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FWF 410: “Populations and Communities” (Molles 2002 after Gause 1934) Matthew J. Gray, Ph.D. College of Agricultural Sciences and Natural Resources University of Tennessee-Knoxville Goal of the Lecture To familiarize students with biological organization terms, population distributions and growth, and basic concepts in community ecology. Reading Assignments: No Assigned Reading for Lecture Lecture Structure I. Levels of Biological Organization What is an individual, population and community? II. Population Distributions and Growth How are individuals distributed in space and how do populations fluctuate through time? III. Community Ecology What factors influence multiple species assemblages? 1 Levels of Biological Organization Individual: A living single or multi-celled organism. Population: A collection of individuals of the same species that have a some probability* of interaction and influence. Community: A collection of populations of different species that have a some probability of interaction and influence. Population Biology Introduction Study of the distribution and abundance of organisms. Terms Population: Group of individuals of the same species that have some probability to interact and reproduce (without dispersing). Distribution: The size, shape, and location of the area where a population exists. Abundance: Number of individuals in a population per unit time, Nt . Density: Number of individuals per unit area (e.g., nt / m2) for a population. Why Study Populations? Populations provide an ecological entity of quantification for management & experiments. •Prudent human natural resource use (harvesting, non-consumptive use) Evolutionary Unit •Human influences on organisms (disturbed vs. undisturbed) Species of Conservation and Management Interest •Regulating mechanisms (competition, predation, habitat availability & quality) Distribution Patterns Scale & Pattern Types Scale Small: Area (e.g., <100 m) over which the environment does not change. Large: Area (e.g., continent) over which the environment does change. Pattern “Home range” “Sp. Dist.” 1) Random: All locations have equal probability (no biological interaction). 2) Regular: All locations have equal probability (‘−’ biological interaction or 3) Clumped: Locations have unequal probability uniform resources). (‘+’ biological interaction or clumped resources). Important for Sampling Design! Equal P[loc] Small Scale (1−3) Large Scale (#3) Antagonistic Interaction Clumped Resources 2 Population Growth Introduction Graphical behavior of population abundance over time. 2 Types: Plankton Blooms 1) Unlimited Population Growth: a) Geometric b) Exponential Rapid increase in population numbers due an abundance of resources typically at low population numbers. Adaptation: 1) Colonization of new habitats. 2) Exploitation of transient resources. 3) Recovery from population crash. 2) Limiting Population Growth: Logistic Growth Rapid increase in population numbers to a stabilizing, limiting abundance (K). Carrying Capacity (K): The maximum number of individuals that can be sustained in an environment at a given time. Abiotic ⇔ Biotic Factors A mathematical reality of all biological populations due to a decrease in available resources necessary for survival and reproduction. Survival + Reproduction Rate of Population Change (by Age Class) Survival Rate, lx: proportion of the population surviving to age x. Fecundity Rate, mx: average number of young produced per individual for age x. Net Reproductive Rate, R0: average number of young produced by an individual in its lifetime. •R0 > 1 ⇒ Population Increasing n •R0 ≅ 1 ⇒ Population Stable R0 = ∑ l x m x •R0 < 1 ⇒ Population Decreasing x =0 ( ) Geometric Rate of Increase, λ: rate of change of the population trajectory. N2 = 25 N1 = 10 o Sl λt = N t +1 pe Generation Time, T : n Nt λ= 25 = 2.5 10 T= ∑ ( xl x m x ) Average Time from Offspring to Offspring x =0 R0 Realized Rate of Increase, r : per capita rate of increase r= ln( R 0) •r > 0 ⇒ Population Increasing •r ≅ 0 ⇒ Population Stable •r < 0 ⇒ Population Decreasing T Phlox (births−deaths) Population Growth Geometric Growth Many Wildlife Non-overlapping Generations Conditions: Recall: Unlimited population growth by species that produce a single generation per year (i.e., 1 reproductive event thus population grows in DISCRETE annual pulses). λ = N t +1 ⇔ N t + 1 = N t λ Nt N m = N 1 λm The abundance at t + 1 can be calculated using the previous abundance & geometric rate of increase. Exponential Growth Overlapping Generations Conditions: Unlimited population growth by species that reproduce continuously (i.e., no discrete annual reproductive event thus population