Biology in Focus - Chapter 40

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CAMPBELL BIOLOGY IN FOCUS

© 2014 Pearson Education, Inc.

Urry • Cain • Wasserman • Minorsky • Jackson • Reece

Lecture Presentations by Kathleen Fitzpatrick and Nicole Tunbridge

40Population Ecology and the Distribution of Organisms

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Overview: Discovering Ecology

Ecology is the scientific study of the interactions between organisms and the environment

These interactions determine the distribution of organisms and their abundance

Modern ecology includes observation and experimentation

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The rediscovery of the nearly extinct harlequin toad in Costa Rica raises many ecological questions What environmental factors limit their geographic

distribution? What factors (food, pathogens) affect population

size?

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Figure 40.1

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Figure 40.2

Global ecology

Landscape ecology

Ecosystem ecology

Community ecology

Population ecology

Organismal ecology

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Global ecology is concerned with the biosphere, or global ecosystem, which is the sum of all the planet’s ecosystems

Global ecology examines the influence of energy and materials on organisms across the biosphere

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Figure 40.2a

Global ecology

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Landscape ecology focuses on the exchanges of energy, materials, and organisms across multiple ecosystems

A landscape (or seascape) is a mosaic of connected ecosystems

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Figure 40.2b

Landscape ecology

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Ecosystem ecology emphasizes energy flow and chemical cycling among the various biotic and abiotic components

An ecosystem is the community of organisms in an area and the physical factors with which they interact

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Figure 40.2c

Ecosystem ecology

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Community ecology deals with the whole array of interacting species in a community

A community is a group of populations of different species in an area

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Figure 40.2d

Community ecology

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Population ecology focuses on factors affecting population size over time

A population is a group of individuals of the same species living in an area

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Figure 40.2e

Population ecology

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Organismal ecology studies how an organism’s structure, physiology, and (for animals) behavior meet environmental challenges

Organismal ecology includes physiological, evolutionary, and behavioral ecology

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Figure 40.2f

Organismal ecology

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Concept 40.1: Earth’s climate influences the structure and distribution of terrestrial biomes

The long-term prevailing weather conditions in an area constitute its climate

Four major abiotic components of climate are temperature, precipitation, sunlight, and wind

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Abiotic factors are the nonliving chemical and physical attributes of the environment

Biotic factors are the other organisms that make up the living component of the environment

Macroclimate consists of patterns on the global, regional, and landscape level

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Global Climate Patterns

Global climate patterns are determined largely by solar energy and the planet’s movement in space

The warming effect of the sun causes temperature variations, which drive evaporation and the circulation of air and water

This causes latitudinal variations in climate

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Latitudinal variation in sunlight intensity is caused by the curved shape of Earth

Sunlight strikes the tropics, regions between 23.5 north and 23.5 south latitude, most directly

At higher latitudes, where sunlight strikes Earth at an oblique angle, light is more diffuse

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Figure 40.3a

Atmosphere

Low angle of incoming sunlight

Sun overhead at equinoxes

Low angle of incoming sunlight

Latitudinal variation in sunlight intensity

90S (South Pole)

0 (Equator)

23.5S (Tropic of Capricorn)

90N (North Pole)

23.5N (Tropic of Cancer)

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Global air circulation and precipitation patterns are initiated by intense solar radiation near the equator

Warm, wet air rising near the equator creates precipitation in the tropics

Dry air descending at 30 north and south latitudes causes desert conditions

This pattern of precipitation and drying is repeated at the 60 north and south latitudes and the poles

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Figure 40.3b

Global air circulation and precipitation patterns

30N

Northeast trades

66.5N (Arctic Circle)

30S

60S

Southeast trades

Westerlies

0

66.5S (Antarctic Circle)

30N

60N

Westerlies

Ascendingmoist airreleases moisture.

0

Descendingdry airabsorbs moisture.

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Variation in the speed of Earth’s rotation at different latitudes results in the major wind patterns

Trade winds blow east to west in the tropics Westerlies blow west to east in temperate zones

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Figure 40.3ba

Northeast trades

66.5N (Arctic Circle)

30S

60S

Southeast trades

Westerlies

0

66.5S (Antarctic Circle)

30N

60N

Westerlies

Global air circulation and precipitation patterns

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Figure 40.3bb

Global air circulation and precipitation patterns

30N

Ascendingmoist airreleases moisture.

0

Descendingdry airabsorbs moisture.

