Ecological communities Guild Community Assembly A community is a local assembly of species that potentially interact. Generally these species are on the same trophic level. A community of species with similar niches is called an ecological guild Counter examples Examples of ecological communities Insect eating birds in a forest. Fish in a pond. Butterflies on a meadow. Birds in a forest form an assembly Aphids and Ladybeetles are on different trophic levels. Fish in an archipelago form metacommunities. Examples of communities Plant visitors and pollinators Nepenthes pitcher plants Deep sea bacterial communities Calcareous grassland Assemblages but not communities Mutual effects of species interections: Direct effects refer to the impact of the presence (or change in abundance) of species A on species B in a two-species interaction. Indirect effects refer to the impact of the presence (or change in abundance) of species A on species C via an intermediary species (A --> B --> C). Cascading effects are those which extend across three or more trophic levels, and can be top-down (predator --> herbivore --> plant) or (plant --> herbivore --> predator). Keystone species are those which produce strong indirect effects. Kelp (brown algae) forest The starfish Pisaster predates on Mytilus mussels and makes space for many other species to colonise. The top predator Pistaster is a keystone species. _ _ _ + Fisherig Sea otter (Enhydra) Sea urchin (Echinus) _ Abiotic and isolation filters at different spatial scales determine local species composition Global species pool Isolation filter Abiotic filters Abiotic filters Local species pools Community Regional species pools Members of single communities pass these filters. Abiotic filters The distribution of species abundances Rank – abundance plot Relative abundances in a sequence of plant succession (Bazzaz 1975) In natural comunities species abundances often differ by factors of more than 1000. That means that the most abundant species are 1000 times more abundant than the rare species Three types of relative abundance distributions (SAD) Log-series SAD Heavy tail π = π0 π −π Log abundance Log abundance Power function SAD ππ π(π) = πΌ π Ronald A. Fisher (1890-1962) P(N) is the probabiity that a species has exctly N individuals Species rank order i Species rank order i Robert May of Oxford (1938-) Log abundance Log-normal SAD π = π −π×ππππ(0,1) Species rank order i Frank W. Preston (1896-1989) Log abundance Power function SAD Parasitic Hymenoptera in a beech forest Heavy tail π = π0 π −π Species rank order i Power function SADs • have a high number of rare species (heavy • • • tail) are input (colonization) driven have a high degree of species turnover often characterize species assemblages but not true communities have a small number of very abundant species lack a larger number of intermediate abundant species Examples • Incomplete samples • Arthropod assemblages • Disturbed habitats Log abundance Log-series SAD Northern German Grassland spiders (Finch 2001) ππ π(π) = πΌ π Species rank order i The log series is a sample distribution. It describes the expected abundance of species in a sample from a large community. It applies to assemblages. For fully censused assemblages it occurs most often • at early stages of succession • in disturbed habitats • in heterogeneous assemblages Examples • Incomplete samples • Heterogeneous assemblages • Large arthropod samples Beetles in a Norwegian spruce forest (Ottesen 1996) Log abundance Log-normal SAD High number of species with intermediate abundance π = π −π×ππππ(0,1) Species rank order i Breeding birds of Ohio (Hicks 1935) π = π0 π −ππ veil line 2 Lognormal SADs are derived from the central limit theorem of statistics that predict normal distributions Lognormal distributions occur most often in • closed and stable communities • undisturbed habitats • K- species dominated communities • Communities influenced by a large number of divergent environmental factors. Often the distribution is not symmetrical having an excess of rare species. Diversity and evennness A measure of diversity is the number of species Abundance High evenness Simpson index of diversity π»= 1 π 1 ππ Shannon index of diversity π π»=− ππ ln(ππ ) 1 Log-series index of diversity Species Abundance Lower evenness π π = π»ππ(1 + ) π» Diversity indices are measures of encounter probability Evenness π» πΈ= πππ Species Alpha, beta and gamma diversity Species richness Alpha diversity refers to the local number of species Beta diversity refers to