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AP Environmental Science Community Interactions, Species Diversity and Succession Community Ecology Community = an association of interacting species inhabiting some defined area Community structure = includes attributes such as the number of species and the kinds of species comprising a community Community function = includes attributes such as energy flow through food webs Are ecological communities “real” functional units? Yes! Frederick E. Clements (1874 – 1945) Holistic community concept (Closed community) Community as a superorganism Based his conclusion on a study of vegetation zones along altitudinal gradients Basis of the modern-day “Gaia concept” Vegetation Zones Closed Communities Along an environmental gradient (e.g. temperature, moisture) there are sometimes sharp boundaries between species assemblages Areas between assemblages are known as ecotones Best seen where there are sharp physical boundaries between species assemblages Between aquatic and terrestrial areas Between distinct soil types Between north and south facing slopes Closed Communities No! Henry A. Gleason (1882-1975) Individualistic concept (Open community) The appearance of communities are simply due to the individual interactions of species and don’t reflect any innate organization above the species level Open Communities Closer examination of the deciduous forests of eastern North America revealed no sharp boundaries between forest types There were generally two gradients of forest types: A north-south gradient based on temperature An east-west gradient based on moisture Gradient analysis is a technique invented by Robert H. Whittaker to analyze changes in (plant) species abundance along gradients Gradients are variables that affect plant community structure (i.e temperature, light & water availability, soil nutrients, etc.) Put to rest the extreme closed community concept of Clements Open Communities Factors Affecting Biological Community Structure Species diversity – function of the number of different species that a community contains (species richness) and the relative abundance of individuals of each species (species evenness). Niche structure: J.H. Brown (1981) bases this on ‘capacity rules’ (how many potential ecological niches occur) and ‘allocation rules’ (how these ecological niches can be divided up among species). Geographical location Consistent climates foster high diversity but ‘specialist species’ Variable climates foster low diversity but ‘generalist species’ Species Richness Species Richness is a measure of the total number of different species in a community Observations regarding species richness: Gradual increase in diversity/richness Pole to Equator In a given ecosystem, species richness increases over time. Larger ecosystems tend to have greater species richness. Usually increases with ecological succession. Why these trends? Why These Trends? Productivity? Spatial complexity/heterogeneity? However, an aspect of environmental structure that is important to one group may not be important to another group Therefore, you must know something about the ecological requirements of species to predict how environmental structure affects their diversity. Species Richness Factors that affect species richness saturation rates of colonization and extinction (MacArthur & Wilson (1967) • Depend on size of ecosystem and • Distance from a source of colonists disturbance severity, variability, and predictability (Slobodkin and Sanders, 1969) • i.e. rainforests – favorable and constant; therefore predictable; polluted stream – severe and variable; therefore unpredictable Calculating Species Richness Generally exclude “accidental species” in the count Does not take into account abundance patterns among the various species Species counts depend on the sample size or sample area (rarely do you get every one!) “Equilibrium model of island biogeography” (MacArthur & Wilson, 1967) # of species (species richness) on an “island” is determined by immigration and emigration of species Larger islands and islands closer to “mainland” will have greater species richness. Equilibrium Model of Island Biogeography Species-Area Curves • Species-area curve • S = CAz • log S = log C + z log A • An equation for a straight line where log c = Y intercept and z = the slope of the line S Species Richness C constant A Area z constant Species Area Curves Diversity Indices Shannon-Weiner Index Also known as the “Shannon Index” or the “Shannon-Weaver Index” Based on Information Theory Measures the amount of order in a system in bits of information Determines rarity or commonness of species Combines measure or species richness with species evenness. Calculating the Shannon-Weiner Index H = -∑(pi) (ln pi) H = Shannon-Weiner