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Regional Effects on Climate

Climate is affected by seasonality, large bodies of water, and mountains

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Seasonality

Seasonal variations of light and temperature increase steadily toward the poles

Seasonality at high latitudes is caused by the tilt of Earth’s axis of rotation and its annual passage around the sun

Belts of wet and dry air straddling the equator shift throughout the year with the changing angle of the sun

Changing wind patterns affect ocean currents

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Figure 40.4

Constant tiltof 23.5

June solstice30N

December solstice

September equinox

60N

0 (equator)30S

March equinox

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Bodies of Water

Oceans, their currents, and large lakes moderate the climate of nearby terrestrial environments

The California Current carries cold water southward along western North America

The Gulf Stream carries warm water from the equator to the North Atlantic

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Figure 40.5

CaliforniaCurrent

PACIFICOCEAN

Gulf Stream

ATLANTICOCEAN

LabradorCurrent

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During the day, air rises over warm land and draws a cool breeze from the water across the land

As the land cools at night, air rises over the warmer water and draws cooler air from land back over the water, which is replaced by warmer air from offshore

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Figure 40.6

Mountainrange

Air flow

Ocean

Leeward sideof mountains

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Mountains

Rising air releases moisture on the windward side of a peak and creates a “rain shadow” as it absorbs moisture on the leeward side

Many deserts are found in the “rain shadow” of mountains

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Mountains affect the amount of sunlight reaching an area

In the Northern Hemisphere, south-facing slopes receive more sunlight than north-facing slopes

Every 1,000-m increase in elevation produces a temperature drop of approximately 6C

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Biomes are major life zones characterized by vegetation type (terrestrial biomes) or physical environment (aquatic biomes)

Climate is very important in determining why terrestrial biomes are found in certain areas

Climate affects the latitudinal patterns of terrestrial biomes

Climate and Terrestrial Biomes

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Figure 40.7

Temperate broadleaf forest

30S

Equator

Tropic of Capricorn

30NTropic ofCancer

Northern coniferous forest

High mountainsTundra

Polar ice

Tropical forestSavanna

ChaparralDesert

Temperate grassland

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A climograph plots the temperature and precipitation in a region

Biomes are affected not just by average temperature and precipitation, but also by the pattern of temperature and precipitation through the year

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Figure 40.8

Temperatebroadleafforest

Northernconiferousforest

Annual mean precipitation (cm)

15

Tropical forestDesert Temperate grassland

Arctic andalpinetundraA

nnua

l mea

n te

mpe

ratu

re (

C)

15

0

0

30

100 200 300 400

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Natural and human-caused disturbances alter the distribution of biomes

A disturbance is an event that changes a community by removing organisms and altering resource availability

For example, frequent fires kill woody plants preventing woodlands from establishing

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General Features of Terrestrial Biomes

Terrestrial biomes are often named for major physical or climatic factors and for vegetation

Terrestrial biomes usually grade into each other, without sharp boundaries

The area of intergradation, called an ecotone, may be wide or narrow

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Vertical layering is an important feature of terrestrial biomes, and in a forest it might consist of an upper canopy, low-tree layer, shrub understory, ground layer of herbaceous plants, forest floor, and root layer

Layering of vegetation provides diverse habitats for animals

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Terrestrial biomes can be characterized by distribution, precipitation, temperature, plants, and animals

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Tropical forest occurs in equatorial and subequatorial regions

Temperature is high year-round (25–29C) with little seasonal variation

In tropical rain forests, rainfall is relatively constant, while in tropical dry forests precipitation is highly seasonal

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Figure 40.9a

A tropical rain forest in Costa Rica

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Tropical forests are vertically layered, and competition for light is intense

Tropical forests are home to millions of animal species, including an estimated 5–30 million still undescribed species of insects, spiders, and other arthropods

Rapid human population growth is now destroying many tropical forests

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Savanna occurs in equatorial and subequatorial regions

Precipitation is seasonal Temperature averages 24–29C but is more

seasonally variable than in the tropics

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Figure 40.9b

A savanna in Kenya

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Grasses and forbs make up most of the ground cover

The dominant plant species are fire-adapted and tolerant of seasonal drought

Common inhabitants include insects and mammals such as wildebeests, zebras, lions, and hyenas

Fires set by humans may help maintain this biome

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Deserts occur in bands near 30 north and south of the equator and in the interior of continents

Precipitation is low and highly variable, generally less than 30 cm per year

Deserts may be hot (50C) or cold (–30C) with seasonal and daily temperature variation

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Figure 40.9c

Organ Pipe Cactus National Monument, Arizona

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Desert plants are adapted for heat and desiccation tolerance, water storage, and reduced leaf surface area

