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Chapter 1, 53: Lecture 1 Ecology: study of processes that determine the geographic distribution of organisms, their abundances, and why they look and act the way they do. Ecosystems and Biomes: 1. ECOSYSTEM: the biotic [living] and abiotic [non-living] factors interacting in a given area. 2. Basic Levels of Organization: individual, family, lines, and species. a. SPECIES: one or more groups of individuals derived from a common ancestor that are capable (actual or potential) of interbreeding and producing fertile offspring. i. Species are reproductively isolated from other species. ii. Each species has a geographic range and within that range, occupies specific habitats. A SPECIES CAN BE COMPOSED OF MANY POPULATIONS, BUT ONLY ONE SPECIES IN A POPULATION. 1. Population: a group of individuals of the same species potentially capable of breeding with one another. a. Has dimensions in time and space: i. Individuals can move between populations (immigration and emigration) but must not get hung up on the exceptions or tails of the distribution. b. INTERACTIONS Species A Species B Competition + Predation, Parasitism + Mutualism;Symbiosis + + Commensalism 0 Amensalism (rare) 0 2. Community: an assemblage of potentially interacting populations or organisms forming an identifiable group within an area. COMMUNITIES ARE INTERACTING SPECIES THAT ARE ADAPTED TO THE SAME ENVIRONMENTAL CONDITIONS SO THEY ALSO ARE RESTRICTED TO A HABITAT AND GEOGRAPHICAL RANGE. Adaptation to the environment a. Temperature and Precipitation b. Energy Flow: movement of suns energy or carbon c. Biogeochemical cycles: movement of nutrients within and among the various biotic and abiotic components of the system. d. Ecological niche: a species` utilization of biotic and abiotic resources in the environment. ORGANISMS ADAPT TO THE FEATURES OF THEIR ENVIROMENT THAT ARE PREDICTABLE Hypothesis What determines the nature of communities, interacting populations? 1. Proximity: are all of the communities found in a particular geographical area more similar to each other than they are to communities in another geographical area? a. FALSE: Ecological Communities are different 2. Arbitrary mix of phenotypes Are communities a random mixture of organisms found in a particular geographical area? a. False 3. Similar climate regions tend to have communities with similar characteristics. The phenotypes species inhibiting similar environmental conditions tend to be similar; there is morphological convergence. a. PHENOTYPES: the morphological, physiological, and behavioral characteristics of individuals within a particular species. Phenotypes results from an interaction between an individuals genotype and the environment. b. EXAMPLE CONVERGENCE: plants near Riverside resemble those near Perth, Australia-Athens, Greece-Cape Town, South Africa. Native to South Africa.

3. Biomes: Characteristics large scale communities occurring in a given climate region

a. The recognizable types of communities or ecosystem(s) that are typical of a geographical region. i. Biomes are characterized by ecologically-equivalent species 1. EXAMPLE: species in tropical rain forests in S.A differ from S.E Asia; but the species in both forests show similar adaptations to the environment: morphology of leaves, growth form (buttressed roots) 2. Boundaries are often not sharp, but can be dramatic ECOTONE 3. Several communities can occur within a biome


1. Tropical forests a. Dry tropical forest b. Tropical rain forest, wet c. Tropical deciduous forest: distinct wet, dry i. Occur within 23.5 latitude of the equator ii. Avg. temp and daylength vary little; rainfall can vary substantially iii. Competition for light is intense, poor nutrient soils iv. Diverse communities, species 2. Savanna: grassland with scattered trees a. Tropical and Subtropical b. 3 Seasons: cool and dry, hot and dry, warm and wet c. Porous soils with a tin later of hummus d. Large predators and herbivores e. Periodic fires are important 3. Desert: driest of all terrestrial biomes: low and unpredictable rainfall (<30cm/yr) a. Hot deserts (soil surface temp (>60 C) i.e., Sahara, Sonoran) v.s cold deserts (west of the Rockies, eastern Argentina (Atacama), central Asia (Gobi) b. Drought-resistance plants/ succulents w/h store water: cacti c. Animals adapted to scarcity of water: physiologically and behaviorally 4. Chaparral: 30-40 latitiude; influence by cool ocean currents a. Mild rainy winters and hot, dry summers b. Dominated by dense, spiny shrubs w/ tough evergreen leaves c. Periodic fire are important: food reserves are often stored in a fire-resistant root crown; seeds may germinate after hot fires d. Seed-eating animals and their predators e. Riverside 5. Temperate grassland: relatively cold winters a. What keeps a grassland devoid of trees: seasonal drought, occasional fires, grazing b. Soils tend to be thick and nutrient rich 6. Temperate forest: species rich a. Occur in the mid-latitudes: cold winters, hot summers => 5-6 month growing season b. Deciduous trees => litter enhances soil c. Unlike tropical forest where the understory is very shaded, temperate forests support several strata of canopy plants, an understory of shrubs, forbs 7. Taiga: coniferous or boreal forests a. cool high elevations and high latitudes b. long cold winters; short, wet summers c. thin, nutrient-poor, often acidic, soils d. trees adapted to snowfall and cold winters e. species-poor: dense stands of only a few species 8. Tundra: northernmost limits of plants growth a. low, shrubby vegetation i. arctic tundra: permafrost => permanently frozen subsoil 1. low evapotranspiration => saturated soils 2. brief summers with nearly continuous light ii. light tundra: at all latitudes with very high elevation 1. above treeline

