B C D E F G H I J K L M N O P Q R S T U V W X Y ZA B C D E F G H I J K L M N O P Q R S T U V W X Y Z2626

A B C D E F G H I J K L M N O P Q R S T U V W X A B C D E F G H I J K L M N O P Q R S T U V W X 2424

A B C D E F G H I J K L M N O A B C D E F G H I J K L M N O ++ Q R S T U V W X Q R S T U V W X 2323

A B C D E F G H I J K L M N O P Q R S T U A B C D E F G H I J K L M N O P Q R S T U ++2121

A B C D E F G H I J K L M A B C D E F G H I J K L M ++ O P Q R S T O P Q R S T ++1919

A B C D E A B C D E ++ G H I J K G H I J K + ++ + N O P N O P ++ 1313

A B C D E A B C D E ++ G G ++ I J K I J K + ++ + N O N O ++ Q Q ++ V V 1414

A B C D E A B C D E + ++ + H I J K L H I J K L ++ N O N O ++ ++ 1212

A B C D E F G H A B C D E F G H + ++ +K L K L + ++ + O P O P ++ 1111

A B C D E F G A B C D E F G ++ I J K I J K + + + + ++ + + + + 1010

A B C D E F G H I J A B C D E F G H I J + + + + ++ + + + + 1010

A B C D E F G A B C D E F G + + ++ + + ++ M M ++ R R 9 9

A B C D E F A B C D E F ++ H H + ++ + L L + ++ + P P 9 9

A B C A B C ++ E E ++ I J I J ++ N N 7 7

A B C D E F I A B C D E F I ++ 77

A B C D E F A B C D E F ++ M M 7 7

A B C D E A B C D E ++ G G ++ M M 7 7

A B C D E F H A B C D E F H ++ 77

A B C D E F H A B C D E F H ++ 77

A B C D E FA B C D E F ++ 66

A B C D A B C D + + F LF L 6 6

A B C D E A B C D E ++ 55

A B C A B C + ++ + F L F L 5 5

A B A B ++ D E D E 4 4

A B A B + + F F 3 3

A B A B ++ 22

CC 1 1

CC 1 1

NESTEDNESSNESTEDNESS (Patterson and Atmar 1986)(Patterson and Atmar 1986)

Goby 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1Gudgeon 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1Catfish 1 1 1 1 1 1 1 1 1 1 1 1 1 1Hardyhead 1 1 1 1 1 1 1 1 1 Perch 1 1 1 1 1 1 1

NESTEDNESS AND NICHE SPACENESTEDNESS AND NICHE SPACE(Kodric-Brown and Brown 1993)(Kodric-Brown and Brown 1993)

NESTEDNESS AND NICHE SPACE(Kodric-Brown and Brown 1993)

GobyGudgeon

Catfish

Hardyhead

Perch

- Why is this important?

- Conserving Diversity: the SLOSS debate - "Nestedness" (Darlington (1957); Patterson and Atmar (1986)

- Fragmentation causes decreased diversity

- non-random loss of predators

- subsequent declines – keystone effects

- Why is this important?

- Conserving Diversity: the SLOSS debate - "Nestedness" (Darlington (1957); Patterson and Atmar (1986)

- Fragmentation causes decreased diversity

- increased stress decreases diversity non-randomly

• Small (4-6g)

• Medium (10-15g)

• Large (21-32g)

MYCOPHAGOUS FLY COMMUNITIES:SPECIES-AREA AND NESTEDNESS PATTERNS

(Worthen, Carswell and Kelly 1996)

L L L L L L L L L L L L L L L

MMMMM M MM MMMMMMM

S S S S S S S S

RICHNESS 5 5 4 4 4 4 4 4 4 4 4 4 4 3 3 3 3 2 2 2 2 2 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1

Phorid sp. 1 1 1 1 + 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

D. putrida 1 1 1 + 1 1 1 1 1 1 1 1 1 + 1 1 1 1 1 1 1 1

L. varia + 1 1 1 1 1 1 1 1 1 1 1 1 1 + 1 1 + + + + + 1 1 1 1

D. tripunctata 1 1 + + + + + + 1 1 1 1 1 1 1

M. dimidiata 1 1 1

D. falleni + + 1 1

Muscid sp. + + 1 1

Leptocera sp. ‘B’ 1 + 1

Leptocera sp. ‘A’ + + 1 1

MYCOPHAGOUS FLY COMMUNITIES:SPECIES-AREA AND NESTEDNESS PATTERNS

(Worthen, Carswell and Kelly 1996)

MYCOPHAGOUS FLY COMMUNITIES:DIFFERENCES IN DESICCATION TOLERANCE?