Common desert animals include scorpions, ants, beetles, snakes, lizards, migratory and resident birds, and seed-eating rodents; many are nocturnal

Urbanization and conversion to irrigated agriculture have reduced the natural biodiversity of some deserts

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Chaparral occurs in midlatitude coastal regions on several continents

Precipitation is highly seasonal with rainy winters and dry summers

Summer is hot (30C); fall, winter, and spring are cool (10–12C)

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Figure 40.9d

An area of chaparral in California

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The chaparral is dominated by shrubs and small trees; many plants are adapted to fire and drought

Animals include browsing mammals, insects, amphibians, small mammals, and birds

Humans have reduced chaparral areas through agriculture and urbanization

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Temperate grasslands occur at midlatitudes, often in the interior of continents

Precipitation is highly seasonal Winters are cold (often below −10C) and dry;

summers are hot (often near 30C) and wet

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Figure 40.9e

A grassland in Mongolia

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The dominant plants, grasses and forbs, are adapted to droughts and fire

Native mammals include large grazers such as bison and wild horses and small burrowers such as prairie dogs

Most grasslands have been converted to farmland

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Northern coniferous forest, or taiga, spans northern North America and Eurasia and is the largest terrestrial biome on Earth

Precipitation ranges from 30–70 cm Winters are cold; summers may be hot (e.g., Siberia

ranges from −50C to 20C)

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Figure 40.9f

A coniferous forest in Norway

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Conifers such as pine, spruce, fir, and hemlock dominate

The conical shape of conifers prevents too much snow from accumulating and breaking their branches

Animals include migratory and resident birds and large mammals such as moose, brown bears, and Siberian tigers

Periodic insect outbreaks kill vast tracts of trees Some forests are being logged at an alarming rate

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Temperate broadleaf forest is found at midlatitudes in the Northern Hemisphere, with smaller areas in Chile, South Africa, Australia, and New Zealand

Significant amounts of precipitation fall during all seasons as rain or snow

Winters average 0C; summers are hot and humid (near 35C)

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Figure 40.9g

A temperate broadleaf forest in New Jersey

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Dominant plants include deciduous trees in the Northern Hemisphere and evergreen eucalyptus in Australia

In the Northern Hemisphere, many mammals hibernate in the winter; birds migrate to warmer areas

These forests have been heavily settled on all continents but are recovering in places

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Tundra covers expansive areas of the Arctic; alpine tundra exists on high mountaintops at all latitudes

Precipitation is low in arctic tundra and higher in alpine tundra

Winters are cold (below −30C); summers are relatively cool (less than 10C)

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Figure 40.9h

Dovrefjell National Park, Norway

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Vegetation is herbaceous (mosses, grasses, forbs, dwarf shrubs, trees, and lichen)

Permafrost, a permanently frozen layer of soil, restricts growth of plant roots

Mammals include musk oxen, caribou, reindeer, bears, wolves, and foxes; many migratory bird species nest in the summer

Settlement is sparse, but tundra has become the focus of oil and mineral extraction

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Concept 40.2: Aquatic biomes are diverse and dynamic systems that cover most of Earth

Aquatic biomes account for the largest part of the biosphere in terms of area

They show less latitudinal variation than terrestrial biomes

Marine biomes have salt concentrations of about 3% The largest marine biome is made of oceans, which

cover about 75% of Earth’s surface and have an enormous impact on the biosphere

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Freshwater biomes have salt concentrations of less than 0.1%

Freshwater biomes are closely linked to soils and the biotic components of the surrounding terrestrial biome

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Video: Clownfish Anemone

Video: Coral Reef

Video: Hydrothermal Vent

Video: Shark Eating a Seal

Video: Swans Taking Flight

Video: Tubeworms

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Aquatic biomes can be characterized by their physical and chemical environment, geological features, photosynthetic organisms, and heterotrophs

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Wetlands and estuaries are among the most productive habitats on Earth

Wetlands are inundated by water at least sometimes and support plants adapted to water-saturated soil

An estuary is a transition area between river and sea; salinity varies with the rise and fall of the tides

High organic production and decomposition in these biomes result in low levels of dissolved oxygen

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Figure 40.10a

A basin wetland in the United Kingdom

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Wetlands can develop in shallow basins, along flooded river banks, or on the coasts of large lakes

Estuaries include a complex network of tidal channels, islands, and mudflats

Plants are adapted to growing in periodically anaerobic, water-saturated soils

Wetland plants include cattails and sedges; estuaries are characterized by saltmarsh grasses