LECTURE 3, 4 CLIMATE PATTERNS 1. Temperature 2. Precipitation 3. Climate and biomes TRENDS IN CLIMATE ASSOCIATED WITH LATITUDE HAVE A LARGER ROLE IN DETERMINING THE BIOMES FOUND AT PARTICULAR PLACE ON EARTH CLIMATE: the average state of weather at a particular location on the planet a. Determination of climate 1. Gradients in solar energy input 2. Earth tilt and spin (seasons) 3. Spatial arrangement of water and land a. Light b. Water c. Wind d. Temperature i. GMT Global Mean Temperature and Greenhouse effect 1. GMT = Average Earth surface temperature averaging day, night, season, and latitude ii. Heat Input: from Earth core, radioactive decay, and sun. 1. Sun most important; average solar input at top of atmosphere iii. Heat loss: from black body radiation any body above absolute zero (0K or -273 C) radiates electromagnetic radiation at a wavelength proportion to its temperature. 1. Earth re-radiates the energy obtained from sun at a longer wavelength (lower E) than receiving wavelength because some of the arriving energy is absorbed by the earth (the missing E), re-radiates the lower energy in the infrared range. As earth warms further, the wavelength of re-radiated energy shorten and rate of energy loss increases (shorter wavelength with more energy) until an equilibrium is reached (ENERGY IN= ENERGY OUT) a. Example: days are warmer than nights because input is higher during the day. i. Cold clear nights vs warmer cloudy nights. 1. Clouds absorb and re-radiate some of the head reradiated from the earth! 2. Humidity can keep temperature warmer. (Water vapor in the air absorbing E) a. Water molecule effect on temperature: little humidity (water) to maintain the heat at night and prevent it from being re-radiated so it cools off reasonably quick. iv. Atmosphere effect (Greenhouse effect) : 1. Insulating blanket due to a. Greenhouse gasses: CO2 CH4 H2O: absorb the re-radiated energy from earth and radiates it back to the surface 2. Greenhouse Effect: atmosphere moderates the global temperature (~16 C) a. w/o the atmosphere, the average temperature would be ~0 F/-20 C b. the glass in the greenhouse allows light energy to pass through, but reflects re-radiated energy in longer wavelength trying to exit back to the interior warming the space. c. POSITIVE EFFECT: makes complexity and abundance of life on earth possible. d. Too much isnt a good thing => global warming. i. Global Warming: increase in sea level, extinction, increase in diseases in the tropics, marked and unpredictable variations in temperature and precipitation.

1. Temperature and EFFECTS a. Planetary and Solar Effects on Temperature i. Long-term Rhythm Factors in the orientation of the earths axis within the earths orbit about the sun that have influences of long-term periodic changes in GMT (global mean temperature) include 1. Tilt a. Obliquity of the axis b. Tilt changes oscillates between 21.5 and 24.5 (Average 23.5 degrees) c. The more tilt the earths axis, the greater the difference in distance from sun at different time of year, greater differences of summer & winter radiation of the same hemisphere, and greater the differences between simultaneous temperatures in the northern & southern hemisphere. 2. Eccentricity a. Orbit changes from elliptical to almost circular changes distance from the sun 3. Precession a. Circular arc of the pole (= to the wobble of a spinning top) moves earth closer/farther away from the sun b. Latitude and Temperature i. Local temperatures depends on latitude ii. @ EQUATOR suns rays strike the Earth from straight overhead and perpendicular to the surface 2 1. Little of atmosphere to travel through and more energy per m arriving at surface and the higher the thermal equilibrium (higher average temperature) iii. @ Poles : the greater the angle to the light striking the surface, the greater the surface area over which 2 the light is spread, the lower the energy per m arriving at surface and the lower the thermal equilibrium (lower average temperature) c. Elevation and Temperature i. Local temperature depends on elevation ii. @ high elevation, fixed number of molecules occupy a specific volume (cooler than lower) iii. @lower elevation, the molecules move downs and compresses into a smaller volume (due to increase in air pressure) iv. TEMPERATURE OF THE AIR INCREASES EVEN IN ABSENCE OF ADDITIONAL APPLICATION OF HEAT FROM OUTSIDE (ADIABATIC HEATING) because molecules bounce of each other more frequently in smaller space

d. Altitude and Temperature i. Moving up in altitude, moving away from equator e. Seasons i. Change due to tilt of the Earth relative to the sun 1. Tilt means a. fewer hours of sunlight in northern hemisphere in winter, fewer hours to absorb energy (December 21) b. longer hours of sunlight in northern hemisphere in summer, longest day June 21 (closest to the sun) ii. region between 23.5 N and 23.5 S of the equator receives sunlight from directly overhead 1. greatest annual input 2. least seasonal variation f. Proximity to Water and Temperature i. Local Temperature is influenced by the proximity to water. 1. Moderation of T a. Water with higher heat capacity holds more KE per unit volume than land so water heats and cools more slowly b. Water cools adjacent to land in summer and warms it in winter c. Produces MARITIME climates

