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? - Why is this important? - Conserving Diversity: the SLOSS debate • Large > Small • Minimize Edge • Clumped • Corridors Wilson and Willis (1975) - Why is this important? Number of Species - 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 B C D E F G H A All species preserved while accommodating the species area effect! B CD EFG H - 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 CD EFG 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 speciespoor islands are also found on species-rich islands. A B C D E F G H A B CD EFG 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 speciespoor islands are also found on specie-rich islands. NOT NESTED A B C D E F G H A B A CD EFG NESTED A AB H ABC A 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 - Why is this important? - Conserving Diversity: the SLOSS debate NESTED-SUBSET STRUCTURE: (Darlington 1957, Patterson andand Atmar - "Nestedness" (Darlington (1957); Patterson Atmar 1986) (1986) A B C D E Communities are ‘nested’ if the species in depauperate assemblages are also found in progressively more species rich communities A B C A A B CD NESTEDNESS (Patterson and Atmar 1986) ABCDEFGHIJKLMNOPQRSTUVWXYZ ABCDEFGHIJKLMNOPQRSTUVWX ABCDEFGHIJKLMNO+QRSTUVWX ABCDEFGHIJKLMNOPQRSTU+ ABCDEFGHIJKLM+OPQRST + ABCDE+GHIJK++ NOP + ABCDE+G+IJK++ NO+Q+ V ABCDE++ HIJKL+ NO+ + A B C D E F G H + +K L + + O P + ABCDEFG+IJK ++ + + + ABCDEFGHIJ ++ + + + ABCDEFG+++ +M+ R ABCDEF+H++ L ++ P ABC+E+ IJ + N ABCDEF I + ABCDEF +M ABCDE+G +M ABCDEF H + ABCDEF H + ABCDEF + ABCD+F L ABCDE+ ABC ++F L AB+DE AB + F AB + C C 26 24 23 21 19 13 14 12 11 10 10 9 9 7 7 7 7 7 7 6 6 5 5 4 3 2 1 1 NESTEDNESS AND NICHE SPACE (Kodric-Brown and Brown 1993) Goby Gudgeon Catfish Hardyhead Perch 111111111111111111111111111 111111111111111111 11111111111111 111111 111 1111111 NESTEDNESS AND NICHE SPACE (Kodric-Brown and Brown 1993) Goby Catfish Gudgeon 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) LLLLLLL LLL LLL MMMMM S L M MM SS L MMMMMMM S SSSS RICHNESS 554444444444433332222 22211111111111111 Phorid sp. D. putrida L. varia D. tripunctata M. dimidiata D. falleni Muscid sp. Leptocera sp. ‘B’ Leptocera sp. ‘A’ 1111+1111111111111111 111 111111111111 111+111111111+1111111 1 +1 1 1 1 1 11 1 1 1 1 11 + 1 1+ ++ ++ 1 1 11 11++++++1111111 11 1 ++ 1 1 ++ 1 1 1+ 1 ++ 1 1 MYCOPHAGOUS FLY COMMUNITIES: DIFFERENCES IN DESICCATION TOLERANCE? (Worthen and Haney 1999) 41.5 D. putrida 41.0 40.5 D. falleni 40.0 D. tripunctata 39.5 39.0 13 15 20 25 30 33 o ACCLIMATION TEMPERATURE (oC) EFFECT OF DESICCATION ON NESTEDNESS (Worthen, Jones and Jetton 1998) 76 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 1x x1 1 1 1 1 1 1 1x1 x 1 1 1 x 1 1 x 1 1 1 1 1x x x x x 1 x 1 x x1 1 1111111111 x1 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 1111 1x 1 1 1 1 1 1 1x x x1 x 1 1 x1 1 x x x 1 1 x 11 11 11 1 1x 1 1 1 1x x xx 1 x1 xx xx 1 xx 1 1 1 1 1x 1 x 1 1 1 1 1 1 x x x x x 1 x1 x x x x1 x x 111 Mycetophilid xx x1 x x x 1 x x x x x x 1 x x x x x x 1 x x x x x x1 x x 1 1 x x x x x 1111 S. alternatus 11 1 1x 1 1x x1 xx x 1 x xx xx xx x x xx x1 x xx xx x xx xx x 1 Tipulid 11 1xx x xx xx 1 1 x 1 xx xx xx 1 x x xx xx 1 x xx xx x xx xx 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. x1 x x x x x x x 1 x x x x x x1 N = 231 P&A (1986) RANDOM1 = 265.4 + 23.4 z = -1.45 ns EFFECT OF DESICCATION ON NESTEDNESS (Worthen, Jones and Jetton 1998) 4432222222222111111111111111111 L.varia Phorid Muscid Tipulid D. putrida M. dimidiata Psychodid S. alternatus N = 22 1x11xx1111111 1111111 x1xx11xxx 1111 11111111 1xxxxx111 xxxx111 x11 11 11x1 xx1 1 1 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 C9 C8 T4 Distance Richness 7.2 7 6.3 5 C7 C6 C5 C4 T1 3 4.7 3 3.4 3 2.4 3 1.8 3 T2 2 T3 1 C3 1 0.7 1 N Cordulegaster Progomphus Macromia Boyeria Gomphus Stylurus Hagenius 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 x 1 1 1 x 1 1 1 x 1 1 1 1 1 1 x 1 1 1 1 R1 = 10.41 + 3.78; N = 4, ns x + 1.96 sd = 2.59 to 17.82 Not nested 1 0 0 0 4 0 0 0 NESTEDNESS, STRESS, AND THE STRUCTURE OF LARVAL ODONATE ASSEMBLAGES C9 C8 C7 C6 C5 C4 C3 Distance Richness 7.2 7 6.3 5 4.7 3 3.4 3 2.4 3 1.8 3 0.7 1 N Cordulegaster Progomphus Macromia Boyeria Gomphus Stylurus Hagenius 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 R1 = 3.00 + 1.50; N = 0, p < 0.05 x + 1.96 sd = 0.04 to 8.98 Nested 0 0 0 0 0 0 0 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.