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Wetlands are home to diverse invertebrates and birds, as well as frogs and alligators

Estuaries support an abundance of marine invertebrates, fish, waterfowl, and marine mammals

Humans activities have destroyed up to 90% of wetlands and disrupted estuaries worldwide

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Lakes vary in size from small ponds to very large lakes

Temperate lakes may have a seasonal thermocline; tropical lowland lakes have a year-round thermocline

Oligotrophic lakes are nutrient-poor and generally oxygen-rich

Eutrophic lakes are nutrient-rich and often depleted of oxygen if ice covered in winter

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Figure 40.10b

An oligotrophic lake in Alberta, Canada

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Eutrophic lakes have more surface area relative to depth than oligotrophic lakes

Rooted and floating aquatic plants live in the shallow and well-lighted littoral zone close to shore

Water is too deep in the limnetic zone to support rooted aquatic plants; the primary producers are phytoplankton

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Zooplankton are drifting heterotrophs that graze on the phytoplankton

Invertebrates live in the benthic zone Fishes live in all zones with sufficient oxygen Human-induced nutrient enrichment can lead to

algal blooms, oxygen depletion, and fish kills

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Streams and rivers have varying environmental conditions from headwater to mouth

Headwater streams are generally cold, clear, turbulent, swift, and oxygen-rich; they are often narrow and rocky

Downstream waters form rivers and are generally warmer and more turbid; they are often wide and meandering and have silty bottoms

Salt and nutrients increase from headwaters to mouth; oxygen content decreases from headwaters to mouth

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Figure 40.10c

A headwater stream in Washington

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Headwater streams may be rich in phytoplankton or rooted aquatic plants

A diversity of fishes and invertebrates inhabit unpolluted rivers and streams

Pollution degrades water quality and kills aquatic organisms

Dams impair natural flow of stream and river ecosystems

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Intertidal zones are periodically submerged and exposed by the tides

Intertidal organisms are challenged by variations in temperature and salinity and by the mechanical forces of wave action

Oxygen and nutrient levels are high Substrate varies from rocky to sandy

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Figure 40.10d

A rocky intertidal zone on the Oregoncoast

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Protected sandy zones support sea grass and algae; rocky zones support attached marine algae

In rocky zones, many animals have structural adaptations for attaching to rock

In sandy zones, worms, clams, and crustaceans bury themselves in sand

Other animals include sponges, sea anemones, and small fishes

Oil pollution has disrupted many intertidal areas

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Coral reefs are formed from the calcium carbonate skeletons of corals (cnidarians)

Shallow reef-building corals live in the photic zone in warm (about 20–30C), clear water; deep-sea corals live at depths of 200–1,500 m

Corals require high oxygen concentrations and a solid substrate for attachment

A coral reef progresses from a fringing reef to a barrier reef to a coral atoll

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Figure 40.10e

A coral reef in the Red Sea

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Corals are the predominant animals on the reef They form mutualisms with symbiotic algae that

provide them organic molecules A high diversity of fish and invertebrates inhabit coral

reefs Coral collection, overfishing, global warming, and

pollution all contribute to reduction in coral populations

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The oceanic pelagic zone is constantly mixed by wind-driven oceanic currents

Oxygen levels are high Turnover in temperate oceans renews nutrients in

the photic zones This biome covers approximately 70% of Earth’s

surface

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Figure 40.10f

Open ocean near Iceland

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Phytoplankton and zooplankton are the dominant organisms in this biome; also found are free-swimming animals

Zooplankton includes protists, worms, krill, jellies, and invertebrate larvae

Other animals include squids, fishes, sea turtles, and marine mammals

Overfishing has depleted fish stocks Humans have polluted oceans with dumping of waste

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The marine benthic zone consists of the seafloor Organisms in the very deep benthic (abyssal) zone

are adapted to continuous cold and high water pressure

Substrate is mainly soft sediments; some areas are rocky

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Shallow areas contain seaweeds and filamentous algae

In dark, hot environments near deep-sea hydrothermal vents of volcanic origin, the food producers are chemoautotrophic prokaryotes

Giant tube worms, arthropods, and echinoderms are abundant heterotrophs near hydrothermal vents

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Figure 40.10g

A deep-sea hydrothermal vent community

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Coastal benthic communities include invertebrates and fishes

Overfishing and dumping of organic waste have depleted fish populations

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Zonation in Aquatic Biomes

Many aquatic biomes are stratified into zones or layers defined by light penetration, temperature, and depth