2. PRECIPITATION PATTERNS a. Air Temperature Effect: i. Warm airs holds more water vapor than cold air. 1. If warm saturated air is cooled the water turns to liquid and falls as precipitation. a. Example, warm air forming clouds and thunderstorms 2. If cool dry air is warmed if can absorb/hold more water vapor. a. Sucks moisture from surroundings b. Global Air Circulation/ Creation of Convection Cells i. Warm air rises. ii. More intense solar input @EQUATOR warms equatorial air causing it to rise. Rising air expands and cools producing 1. PRECIPITATION & CIRCULATION a. Rising air at equator creates a LOW PRESSURE ZONE (tropical convergence)

i. Replaced by air sucked in from north and south

ii. Rising air now cool and falls at 30 N and 30 S latitude

iii. As it falls it compresses, heats, and creates a region of high pressure b. 90 N and 90 S weak convection cells create heavy cold polar air that sinks at poles creating region of high pressure i. It warms as it moves towards the equators ii. Creates subpolar lows at 60 N and 60 S latitude 1. Jet streams that blow W to E at 5 to 8 miles high and up to 200 mph 30 N and 60 N and 30 S and 60 S latitude as air flows from areas of high pressure to ares of low pressure 2. EAST WEST CIRCULATION (not jet streams) a. Physics of spinning Earth and air movement towards or away from the equator prevailing wind patterns. These patterns (CLOCKWISE ROTATION @ N HEMISPHERE, COUNTERCLOCKWISE @ S HEMISPHERE) create areas near the i. Equator where winds are calm. ii. Above and below the equator, the winds blow from E to W iii. Areas between 30 to 60 N and S the winds blows from W to E c. Ocean Currents and Influence on Climate i. Surface water pushed by prevailing winds and deflected by continents. ii. Clockwise circulation pattern in North (GYRE) and counterclockwise in S iii. Warm water from equator is circulated towards the poles along the eastern continental margins and cold water is circulated alone western continental margins c.

3. BIOMES AND CLIMATES a. Why are the deserts around 30 degrees N & S latitude? i. As warm moist tropical air rises and cools, it loses water as rain along the equator. ii. As it cools and descends around 30 degrees N & S latitude from the tropical convection cell, it is dry. b. Why do Mediterranean climates (cool wet winters and hot dry summers) occur on the western edges of continents at mid latitudes? i. Prevailing westerly winds bring moisture from the ocean. 1. In winter, the winds are warmer than land and drop moisture as they cool. 2. In summer, the winds are cooler than land and retain moisture as land warms the air (dry humidity) c. Why is it so dry to the East of mountains on the western border of continents at mid-latitudes? i. Prevailing west winds bring moisture from ocean. 1. When winds hit mountains, the air rises and cools, dropping moisture on west slopes and leaving cool dry air to flow over east slopes. Phenomenon is a rain shadow. d. Why thunderstorms in the middle of the continent? i. Local convection cells formed as land heats, air above it warms and rises, creating an area of local low pressure pulling in warm, moist surface air from Gulf of Mexico. ii. As warm mist air rises, it cools and water precipitates out. iii. Same physics (without Gulf air) occur along mountains and formation of summer thunderstorms.


Lecture 5 ENERGY FLOW: 1. The biosphere is not a closed system. a. Its function is dependent on the continuous supply of energy from outside the system to maintain the complex structure. i. EXAMPLES 1. Algae Fish Human 2. Grass Cow Human 3. Forest Deer Human 2. PLANTS PROVIDE ENERGY a. Plants essential to make elements available to animals and decomposers b. Plants also essential in providing the most important need of all life energy. c. Energy cannot be recycled efficiently d. Continuous influx of energy is required; the source is typically the sun TROPHIC LEVELS: 1. Food Chain: a pathway along which food is transferred from one trophic level to the next trophic level. a. Plants and Phytoplankton are the Primary producers (AUTOTROPHS) i. Plant Herbivore Carnivore Carnivore ii. 1 producers 1 consumer 2 consumers 3 consumers iii. 1, 2 and 3 consumers (HETEROTROPHS) b. Autotrophs: Self-feeders use energy to convert carbon dioxide into biomass. i. EXAMPLE 1. Photosynthetic plants or chemoautotrophic bacteria (Primary producers) c. Heterotrophs: other feeders depend directly or indirectly on autotrophs. i. Primary Consumers: Herbivores that eat primary producers ii. Secondary Consumers: Carnivores that eat primary consumers iii. Tertiary Consumers: Top carnivores that eat carnivores d. Decomposers: feed on nonliving organic matter. i. EXAMPLE 1. Bacteria, fungi and some animals TROPIC RELATIONSHIPS ARE MORE COMPLEX THAN SIMPLE LINEAR CHAINS 2. Food webs: branched pathways that are used to represent how food is transferred among organisms in a community or an ecosystem. TROPHIC LEVELS TEND TO BREAK DOWN IN NATURE 3. Omnivores: feed on more than one trophic level a. EXAMPLE i. Coyote, bear, or human TROPHIC LEVELS MAY CHANGE WITH AGE INEFFICIENCY OF ENERGY TRANSFER: 1. Energy is lost as biomass is transferred up the food chain or through a food web. 2. Of the solar energy striking the atmosphere, approx. 53% reflected back into space by clouds, dust, molecules in atmosphere. Approx 47% hits Earth surface. (100% Energy hitting plant 80% reflected) x 5% fixed as biomass = 1% of initial energy hitting plant fixed as biomass = GPP 3. Gross Primary Production (GPP): energy fixed by photosynthesis a. Not all of the energy fixed by photosynthesis is available to herbivores. Some of the energy is lost to plant respiration and metabolic processes. 4. Net Primary Production (NPP): GPP R; available energy = energy fixed by photosynthesis energy lost via respiration (ultimately heat). Represents the surplus available to the herbivore trophic level. a. NPP ranges between 50-90% GPP: high for algae and grasses and lower fro tress (support a large amnt. of nonphotosynthetic biomass) b. NPP for earth is very large and expressed as biomass per unit time 5. Biomass: the weight or standing crop of species 6. Production: change in biomass per unit time, or the change in energy per unit area per unit time.

