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BIODIVERSITY

"species - area relationship"

S = CA z log 10 S = log 10 C + z log 10

where C is the y intercept and z is the slope of the line.

"species - area relationship"

Breedings Birds - North Am.

"species - area relationship"

Island Area log(square km)

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

MacArthur and Wilson (1967) THEORY OF ISLAND BIOGEOGRAPHY

Edward O. Wilson Prof. Emer., Harvard Robert MacArthur 1930-1972

MacArthur and Wilson (1967) THEORY OF ISLAND BIOGEOGRAPHY - Species Richness is a balance between

COLONIZATION (adds species) and EXTINCTION (subtracts species)

- Colonization Increases with Area - larger target - more habitats

Mainland

confirmation: greater immigration rate on larger islands

- Colonization Increases with Area - larger target - more habitats

- Colonization Increases with Area - larger target - more habitats (except very small)

Niering, W.A. 1963. Terrestrial ecology of Kapingamarangi Atoll, Caroline Islands. Ecological Monographs 33:131-160.

- Colonization Increases with Area - larger target - more habitats

- Extinction Decreases with Area - more food means larger populations that are less likely to bounce to a size of "0" (extinction)

- Extinction Decreases with Area

Reduced Turnover on larger islands

Wright, S.J. 1980. Density compensation in island avifaunas. Oecologia 45: 385 389. Wright, S. J. 1985. How isolation affects rates of turnover of species on islands. Oikos 44:331-340.

COL - large COL - small SMALL species richness LARGE EXT - small EXT - large

- Colonization Decreases with Distance - fewer species can reach

Mainland

- Colonization Decreases with Distance - fewer species can reach

saturation is the % of species found on a patch of mainland that size

- Extinction Increases with Distance - recolonization less likely at distance

Mainland

"Rescue Effect"

- Extinction Increases with Distance - recolonization less likely at distance

Wright, S.J. 1980. Density compensation in island avifaunas. Oecologia 45: 385 389. Wright, S. J. 1985. How isolation affects rates of turnover of species on islands. Oikos 44:331-340.

COL - close COL - far far species richness close EXT - far EXT - close

Equilbrium Island Biogeography & Turnover

Turnover on "Landbridge" islands (California Channel Islands)

equilibria Island

Los Coronados San Nicholas San Clemente Santa Catalina Santa Barbara San Miguel Santa Rosa Area km2 2.6

Distance km Bird Spp. 1917 13 11 Bird Spp. 1968 11 Extinctio ns Human Introd.

4 0 Immigratio ns Turnover % 4 36 57 145 194 2.6

36 218 98 79 32 61 42 44 11 28 30 10 11 14 11 24 34 6 15 25 6 9 6 7 4 1 2 1 1 0 0 1 4 4 9 3 8 11 50 25 24 62 46 32 Santa Cruz 249 31 36 37 6 1 5 17 Anacapa 2.9

21 15 14 5 0 4 31 Diamond, J.M. 1969. Avifaunal equilibria and species turnover rates on the Channel Islands of California. Proc. Natl. Acad. Sci 64: 57-63. Jones, H.L. and Diamond, J.M. 1976. Short-time-base studies of turnover in breeding bird populations on the Channel Islands of California. Condore 73: 526-549. [ + ]

Equilbrium Island Biogeography & Turnover

Turnover on "Landbridge" islands (California Channel Islands)

equilibria and turnover Island

Los Coronados San Nicholas San Clemente Santa Catalina Santa Barbara San Miguel Santa Rosa Area km2 2.6

Distance km Bird Spp. 1917 13 11 Bird Spp. 1968 11 Extinctio ns Human Introd.

4 0 Immigratio ns Turnover % 4 36 57 145 194 2.6

36 218 98 79 32 61 42 44 11 28 30 10 11 14 11 24 34 6 15 25 6 9 6 7 4 1 2 1 1 0 0 1 4 4 9 3 8 11 50 25 24 62 46 32 Santa Cruz 249 31 36 37 6 1 5 17 Anacapa 2.9

Dramatic evidence that, although the communities had recovered in terms of species richness, the composition was very different with typically about 80% of the species turning over.

- Why is this important?

- all habitats except the atmosphere are islands.

Continents big islands

White-faced Saki (Pithecia pithecia)

Monk Saki (Pithecia monachus) White-faced Saki (Pithecia pithecia)

Monk Saki (Pithecia monachus) White-footed Saki (Pithecia albicans) White-faced Saki (Pithecia pithecia)

Monk Saki (Pithecia monachus) White-footed Saki (Pithecia albicans) White-faced Saki (Pithecia pithecia) Rio Tapajos Saki (Pithecia irrorata)

Minnesota: Land O'Lakes

"Sky Islands" High elevation habitats separated by inhospitable (desert) habitat.

- Why is this important?

- all habitats except the atmosphere are islands.

- human activity fragments a landscape, making lots of islands, too.

Bolivia has lost 50% of its rainforest in last 30 years

Even Costa Rica has lost 95% of its old growth forest that is outside of national parks...

- 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?

- Why is this important?

- Conserving Diversity: the SLOSS debate

• Large > Small • Minimize Edge • Clumped • Corridors Wilson and Willis (1975)

- Why is this important?