The upper photic zone has sufficient light for photosynthesis, while the lower aphotic zone receives little light

The photic and aphotic zones make up the pelagic zone

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The organic and inorganic sediment at the bottom of all aquatic zones is called the benthic zone

The communities of organisms in the benthic zone are collectively called the benthos

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Figure 40.11

Zonation in a lake

Littoralzone Limnetic

zone

Pelagiczone

Photiczone

Aphoticzone

Benthiczone

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In oceans and most lakes, a temperature boundary called the thermocline separates the warm upper layer from the cold deeper water

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Communities in aquatic biomes vary with depth, light penetration, distance from shore, and position in the pelagic or benthic zone

Most organisms occur in the relatively shallow photic zone

The aphotic zone in oceans is extensive but harbors little life

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Concept 40.3: Interactions between organisms and the environment limit the distribution of species

Species distributions are the result of ecological and evolutionary interactions through time

Ecological time is the minute-to-minute time frame of interactions between organisms and the environment

Evolutionary time spans many generations and captures adaptation through natural selection

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Events in ecological time can lead to evolution For example, Galápagos finches with larger beaks

were more likely to survive a drought, as they could eat the available larger seeds

As a result, the average beak size was larger in the next generation

This resulted in an evolutionary change

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Ecologists ask questions about where species occur and why species occur where they do

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Figure 40.12-1

Why is speciesX absent froman area?

Does dispersallimit its

distribution?

Area inaccessibleor insufficient time

Do biotic factors(other species)

limit itsdistribution?

Yes

No

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Figure 40.12-2

Why is speciesX absent froman area?

Does dispersallimit its

distribution?

Area inaccessibleor insufficient time

Predation, parasitism,competition, disease

Do biotic factors(other species)

limit itsdistribution?

Do abiotic factorslimit its

distribution?

Yes

No

YesNo

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Figure 40.12-3

Chemicalfactors

Why is speciesX absent froman area?

Does dispersallimit its

distribution?

Area inaccessibleor insufficient time

Predation, parasitism,competition, disease

Water, oxygen, salinity,pH, soil nutrients, etc.

Do biotic factors(other species)

limit itsdistribution?

Temperature, light,soil structure, fire,moisture, etc.

Do abiotic factorslimit its

distribution?

Physicalfactors

Yes

No

YesNo

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Dispersal and Distribution

Dispersal is the movement of individuals away from centers of high population density or from their area of origin

Dispersal contributes to the global distribution of organisms

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Transplants indicate if dispersal is a key factor limiting species distributions

Transplants include organisms that are intentionally or accidentally relocated from their original distribution

If a transplant is successful, it indicates that the species’ potential range is larger than its actual range

Species transplants can disrupt the communities or ecosystems to which they have been introduced

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Biotic Factors

Biotic factors that affect the distribution of organisms may include Predation Herbivory

For example, sea urchins can limit the distribution of seaweeds

Mutualism Parasitism Competition

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Figure 40.13

Control (both urchinsand limpets present)

100 Both limpets and urchinsremoved

Only urchinsremoved

Only limpets removed

Limpet

Sea urchin

August1982

August1983

February1983

February1984

80

60

40

20

0

Seaw

eed

cove

r (%

)

Results

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Abiotic Factors

Abiotic factors affecting the distribution of organisms include Temperature Water and oxygen Salinity Sunlight Rocks and soil

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Temperature is an important environmental factor in the distribution of organisms because of its effects on biological processes

Cells may freeze and rupture below 0C, while most proteins denature above 45C

Most organisms function best within a specific temperature range

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Availability of water and oxygen is an important factor in species distribution

Desert organisms exhibit adaptations for water conservation

Water affects oxygen availability, as oxygen diffuses slowly in water

Oxygen concentrations can be low in deep oceans and deep lakes

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Salinity, salt concentration, affects the water balance of organisms through osmosis

Most aquatic organisms are restricted to either freshwater or saltwater habitats

Few terrestrial organisms are adapted to high-salinity habitats

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Sunlight is the energy source for photosynthetic organisms and, as such, can limit their distribution

In aquatic environments most photosynthesis occurs near the surface where sunlight is available

Shading by the canopy drives intense competition for light in forests

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Rocks and soil have many characteristics that limit the distribution of plants and thus the animals that feed on them Physical structure pH Mineral composition

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Concept 40.4: Dynamic biological processes influence population density, dispersion, and demographics

Population ecology explores how biotic and abiotic factors influence density, distribution, and size of populations