7. Food Chain Efficiency (Ecological Efficiency): percentage of energy transferred from one trophic level to the next trophic level

F.C.E on average, its around 10% (or, 90% of E entering is lost TROPHIC PYRAMID 1. Energy decreases as one moves through higher trophic levels, so do numbers and biomass in most system. a. bottom-heavy pyramid 2. In food chains where the lower trophic level organisms are shorter-lived than their predators and the food organisms may turn over at a rapid rate. a. inverted biomass pyramid 3. Ecological efficiencies are usually lower in terrestrial environments than in aquatic environment. 4. Also lower in tropical systems, but high E input results in high productivity. 5. Comparisons b/w temperate and tropical systems a. Photoinhibition of photosynthesis in the tropics (aquatics); higher relative metabolic costs; nutrient limitation b. Plants metabolic costs are < animals (30-85% less) 6. Aquatic ecosystem can support more trophic levels than the average terrestrial system a. Cow: temperate grassland (highest NPP) b. Fish: temperate algae (high NPP), most efficient ectotherm transfer c. Deer: temperate forest (low NPP), least efficient endotherm transfer 7. Ectotherm: Any so-called cold-blooded animal; that is, any animal whose regulation of body temperature depends on external sources, such as sunlight or a heated rock surface. a. EXAMPLE i. Fishes, amphibians, reptiles and invertebrates 8. Endotherm: Any so-called cold-blooded animal; that is, any animal whose regulation of body temperature depends on external sources, such as sunlight or a heated rock surface. a. EXAMPLE i. Birds and mammals BIOACCUMULATION AND BIOMAGNIFICATION 1. Organisms accumulate materials from their environment and from the things that they eat 2. Because of the inefficiencies in energy exchange, more biomass must be consumed from a lower trophic level to maintain a biomass at the next trophic level. 3. Chemicals that are not metabolized may be more and more concentrated in successive trophic levels a. EXAMPLE i. DDT (Dithiothreitol)

Lecture 6 1. Biogeochemical cycles a. Besides input of solar energy, Earth is a CLOSED SYSTEM b. Nutrient, element cycles i. Biotic and abiotic components c. Structure of ecosystems and (Net primary production) NPP are strongly influenced by the rate of recycling of nutrients as a given place. d. BASIC FEATURES OF BIGEOCHEMICAL CYCLES i. Approx.. balanced on a global scale ii. Driven by energy iii. Have biological accessible/inaccessible reservoirs 1. Recycling is critical because in some forms a. Nutrients may be an accessible form whereas in other they are inaccessible to living things i. Nutrients are converted from inaccessible to accessible forms by biological, geological, and biogeochemical cycles (chemical processes). e. Cycles i. Carbon Cycle

1. Left: depict the storage of carbon in the ocean as dissolved CO2 and as calcium carbonate in the form of shells, rocks, etc. 2. Right: Exchanges in cycle by photosynthesis, respiration, consumption, erosion, death, weathering, combustion (fossil fuels), sedimentation, and fire (natural). ii. Important points to remember about the carbon cycle: 1. a readily accessible reservoir exists in the atmosphere. Cycling time of an atom of carbon in the atmosphere is approximately 3-5 years. 2. The reciprocal processes of photosynthesis and respiration are important for exchange of carbon dioxide between the biota and the atmosphere. 3. The oceans are a buffer for the atmosphere. Even though the cycling time of a carbon atom dissolved in seawater is ~300-400 years, the amount of carbon dissolved in seawater is about 50 times the pool in the atmosphere. 4. The amount of carbon at the bottom of ocean as calcium carbonate is 5 orders of magnitude greater than the amount of carbon in the atmosphere. This carbon recycles extremely slowly (available by weathering or burning); for all practical purposes, the carbon in calcium carbonate is an inaccessible reservoir. This also was true for fossil fuels...until man came along. 5. Human activity accounts for only 2% of the carbon cycled each year

2. Nitrogen Cycle th a. 4 most common element b. Component of proteins, dna, chlorophyll, and other organic molecules c. large pool of N in the atmosphere (78%) cannot be assimilated by most organisms: i. Two major sources are nitrogen 1. fixing organisms 2. lightning (14-28X < N-fixers per annum) d. nitrogen is not directly involved in the release of chemical energy by respiration: role is linked to protein molecules and nucleic acids e. breakdown of nitrogenous organic compounds to inorganic forms requires many steps often involving specialized bacteria. f. most of the biochemical transformations involved in the decompostion/transformation of nitrogen take place in the soil and oceans. i. Problem: inorganic nitrogen is very soluble and forms of nitrogen (NH 3 and NO3) are toxic in high concentrations. 1. Leaching: positively charged ions are replaced by H+ ions; negatively charged ions do not bind to clay particles such as calcium, magnesium, potassium.

g. Fixation converts nitrogen gas N2 to ammonia and ammonium i. Processes: Rainout deposition, fixation, nitrification, absorption, consumption, erosion, excretion, sedimentation, denitrification

3. Phosphorus Cycle

a. P used in nucleic acids, phospholipids, many proteins (~1/10 the N requirement: still a high level), bones & teeth b. Processes: Consumption, death, absorption, human activity, weathering, erosion, chemical precipitation, sedimentation