Diversity Index pi = proportion of total sample belong to the ith species ln pi = the natural log of pi An Example Community A: 99 individuals in species 1; individual in species 2 “Uneven” community • H = -[(p1)(ln p1) + (p2)(ln p2)] • H = -[0.99 (ln 0.99) + 0.01 (ln 0.01)] • H = -[(-0.0099) + (-0.0461)] = +0.056 Community B: 50 individuals in species 1; 50 individuals in species 2 “Even” community • H = -[(0.5) (ln 0.5) + (0.5) (ln 0.5)] • H = -[(-0.3466) + (-0.3466)] = +0.693 1 Comparison of Shannon-Weiner Index Values for Even and Uneven Communities 3.5000 3.0000 2.5000 H 2.0000 1.5000 1.0000 0.5000 0.0000 0 5 10 15 # of species H - even community H- uneven community 20 25 Pielou’s Evenness Index Compares the evenness component of species diversity in a community with the maximum amount of evenness possible given the same species richness Also known as Shannon’s equitability (EH) EH = H/Hmax = H/ln S Hmax = ln S Community A--> J = 0.056/0.693 = 0.081 Community B--> J = 0.693/0.693 = 1.000 EH ranges from 0-->1.0 with 1.0 indicating maximum evenness Comparison of Pielou's Evenness Index for Even and Uneven Communities 1.2 1 1 1 1 1 0.0434 0.0338 0.0304 0.0285 Evenness 0.8 0.6 0.4 0.2 0 #of species 5 10 # of species EH - even community EH - uneven community 15 Use of Diversity Indices Used in applied ecology to measure differences in diversity between two or more communities Also used to measure changes in diversity within a community after some environmental modification or disturbance Robert MacArthur (1930-1972) UPenn & Princeton Studied the diversity of warblers in northeastern forests Discovered that different warbler species fed in different parts of the forest canopy MacArthur’s Warblers Bird Species Diversity vs. Foliage Height Diversity Also showed that the greater the Foliage Height Diversity of the forest the greater the Bird Species Diversity Ecological Succession Plant (and animal) communities develop in often predictable ways following disturbances, a process known as succession Natural, gradual changes in the types of species that live in an area Classified as primary or secondary succession Indiana Dunes Succcession Henry Chandler Cowles (1869-1939) American botanist and ecological pioneer Born in Kensington, Connecticut, he attended Oberlin College in Ohio and the University of Chicago He obtained his Ph.D. in 1898 for his study of vegetation succession on the Lake Michigan sand dunes. Kinds & qualitative characteristics of disturbances that impact communities Disturbance magnitude Asteroid impact Volcanism Glaciation Landslide Ice Storm Hurricane Fire Treefall Disturbance frequency Primary Succession General characteristics Begins in a place without any soil Sides of volcanoes Landslides Flooding Remnants of glaciers Starts with the arrival of living things such as lichens that do not need soil to survive Lichens (pioneer species) secrete weak acids which in turn break rocks into soil Called the Pioneer Stage Lichens: Symbiosis of a fungus & an alga Example of a mutualism Primary Succession General characteristics Soil starts to form as lichens and the forces of weather and erosion help break down rocks into smaller pieces When lichens die, they decompose, adding small amounts of organic matter to the rock to make soil Primary Succession General characteristics Simple plants like mosses and ferns can grow in the new soil Primary Succession General characteristics The simple plants die, adding more organic material The soil layer thickens, and grasses, wildflowers, and other plants begin to take over Called Seral Stages Primary Succession General characteristics These plants die, and they add more nutrients to the soil Shrubs and trees can survive now Still called Seral Stages Primary Succession General characteristics Insects, small birds, and mammals have begun to move in What was once bare rock now supports a variety of life Called a Climax Community Climax Community A stable group of plants and animals that is the end result of the succession process Does not always mean big trees Grasses in prairies Cacti in deserts Mosses, sedges and lichens in the tundra Secondary Succession Begins in a place that already has soil and was once the home of living organisms Occurs faster and has different pioneer species than primary succession Example: after forest fires Intermediate Disturbance Hypothesis J.P. Grime (1973), Henry Horn (1975) and Joseph Connell (1978) The Intermediate Disturbance Hypothesis is an ecological hypothesis which proposes that biodiversity is highest when disturbance is neither too rare nor too frequent. With low disturbance, competitive exclusion by the dominant species arises. With high disturbance, only species tolerant of the stress can persist. Role of Disturbance in Succession