(Worthen and Haney 1999)

39.0

39.5

40.0

40.5

41.0

41.5

13 15 20 25 30 33

ACCLIMATION TEMPERATURE (oC)

39.0

39.5

40.0

40.5

41.0

41.5

13 15 20 25 30 33

ACCLIMATION TEMPERATURE (oC)

D. putrida

D. falleni

D. tripunctata

7 6 6 6 5 5 5 5 5 4 4 4 4 4 4 4 3 3 3 3 3 3 3 3 3 2 2 2 2 2 2 2 2 2 2 2 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

L. varia 1 x x 1 1 1 1 1 1 1 1 x 1 x 1 1 1 x 1 1 x 1 1 1 1 1 x x x x x 1 x 1 x x 1 1 1 1 1 1 1 1 1 1 1 1

Phorid x 1 1 1 1 x x x 1 x 1 1 1 1 1 x 1 x x 1 1 1 x x 1 x x x x x x x 1 x 1 1 1 1 1 1 1 1

D. falleni 1 x 1 1 1 1 1 1 1 x x x 1 x 1 1 x 1 1 x x x 1 1 x 1 1 1 1

Muscid 1 1 1 1 x 1 1 1 1 x x x x 1 x 1 x x x x 1 x x 1 1 1 1

D. putrida 1 x 1 x 1 1 1 1 1 1 x x x x x 1 x 1 x x x x 1 x x 1 1 1

Mycetophilid x x x 1 x x x 1 x x x x x x 1 x x x x x x 1 x x x x x x 1 x x 1 1 x x x x x 1 1 1 1

S. alternatus 1 1 1 1 x 1 1 x x 1 x x x 1 x x x x x x x x x x x x 1 x x x x x x x x x x x 1

Tipulid 1 1 1 x x x x x x x 1 1 x 1 x x x x x x 1 x x x x x x 1 x x x x x x x x x x 1

D. tripunctata x x x x 1 x x x x x 1 x 1 x x x 1 1 1

M. dimidiata x x x x x x x x x x 1 1 x x x x x x x x x x x x x 1

Cecidomyiid 1 1 x x x x x x x x x x x x x x x x x x x x x x x 1

D. neotest. x 1 x x x x x x x 1 x x x x x x 1

N = 231 P&A (1986) RANDOM1 = 265.4 + 23.4 z = -1.45 ns

EFFECT OF DESICCATION ON NESTEDNESS(Worthen, Jones and Jetton 1998)

4 4 3 2 2 2 2 2 2 2 2 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

L.varia 1 x 1 1 x x 1 1 1 1 1 1 1 1 1 1 1 1 1 1

Phorid x 1 x x 1 1 x x x 1 1 1 1 1 1 1 1 1 1 1 1

Muscid 1 x x x x x 1 1 1 x x x x 1 1 1

Tipulid x 1 1 1 1

D. putrida 1 1 x 1

M. dimidiata x x 1

Psychodid 1

S. alternatus 1

N = 22 P&A (1986) RANDOM1 = 45.1 + 10.5 z = -2.15*

EFFECT OF DESICCATION ON NESTEDNESS(Worthen, Jones and Jetton 1998)

THE EFFECT OF AN INDUSTRIAL SPILL ON THE MACROFAUNA OF THE UPPER ENOREE

(Worthen, Haney, Cuddy, Turgeon and Andersen 2001)

The Upper Enoree River

NESTEDNESS, STRESS, AND THE STRUCTURE OF LARVAL ODONATE ASSEMBLAGES

R1 = 10.41 + 3.78;

x + 1.96 sd = 2.59 to 17.82

N = 4, ns

Not nested

C9 C8 T4 C7 C6 C5 C4 T1 T2 T3 C3Distance 7.2 6.3 4.7 3.4 2.4 1.8 0.7Richness 7 5 3 3 3 3 3 2 1 1 1

NCordulegaster 1 1 1 1 1 1 1 1 1 1 0Progomphus 1 1 1 1 1 1 1 0Macromia 1 1 1 1 1 1 0Boyeria 1 1 x x x 1 x 1 1 4Gomphus 1 1 0Stylurus 1 0Hagenius 1 0

R1 = 3.00 + 1.50;

x + 1.96 sd = 0.04 to 8.98

N = 0, p < 0.05

Nested

C9 C8 C7 C6 C5 C4 C3Distance 7.2 6.3 4.7 3.4 2.4 1.8 0.7Richness 7 5 3 3 3 3 1

NCordulegaster 1 1 1 1 1 1 1 0Progomphus 1 1 1 1 1 1 0Macromia 1 1 1 1 1 0Boyeria 1 1 1 0Gomphus 1 1 0Stylurus 1 0Hagenius 1 0

NESTEDNESS, STRESS, AND THE STRUCTURE OF LARVAL ODONATE ASSEMBLAGES

Summary: Causes of nestedness - nested niche space

- differences in dispersal capabilities

-differences in extinction probabilities

As these are the same factors that cause the species-area relationship, itself, we should not be surprized that communities distributed across habitats of different size are often nested, too.