Lecture7, 8
ECOLOGICAL COMMUNITIES Def: A group of interacting species that live together in the same area. A community may be all interacting species Communities have characteristics that are above and beyond those of the species it contains. Ecological communities do not all contain the same number of species; yet, some very distinct trends are evident when we compare the number of species found in communities of different biogeographic regions of the Earth. a. EXAMPLE: animals and plants were more abundant and varied in the tropics than in other parts of the world. 5. Small or remote islands have fewer species than large islands and islands closer to the mainland. WHAT DOES SPECIES DIVERSITY MEAN AND HOW IS IT MEASURED? 1. Species Richness: is a number of different species represented in an ecological community, landscape or region. 2. Evenness: a measure of the relative abundance of species in communities. a. When species are more equally (evenly) abundant, we say that the species exhibit a more even distribution. b. We can compare the evenness of the two communities by plotting the relative abundance of the species in each community: ranked species abundance curves. 3. Both richness and evenness should be included in a measure of species diversity. 4. A mathematical index that includes both concepts is the Shannon Index of diversity. WHAT DETERMINE SPECIES DIVERSITY? (3) major factors 1. Abiotic environment 2. Degree of disturbance 3. Habitat patch size and isolation SPICIES DIVERSITY DECREASE W/ ENVIRONMENT HARSHNESS 1. Increasing latitude and disturbance 2. More species occur in tropical communities 3. Higher latitudes tend to have more environment extremes (increasing latitude cold, dry, lower NPP) WHY ARE MORE SPECIES IN TROPICS? 1. Hypotheses: a. Time hypothesis: tropical environment is older, allowing more time for the evolution of a greater variety of plants and animals. (NO GLACIATION) b. Speciation rates are higher in the tropics i. migration (no reason to move to escape harsh conditions) ii. generation (favorable temps, no hibernation, faster development) iii. resource specialization iv. Many tropical populations are relatively sedentary because of the environment is relatively aseasonal environment is favorable; no need to move. c. Extinction rates are lower in the tropics d. Competition might be lower in the tropics because of i. More resources ii. spatial heterogeneity (more niches or spaces to live) iii. Predators cropping prey populations reduces competition: importance of predation. iv. Environmental stability of tropics might allow small populations to persist where they would not in the environmental extreme found in more temperate climates. SPECIES DIVERSTIY IS THOUGHT TO RESULT FROM A BALANCE BETWEEN REGIONAL PROCESS AND MORE LOCAL PROCESS 1. Regional processes: immigration and dispersal 2. Local processes: influence extinction (predation, competition, adaptation and chance events) or contribute to environmental differences within a particular region (productivity, habitat heterogeneity) 1. 2. 3. 4.


1. Pollution effects: Human activities and consequent pollution presents organisms with novel, stressful environments that phenotypes not adapted to. a. EXAMPLE: leaching of phosphates from fertilizers and detergents and Bioaccumulation and Biomagnification

2. Fire effects: increased fire frequency, fire suppression, fuel loading, patterns of vegetation structure and introduced species, catastrophic while fires. 3. Introduced species and changes in ecosystem: homogenized global flora and fauna a. Competition for space, light, water nest sties, and food reduction in species diversity b. Predation reduction in population size with resulting impacts on reproduction c. Reproductive success rats, mongoose, cane toad, cuckoos, brown snake, foxes/cats d. Disease organisms human (west Nile, Asian bird flu, dengue), animal (exotic Newcastle, hoof and mouth, mad cow/BSE), and plant (chestnut blight, Dutch elm disease, sudden oak death) HABIT SIZE AND ISOLATION AND DISTURBACNE

1. Habitat size and isolations: a. Observations: i. Islands contains fewer species than does the nearby mainland ii. Small islands contain fewer species than do large island iii. Islands closer to the mainland contain more species than do similar sized islands located at a greater distance from the mainland. 2. Equilibrium Theory of Island Biogeography a. Hypothesized that these trends in species diversity are caused by a balance (equilibrium) between immigration and extinction rates b. Immigration rate of new species: decrease with number of species present. As more and more colonists arrive from mainland, fewer of them represent a new species. c. Extinction rate of species: increase with the number of species present (more sp. Present, the more likely that any species will go extinct). d. The equilibrium species number is the point where immigration rate balances extinction rate.

e. Proximity and Island Size: i. if immigration is reduced, equilibrium species number will decrease. (i.e., an island far away from the mainland relative to an island nearer the mainland) ii. if extinction rate is increased, equilibrium species number will decrease (i.e., small island or small populations) iii. Large islands have more species than small islands. f. Relevance: conservation biology where we create habitat islands called reserves. i. Natural habitats are being destroyed and fragmented by human land uses. This creates small, isolated habitat fragments in which S# is reduced (=small isles). g. Ripple effect: loss of coyotes increased domestic cats/feral cats which increased mortality on birds, small mammals, especially at he edge of suburban environment. h. Edge effects: recreational hunting by cats illustrates edge effect. i. SMALL RESERVES will be more severely affected by edge effects than will large reserves ii. Location is important because small reserve can serve as islands along a dispersal corridor between larger reserves. iii. Reserves should be as large as possible, as close together as possible, and connected by corridors. 3. Disturbance a. Diversity can depend on disturbance and diversity is highest at an intermediate level of disturbance. b. Succession: a repeatable progressive change through time in the species composition of a community. c. 1 succession: establishment and development of communities in newly formed habitats the habitat is altered by succession and made suitable for succeeding successional stage.

d. Xerarch succession: start off with a habitat characterized by low water retention (xeric = dry), rock, at the early stages of succession (think of volcanic island) e. Hydrarch succession: which the soil is saturated and succession begins in the open water of a shallow lake, bog, etc and is initiated by facter(s) that reduce water depth (plant species w/ h grows out across the water surface) and eventually increase soil aeration i. Bare rock => lichens => moss => grasses + forbs => woody plants (pines => hardwoods: CLIMAX COMMUNITY) ii. Soil: none => traces of organic matter in cracks => thin soil => => soil gets progressively deeper and richer f. Climax community: a self-perpetuation stable community in a particular environment g. 2 succession: regrowth after the climax community is destroyed by a disturbance i. EXAMPLE: tree fall in the forest or a fire ii. 2 succession process is more rapid than in 1. 1. EXAMPLE: fire disturbs a chaparral community. Some annuals only appear after fires seed bank. h. Earlier colonizers: good dispersal mechanisms, rapid germination and growth, poor competitors often thought of as weed species i. Late colonizers: moderate dispersal mechanisms; often require facilitation of the habitat by the early colonizers, good competitors. j. With frequent disturbance only early successional species; Early successional communities less diverse k. No disturbance climate species l. Intermediate disturbance mixture of early successional and climax species: Patchwork that contains species that are characteristic of many successional stage m. Fire disturbance maintenance of a subclimax community by disturbance n. Keystone species: a species that has a disproportionately large effect on its environment relative to its abundance