grows CONTINUOUSLY). Continuous population growth can be represented as a rate of change (i.e., derivative) f ′(N ) = dN = rN dt where, r = realized rate of increase (avg. per capita increase) r = births − deaths (constant!) Exp Growth Cannot Notice: as N increases, dN/dt increases be Sustained 3 Population Growth Logistic Growth Overlapping Generations K All populations are governed by a limiting distribution & unable to grow without bound indefinitely. Limiting Growth b=d Sigmoidal Curve +,∩ K/2 ⇔ b−d = 0 ⇒r=0 ⇒ Nt+1 = Nt ⇒ dN / dt = 0 ⎡ ⎛ N t⎞⎤ N t +1 = N t ⎢1 + r m ⎜ 1 − ⎟⎥ ⎝ K ⎠⎦ ⎣ +,∪ discrete Yeast •Food Availability •Space (competition) •Disease •Parasitism •Predation NOTE: K can fluctuate! Balanus African Buffalo Connell Gause Waste Sinclair Space Grass Limits to Population Growth What affects birth and death rates? Andrewartha & Birch Smith & Nicholson Environment vs. Organisms 1) Density-independent Factors: Food abundance Abiotic factors (e.g., precipitation, temperature) that can affect population growth. Seeds/Caterpillars 2) Density-dependent Factors: Biotic factors (e.g., competition, predation, disease) that can affect population growth. mx Galápagos Finches: El Niño Fledglings Rainfall vs. Nt 4X b>d 85% (starvation) b<d Community Ecology Introduction Study of the how abiotic and biotic factors affect the association and interaction of species in a given area. Community: An association of interacting species inhabiting some defined area. e.g., plant community, amphibian community, insect community Community Structure: The assemblage of a community, which includes: •Number of species (species richness) •Relative abundance per species (Nti, species abundance) •Types of species (composition) Guild: Plant Life Forms Groups of species exploiting a common resource in a similar fashion (i.e., have similar ecological roles). e.g., aquatic insect guilds Complex assembly of component guilds. Why Study Guilds? Shredders (CPOM) Collectors (FPOM) Grazers (algae) Predators 1) Focus on functional role 3) Manageable Unit 2) Focus on interaction 4) Basic Building Blocks 4 Species Diversity A combination of number of species and their relative abundance. Species Evenness 1) Species Richness Components: 2) Species Abundance A measure (comparison) of abundance among species. As abundance per species becomes “more even”, species diversity increases. B A Both have 5 species One species comprises 84% and others 4% of community. Each species comprises 20% of community. Common Misuse of Terms Species Diversity is Greater for Community B MAX ( H ′ ) = ln( SR ) MIN ( H ′ ) = 0 Species Diversity A common method to calculate species diversity: pi = proportion of the ith species Community A Species 1 2 3 4 5 H A′ = −( −0.662) Shannon-Wiener Index Abundance 21 1 1 1 1 = 0.662 s H B′ = −( −1.610 ) i =1 = 1.610 H ′ = − ∑ p i ln p i pi 21/25 = 0.84 0.04 0.04 0.04 0.04 ln(pi) ln(0.84)= −0.174 −3.219 −3.219 −3.219 −3.219 piln(pi) 0.84(−0.174)=−0.146 −0.129 −0.129 −0.129 −0.129 pi 5/25 = 0.20 0.20 0.20 0.20 0.20 ln(pi) ln(0.20)= −1.609 −1.069 −1.609 −1.609 −1.609 piln(pi) 0.20(−1.609)=−0.322 −0.322 −0.322 −0.322 −0.322 −1.610 25 Community B Species 1 2 3 4 5 −0.662 Abundance 5 5 5 5 5 25 Environmental Complexity In general, environmental complexity and species diversity are positively related. FHD and Bird Species Diversity FHD = foliage height diversity, where S = # of foliage layers & pi = biomass or volume per layer Robert MacArthur (1958): Examined niche partitioning of warblers in Maine. Found that number of pairs and warbler diversity increased with amount of vegetation and increasing FHD (increase in K and # of niches [unique habitats]). Volume of Foliage Foliage Height Diversity 5 Disturbance & Diversity Environmental Complexity Explains Part Management Implications! Intermediate Disturbance Hypothesis Equilibrium: A state when environmental conditions are more or less stable. Disturbance: 1) A state when environmental conditions are unstable. “Ecology of Natural Disturbance” “Species Dependent” 2) Any discrete event in time that disrupts an ecosystem, community, or population structure and changes resources, substrate availability, or the physical environment (White & Pickett, 1985). 26 Major Sources (fire, storms, disease, predation, human-induced) Hypothesis: Intermediate levels of disturbance (frequency & intensity) will result in the greatest species diversity. Too Much: Few species that complete life cycle between disturbance events. Too Little: Limited to most effective competitors (other excluded). Joseph Connell Just Right: Sufficient time for a variety of species to colonize & complete life yet short enough to prevent exclusion. High Science 199: 1302−1310 Low 6