- Conserving Diversity: the SLOSS debate Area in Square Meters Simberloff and Gotelli (1983)

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

different

islands?

A C E G B D F H A C D B 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 C E G B D F H A C D B 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 C E G B D F H A C D E F G B H

A C E G - 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.

NOT NESTED NESTED B D F H A C D E F G B H A A B A B C A A

- Why is this important?

- "Nestedness" (Darlington (1957); Patterson and Atmar (1986)

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

A B C D E A B C A A B C D

NESTEDNESS

(Patterson and Atmar 1986) 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 Z 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 + 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 + A B C D E F G H I J K L M + O P Q R S T + A B C D E + G H I J K + + A B C D E + G + I J K + + N O P + N O + Q + A B C D E + + H I J K L + N O + + V A B C D E F G H + + K L + + O P + A B C D E F G + I J K + + + + + A B C D E F G H I J + + + + + A B C D E F G A B C D E F + + + + H + + + M + L + + P R A B C + E + I J + A B C D E F I + A B C D E F + M N A B C D E + G + M A B C D E F H + A B C D E F H + A B C D E F + A B C D + F L A B C D E + A B C + + F L A B + D E A B + F A B + C C 7 7 7 6 6 5 5 4 3 2 1 1 26 24 23 21 19 13 14 12 11 10 10 9 9 7 7 7

NESTEDNESS AND NICHE SPACE

(Kodric-Brown and Brown 1993) Goby Gudgeon Catfish Hardyhead Perch 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 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 1 1 1 1 1

NESTEDNESS AND NICHE SPACE

(Kodric-Brown and Brown 1993) Goby Gudgeon 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

MYCOPHAGOUS FLY COMMUNITIES: SPECIES-AREA AND NESTEDNESS PATTERNS (Worthen, Carswell and Kelly 1996) • 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.

D. putrida L. varia D. tripunctata M. dimidiata D. falleni

Muscid sp.

Leptocera

sp. ‘B’

Leptocera

sp. ‘A’ 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 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 + + + + + 1 1 1 1 1 1 + + + + + + 1 1 1 1 1 1 1 1 1 1 + + + + 1 + 1 1 1 1 1 + + 1 1

MYCOPHAGOUS FLY COMMUNITIES: DIFFERENCES IN DESICCATION TOLERANCE? (Worthen and Haney 1999)

D. putrida D. falleni D. tripunctata

ACCLIMATION TEMPERATURE ( o C)

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

Phorid

D. falleni

Muscid

D. putrida

Mycetophilid

S. alternatus

Tipulid

D. tripunctata

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 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 1 x 1 1 1 1 1 1 1 x x x 1 x 1 1 1 1 x 1 1 1 1 x x x x 1 1 x 1 1 x x x 1 x 1 x x x x 1 1 1 x x x 1 1 1 1 1 1 1 1 1 x 1 x 1 1 1 1 1 1 x x x x x x x x 1 x x x 1 x x x x x x 1 1 x 1 x x x x x x x x x x 1 x x 1 1 1 1 x x x x x x 1 x x 1 1 x x x x x 1 1 1 1 1 1 1 x 1 1 x x 1 x x x x x x x x x x x 1 x x x x x x x x 1 1 1 x x 1 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 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 x x x 1 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 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

D. neotest.

x 1 x x x x x x x 1 x x x x x x 1 1 1 1 1 1 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

Phorid Muscid Tipulid

D. putrida M. dimidiata

Psychodid

S. alternatus

1 x 1 1 x x 1 1 1 1 1 1 1 1 1 1 1 1 1 1 x 1 x x 1 1 x x x 1 1 1 1 1 1 1 1 1 1 1 1 1 x x x x x 1 1 1 x x x x 1 1 1 x 1 1 1 1 1 1 x 1 x x 1 1 1 N = 22 P&A (1986) RANDOM1 = 45.1 + 10.5 z = -2.15*

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

Distance Richness

Cordulegaster Progomphus Macromia Boyeria Gomphus Stylurus Hagenius

C9 7.2

7 C8 6.3

5 T4 3 1 1 1 1 1 1 1 C7 4.7

3 C6 3.4

3 1 1 1 1 1 1 1 1 1 1 1 1 1 x x x 1 C5 2.4

3 C4 1.8

3 T1 2 1 T2 1 T3 1 1 C3 0.7

1 N 1 1 1 1 1 1 1 x 1 1 0 0 0 4 0 0 0 R1 = 10.41 + 3.78; x + 1.96 sd = 2.59 to 17.82

N = 4, ns Not nested

NESTEDNESS, STRESS, AND THE STRUCTURE OF LARVAL ODONATE ASSEMBLAGES

Distance Richness C9 7.2

7 C8 6.3

5 C7 4.7

3 C6 3.4

3 C5 2.4

3 C4 1.8

3 C3 0.7

Cordulegaster Progomphus Macromia Boyeria Gomphus Stylurus Hagenius

R1 = 3.00 + 1.50; 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 N = 0, p < 0.05

1 N 0 0 0 0 0 0 0 x + 1.96 sd = 0.04 to 8.98

Nested

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.