Lecture 9,10 1. Population reproductive group of individuals of the same species living in the same area a. small subset of species b. evolutionary significance 2. Population ecology: measuring changes in population size and composition and identifying factors causing the changes observed. 3. Deme: a local group of interbreeding organisms.

a. The population will function in a narrower range of environmental conditions than will the individuals in a population. i. A population can live and reproduce successfully (a<->a) ii. maintain organism: limits define where an organism can exist indefinitely (b <-> b) iii. critical functions: upper (c) and lower lethal limits (c)

Example: stream insects: adult body size, metabolic efficiency, fecundity, and abundance are maximized at the center of the latitudinal range. Can be asymmetrical: stenothermic vs. eurythermic b. density: number of individuals per unit area or unit volume. 4. Some of the basic questions asked by population biologists include: a. How many organisms are there? i. absolute estimation of population size. ii. Total counts: most direct method to find out how many organisms live in a particular area is to count them. iii. Subsampling methods: there are two general methods...you will use both of them in the lab. 1. quadrats: sampling areas of known size. 2. capture-summation methods: widely used in small mammal studies; although this method is based on some questionable assumptions: Limit discussion/presentation

a. population is stationary b. probability of capture is the same for each animal c. probability of capture remains constant from trapping to trapping d. PROCEDURE i. Set the same number of traps each time the population is sampled. Expect to catch a smaller number every time because the population is depleted as sampling occurs. To estimate population size: plot daily catch against number of animals already caught. 3. capture-recapture: a particularly important method in animal ecology is mark-recapture 4. All capture-recapture models make three critical assumptions: a. marked and unmarked individuals are captured randomly b. marked individuals are subject to the same mortality rate as unmarked individuals no increase in probability of death, not a target for predator c. marks are not lost or overlooked 5. Calculation a. N = total population size (what you are trying to find) b. Time 1, M and release c. Time 2, capture individuals C (total number of individuals in a sample) and R (number of marked individual in C) d. DETERMINATION OF POPULATION SIZE ON THE ASSUMPTION

b. How does population size vary over space? i. DISPERSION: the pattern of spacing of individuals within the geographical boundaries of the population 1. Clumped: aggregated in patches (e.g., insects on a stream bed, fish in schools, insects under logs) often associated with a specific microclimate, food source, or behavioral benefit (herd or school). Most common distribution. 2. Uniform: spacing is even (e.g., penguins at a rookery; trees in orchard [artificial]) often associated with direct interactions between individuals (antagonistic interactions - chemical [allelopathic], physical [shade], or behavioral [aggression]- between individuals competing for resources). 3. Random: spacing varies in unpredictably forest trees: absence of strong attractions, heterogeneity of resources, or repulsion: RARER than others c. How does population size vary over time? / How fast does a population grow? i. Factors which add individuals: 1. births and immigration ii. Factors which subtract individuals: 1. deaths and emigration iii. depends on the composition of the population 1. . One with a high proportion of females in a reproductive age range will grow more rapidly than one composed mostly of post-reproductive and pre-reproductive individuals. iv. PARAMETERS THAT ARE IMPORTANT 1. Age at first reproduction 2. FECUNDITY : a. number of reproductive episodes/lifetime (parity: semelparous = breeding once in a lifetime vs. iteroparous= breeding several times in a lifetime b. average number of offspring per episode (clutch size) c. AGE-SPECIFIC FECUNDITY (bx) 3. SURVIVORSHIP (lx): a. how many females of a particular age, x, are reproducing v. LIFE TABLE 1. Horizontal (cohort) life table: large numbers of newborn individuals are "marked" and followed until they die. One has to be able to estimate deaths from losses due to emigration. Age-specific fecundity data can be collected from marked individuals. 2. Vertical (static) life table: if individuals can be aged directly, as by tree rings, fish scales or otoliths, mammalian teeth, obituary columns, inscriptions on head stones, etc.; then a sample of the population is collected and the age distribution is determined. "Snapshot" in time of the population. Age-specific fecundity has to be estimated indirectly in most cases.

3. LIFETABLE CALCULATION a. NET REPRODUCTIVE RATE (net fecundity) i. R0 = the expected number of female offspring produced during the lifetime of an original female in a cohort 1. Multiplication rate per generation, factor by which population size changes each generation ii. R0 = lxbx 1. where lx is the probability at birth of a female surviving to age x [proportion of cohort surviving to beginning of age x] 2. bx is the expected daughters produced at age x. 3. The value lxbx represents the expected reproduction at age x. iii. If.. 1. R0 = 1, then the population is stable = 1 female offspring/ female parent/ generation 2. R0 < 1, then the population is decreasing; 3. R0 > 1, then the population is increasing. b. MEAN GENERATION TIME: i. T = the mean period elapsing between the birth of parents (mother and birth of offspring daughter) ( )( )