the change in species composition among local habitats Gamma diversity refers to the regional species pool g b a π¦ = πΌπ΄π½ Multiplicative partitioning of diversity πΎ =πΌ×π½ Additive partitioning of diversity πΎ =πΌ+π½ Area Beta diversity is a measure of regional habitat diversity Species interactions or neutrality Pool of individuals Stephen P. Hubbell (1942- Birth Death Neutral models are individual based! Random migration Ecological drift Zero sum multinomial Motoo Kimura (1924-1994) Mutations Individuals are grouped to evolutionary lineages Species are ecologically equivalent Ji ο½ J 0 ο« iJ 0 ο« bJ 0 ο eJ 0 ο dJ 0 ο½ J 0 J is the total number of individuals Neutral models lack any specific biological interaction like competition, mutualism, regulation, species specific survival. Neutral models provide ecological expectations without species interaction Neutral models make explicit predictions about Abundance rank order relationships Diversity and evenness Abundance 10000 Core species 1000 100 10 1 Leistus rufomarginatus 0 Photos by Roy Anderson 5 10 15 20 25 30 Rank order 100 Abundance Satellite species 10 1 0.1 Ground beetles on lake islands in Lake Mamry (Ulrich and Zalewski 2007) 0 5 10 15 20 Rank order 25 30 Neutral models make explicit predictions about Core and satellite species 35 30 Predicted Species 25 20 Observed 15 10 5 0 1 2 4 Sites occupied Dyschirius globosus Ground beetles on lake islands in Lake Mamry (Ulrich and Zalewski 2007) Regional diversity patterns Observed Predicted 8 15 Ecological gradients and the classification of communities Species 1pog 2pog 3pog dab Pterostichus melanarius 0 36 13 17 Pterostichus oblongopunctatus 0 2 7 135 Pterostichus niger 0 0 3 0 Oxypselaphus obscurus 1 0 27 96 Harpalus 4-punctatus 0 41 17 9 Carabus granulatus 0 11 52 11 Patrobus atrorufus 0 6 22 81 Pterostichus antracinus 11 1 0 0 Platynusas similis 0 7 25 4 Pterostichus nigrita 30 2 2 5 Carabus hortensis 0 0 0 0 Pterostichus strennus 0 5 3 47 Distance matrix ful 187 83 191 27 29 12 11 0 48 1 75 13 gil 345 188 167 166 77 110 348 21 39 58 52 30 guc 60 11 135 80 0 25 9 1 2 1 109 5 hel kor lip 169 1199 704 8 1019 180 0 137 0 0 96 7 67 555 69 11 154 113 0 11 37 11 2 2 9 76 117 0 0 2 0 0 0 6 28 24 mil 394 4 0 48 0 0 0 274 0 39 0 22 sos 428 141 530 278 9 59 35 0 9 18 0 14 swi 13 1 3 0 0 1 2 0 0 0 0 4 Ground beetles from Mazuran lake lands The first two eigenvectors Species Pterostichusoblongopunctatus(Fabricius) Pterostichusmelanarius Pterostichusniger(Schaller) Oxypselaphusobscurus(Herbst) Harpalus4-punctatusDejean Carabusgranulatus Patrobusatrorufus(Stroem) Pterostichusantracinus Platynusassimilis(Paykull) Pterostichusnigrita(Paykull) CarabushortensisLinnaeus Pterostichusstrennus(Panzer) Pterostichusmelanarius 0 787.7 1363 1393 1122 1359 1494 1515 1442 1541 1550 1518 Pterostichusoblongopunctatus(Fabricius) 787.7 0 1004 955.5 530.5 887.8 1040 1101 975.5 1062 1066 1030 Pterostichusniger(Schaller) 1363 1004 0 327.5 714.1 533.9 591.8 671 586.5 591 571.9 590.6 Oxypselaphusobscurus(Herbst) 1393 955.5 327.5 0 563.3 281.7 328.9 418.8 344.1 324.4 339 320.4 Harpalus4-punctatusDejean 1122 530.5 714.1 563.3 0 415.3 618.4 628.2 488 567.8 578 538 Carabusgranulatus 1359 887.8 533.9 281.7 415.3 0 299.4 354.5 127.5 215.1 239.6 192 Principal Patrobusatrorufus(Stroem) 1494 1040 591.8 328.9 618.4 299.4 0 438.2 338 307.4 333.9 322.6 Pterostichusantracinus 1515 1101 671 418.8 628.2 354.5component 438.2 0 311.9 239.7 305.9 259.9 Platynusassimilis(Paykull) 1442 975.5 586.5 344.1 488 127.5 338 analysis 311.9 0 157.2 180.7 123 Pterostichusnigrita(Paykull) 1541 1062 591 324.4 567.8 215.1 307.4 239.7 157.2 0 141.3 72.41 CarabushortensisLinnaeus 1550 1066 571.9 339 578 239.6 333.9 305.9 180.7 141.3 0 139.6 Pterostichusstrennus(Panzer) 1518 1030 590.6 320.4 538 192 322.6 259.9 123 72.41 139.6 0 EV 1 0.648 0.368 -0.059 -0.179 0.036 -0.207 -0.202 -0.218 -0.240 -0.276 -0.264 -0.270 EV 2 0.496 0.361 0.298 0.240 0.262 0.218 0.262 0.267 0.228 0.237 0.244 0.232 Principal component analysis PCA serves to identify ecological communities Communities as ecological indicators Ecological indicators are used to provide information about the state and the functioning of ecological systems. Indicators might be single species , sets of species or whole communities. Often used indicators Anthropogenic disturbance Acid rain Eutrophication Invasive species Sedimentation Logging Heavy metals Urbanization Air pollution Air quality Indicator Mosses Aquatic macrophytes Birds Shrubs Bark beetles Protozoa Birds, Carabids Plants Lichen