vi. Geometric Growth 1. Consider non-overlapping generations a. N1 = N0 - N0d + N0b = N0(1 - d + b) = N0R0 i. b = per capita birth rate, d = per capita death rate, (ignoring immigration and emigration) b. If ecological conditions are unchanged, R0 stays the same, then 2 i. N2 = N1R0 = (N0R0)R0 = N0R0 c. therefore, n i. Nn = N0R0 , where n = the number of generations 2. This represents a geometric series (e.g., 1, 3, 9, 27, ...) where change is multiplied by a constant factor (i.,e., the number in each generation is multiplied by a factor of 3) each time unit, not by adding a constant amount (e.g., 3, 6, 9, 12, 15,...) during each time unit. a. Annual Growth Rate i. NT = N0R0; where NT = population size at the end of one generation 1. N0 = population size at time 0 ii. N1 year = N0 1. = is the annual growth rate which produces geometric growth vii. Exponential Growth 1. GENERATIONS OVERLAP: Continuous birth and deaths of an organism a. instantaneous rates of change. b. N/t = B D i. Where B = # births in a population in a time interval and D = # deaths in the population in the same time interval c. dN/dt = (b - d) N = rN i. r = instantaneous growth rate or intrinsic rate of increase (individuals per individual per unit time = instantaneous birth rates instantaneous death rates) 1. r = 0; population is stable, not changing 2. r > 0; population is increasing 3. r < 0; population is declining ii. As t -> 0, N/t -> dN/dt = (b - d)N = rN


LECTURE 11 1. Demography: study of birth and death processes that determine a. Rate of change in population size b. Age structure of populations c. Vital Statistics: probability of death and fecundity of individuals of each age in the population i. important for understanding natural selection, competition, predation, and even ecosystem stability because populations with different birth and death statistics may have very different growth potentials.

d. Demographers often make a distinction between the instantaneous rate of increase (r) and the innate capacity for increase (rmax or r0). The instantaneous rate of increase is calculated from the difference between instantaneous birth rate and death rate (b-d) usually in natural populations or the change in population size per unit time. i. rmax is the maximum biotic potential for reproduction and growth of a species. 2. LOGISTIC POPULATION GROWTH a. what factors limit population growth? i. Birth and death rates are sensitive to change in climate, food supply, habitat availability, abundance of predators on that population b. Logistic incorporates the intrinsic capacity of a population to grow and limitations imposed by the environment i. ii. K, carrying capacity: resource levels will be depressed until the minimum maintenance requirement for each individual is reached and the population stops growing ( ) 1. a. Rate of increase of population per unit time = b. rate of population growth per capita x population size x unutilized opportunity for population growth 2. When.. a. N=K, population growth = 0 therefore population is stable b. N > K, population is declining c. N < K, population is growing


The logistic model predicts the regulation of population as density increases. i. density-dependent factors: regulating factors relating to population size. 1. As population increases, the effect of the specific factor becomes more severe. ii. density-independent factors: factors unrelated to population density. The strength of the impact of the specific factor does not change with changes in population size.

Lecture 12

1. Negative/Negative interaction between two species represents competition IT IS POSSIBLE TO HAVE POSITIVE/NEUTRAL [COMMENSUALISM] OR POSITVIE/POSITVE [MUTUALISM] INTERACTION AS WELL.

2. Intraspecific competition: the reliance of two or more populations of the same species on the same limited resources.

3. Interspecific competition: the reliance of two or more species on similar limiting resources

4. Model prediction: a. Two competitors (species 1 and 2) can only coexist if individuals of species 1 depress the per-capita growth rate of individuals of species 1 more than they depress the per capita growth rate of species 2, vice versa. b. In other words, intraspecific competition >> interspecific competition; otherwise one species will go extinct. 5. Possible outcomes of strong competitive interactions: a. Outcome 1: Coexistence (e.g., at lower carrying capacities) i. Gause (1932): Saccharomyces vs. Schizosaccharomyces 1. Yeast growth stops before the reserves of food and energy are exhausted 2. Accumulation ethyl alcohol (byproduct of breakdown of sugar under anaerobic conditions) 3. Alcohol kills new yeast buds after they separate from the mother cell ii. Gause demonstrated that a simple physiological basis existed and that coexistence at lower Ks occurred when the two competitors were reared together. b. Outcome 2: Competitive exclusion i. P. Aurelia vs. P. caudatum 1. Each grow well separately at 26C and pH = 8; bacteria added daily as a food source. 2. Tow factors limit growth: food and accumulation of waste products (more detrimental to P. caudatum than to P. Aurelia) 3. In mixed cultures: P. caudatum went extinct ii. The competitive exclusion principle: Complete competitors cannot coexist c. Outcome 3: Competitive Displacement i. Fundamental vs. realized niche 1. Fundamental niche: the resources that a population is theoretically capable of using under ideal circumstances

2. Realized niche: the subset of the fundamental niche actually occupied by a species; resources a population actually uses. Constrained by interaction with other living things and the abiotic environment (i.e., disturbance, harshness, etc) 3. If two species have identical fundamental niches, then they cannot coexist 4. Weaker competitors must find an alternative resource(s) if they are not to be driven to extinction ii. Important study: Chthamalus vs. Balanus 1. Balanus balanoides: occupies a lower and less exposed zone of the rocky intertidal a. Cant survive desiccation in the high intertidal b. Grows faster and crowds our Chthalamus in the lower intertidal c. Lower limit is set by predatory whlks d. Fundamental = Realized for Balanus 2. Chathalamus stellatus: attached to rocks in the high intertidal a. Can survive in the lower intertidal when Balanus is absent. b. Balanus overgrowing, undercutting and crushing Chthalamus c. Fundamental > Realized for Chthalamus

d. Outcome 4: Niche Shift i. Niche Shift: Evolutionary response with character displacement or resource partitioning ii. Evolutionary changes that permit coexistence: 1. Changes in life history traits: r, K, competitive abilities; a. i.e., colonizing (r) vs. competing (K) life historie 2. Ecological divergence 3. Resource partitioning: sympatric species consume slightly different foods or use resources in slightly different ways a. i.e., birds may forage in particular regions of a fire tree b. divide resource and specialize potentially one of the reasons for enhanced species diversity in the tropics is the abundance of a variety of resources and the pressure to specialize from competition. 4. Character displacement: most cases of resource partitioning appear to reflect past selection for divergence in resource utilization a. The differential reproduction and survival caused by interspecific competition has led to the divergence between close competitors i. i.e., beak size in Galapagos finches which are allopatric (isolated) and sympatric (together) e. TYPE OF COMPETITION (INTRASPECIFIC) i. Interference (direct, contact) competition: organisms seeking a resource may harm one another in the process; even if the resource is not in short supply. 1. i.e., birds fighting over territories, barnacles fighting for space on rocks, allelopathy in plants, dung flies fighting for mates (remember uniform spacing) ii. Exploitative (indirect, scramble) competition: organisms utilize common resources that are at times in short supply. 1. How quickly or how efficiently an organism can utilize resources will determine the outcome of competitive interaction 2. i.e., yeast, Daphnia, house flies, blow flies, dung beetle, mosquitoes

Lecture 13

PREDATION (CARNIVORY, HERBIVORY) OR PARASITISM [+/- INTERACTION] 1. Predators kill their prey; whereas, a parasite does not necessarily kill its host. 2. Herbivores often crop plant hosts eat some but not all AN IMPORTANT PROCESS 1. Moving energy and materials through the trophic levels remember energy flow, nutrient cycling, and trophic pyramid lecture materials 2. Influences the distribution and abundance of species predation as a population regulation mechanism 3. Strong selective force: arm races between predator and prey Lotka-Volterra predator-prey equations: Prey: ( )

rH = unrestricted reproductive rate of the prey in the absence of predators E = coefficient representing the efficiency of predation p = # of predators; H = # of prey

Predation rate and the # of prey are related in such a way that if THE NUMBER OF PREY DOUBLES, SO DOES THE PREDATION rate


dp = instantaneous death rate of predators in the absence of prey = skill of pred. for catching and transforming prey into new predators P = # of predators; H = # of prey IMMIGRATION OF PREY FROM PREDATOR-FREE SPACE INTO THE POPULATIO IS OFTEN REQURIED TO MAINTAIN THE CYCLE 1. Example of cycles in nature a. Snowshoe hare-lynx: 10 yr cycles. i. Change in hares food supply appear to be the major driving force ii. In this case, plant quality declines w/ increasing herbivory by the rabbits (density dependent response) iii. Hares on island w/o lynx exhibit cycles b. Lemmings jaegers & owls: Arctic tundra: ~4 yr cycle 2. Some example of influence of predators on communities have occurred when man accidentally introduced a new predator or purposefully removed a predator. a. Paines experiments with starfish: Keystone predator species i. Impact of insertion or deletion of predator into community b. Removals: Chinese killing birds during the Cultural Revolution (insect problems increased) 3. Biological Control: Human manipulation of predators for regulation of plants and animals. a. Predators are often used as biological control agent for pest species intentional introduction of predators to establish an upper trophic level to control lower trophic level. i. Keep the pest species below some threshold level; not total elimination ii. Reduce the use of chemical pesticides b. EXAMPLES: i. Biological control of herbivore: lady bird beetle on citrus; CA example 1. Lady bird beetle as biological control agents

2. The beetle decimated the scale populations and saved the citrus industry in CA ii. Biological control of plant/weed: Opuntia-Cactoblastic cactorum (prickly pear cactus-cactus moth: herbivore as regulator) 4. Introduction of Diseases: Fungus for Gypsy moth control, calicivirus for rabbit control 5. Plant recruitment of natural enemies: Exciting new research area a. When some plants are damaged by insect feeding, they also produce chemicals that are attractive to predators and parasitoids yelling for help. b. Plants and some herbivores can also produce food rewards to keep predator or parasite guards. How can predators respond to an increase in prey density? (3) ways 1. Functional response: a change in the rate of exploitation of prey (number consumed they eat more if more prey are available) by an individual predator as a result of a change in prey density a. Constrained by: i. Handling time: the time it takes to catch, kill and eat a prey. Often predation rate decreases at high prey densities. ii. Satiation and digestion: a predator can only eat so many prey: digestion and assimilation take time. iii. Prey distribution predator search behavior 3 types of functional responses: Predator consumes constant proportion of the prey population, regardless of prey density Predation rate decrease and handling times and predator satiation set an upper limit to the number of prey consumed Predator response lags at low prey densities due to low hunting efficiency or absence of a search image. As prey density increases either hunting efficiency or learning takes place. 2. Numerical response: density of predators in a given area increases with increased prey density (number of predators increases with an increase in prey). Can occur because: a. Predator reproduction e.g., higher survival of young b. Aggregation/immigration into an area of high prey density 3. Switching: when a predator has a choice of two different food items, the situation becomes more complex. a. Predator may have a fixed preference for one species of prey over the other, and this preference may not change as the composition of the available prey changes. b. The preference for one prey type depends on the proportions of prey types available. c. The predator switches witches from preferring one prey to preferring the other. d. Switching is important because it may stabilize the density fluctuations of the prey species (prey driven to low abundance does not go extinct, because the predator switches to another prey species)