A population is a group of individuals of a single species living in the same general area

Populations are described by their boundaries and size

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Density and Dispersion

Density is the number of individuals per unit area or volume

Dispersion is the pattern of spacing among individuals within the boundaries of the population

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Density: A Dynamic Perspective

In most cases, it is impractical or impossible to count all individuals in a population

Sampling techniques can be used to estimate densities and total population sizes

Population density can be estimated by extrapolation from small samples or an index of population size (e.g., number of nests)

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Density is the result of an interplay between processes that add individuals to a population and those that remove individuals

Additions occur through birth and immigration, the influx of new individuals from other areas

Removal of individuals occurs through death and emigration, the movement of individuals out of a population

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Figure 40.14

Births andimmigrationadd individuals toa population.

Deaths andemigrationremove individualsfrom a population.

Births

Immigration

Deaths

Emigration

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Patterns of Dispersion

Environmental and social factors influence the spacing of individuals in a population

The most common pattern of dispersion is clumped, in which individuals aggregate in patches

A clumped dispersion may be influenced by resource availability and behavior

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Figure 40.15

(a) Clumped

(c) Random(b) Uniform

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Figure 40.15a

(a) Clumped

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Figure 40.15b

(b) Uniform

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Figure 40.15c

(c) Random

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Figure 40.15d

(a) Clumped (c) Random(b) Uniform

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A uniform dispersion is one in which individuals are evenly distributed

It may be influenced by social interactions such as territoriality, the defense of a bounded space against other individuals

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In a random dispersion, the position of each individual is independent of other individuals

It occurs in the absence of strong attractions or repulsions

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Demographics

Demography is the study of the vital statistics of a population and how they change over time

Death rates and birth rates are of particular interest to demographers

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Life Tables

A life table is an age-specific summary of the survival pattern of a population

It is best made by following the fate of a cohort, a group of individuals of the same age, from birth to death

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Survivorship Curves

A survivorship curve is a graphic way of representing the data in a life table

Survivorship curves plot the proportion or numbers of a cohort still alive at each age

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Survivorship curves can be classified into three general types Type I: low death rates during early and middle life

and an increase in death rates among older age groups

Type II: a constant death rate over the organism’s life span

Type III: high death rates for the young and a lower death rate for survivors

Many species are intermediate to these curves

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Figure 40.16

1,000

Percentage of maximum life span

III

Num

ber o

f sur

vivo

rs (l

og s

cale

)

100

10

01

10050

II

I

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Reproductive Rates

For species with sexual reproduction, demographers often concentrate on females in a population

A reproductive table, or fertility schedule, is an age-specific summary of the reproductive rates in a population

It describes the reproductive patterns of a population

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Table 40.1

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Concept 40.5: The exponential and logistic models describe the growth of populations

Unlimited growth occurs under ideal conditions; in nature, growth is limited by various factors

Ecologists study growth in both idealized and realistic conditions

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Per Capita Rate of Increase

Change in population size can be defined by the equation

If immigration and emigration are ignored, a population’s growth rate (per capita increase) equals birth rate minus death rate

− −Change inpopulation

sizeBirths

Immigrantsentering

populationDeaths

Emigrantsleaving

population

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The population growth rate can be expressed mathematically as

where N is the change in population size, t is the time interval, B is the number of births, and D is the number of deaths

ΔNΔt B −D

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Births and deaths can be expressed as the average number of births and deaths per individual during the specified time interval

where b is the annual per capita birth rate, m (for mortality) is the per capita death rate, and N is population size

B bND mN

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The population growth equation can be revised

ΔNΔt bN −mN

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The per capita rate of increase (r) is given by

r b − m

Zero population growth (ZPG) occurs when the birth rate equals the death rate (r 0)

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Change in population size can now be written as

ΔNΔt rN

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Instantaneous growth rate can be expressed as

where rinst is the instantaneous per capita rate

of increase

dNdt

rinstN

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Exponential Growth

Exponential population growth is population increase under idealized conditions

Under these conditions, the rate of increase is at its maximum, denoted as rmax

The equation of exponential population growth is

dNdt

rmaxN

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Exponential population growth results in a J-shaped curve

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Figure 40.17

1,000

Number of generations

Popu

latio

n si

ze (N

) 1.0N

100

0 155

2,000

1,500

500

0.5N dt dN

dt dN

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The J-shaped curve of exponential growth characterizes populations in new environments or rebounding populations For example, the elephant population in Kruger

National Park, South Africa, grew exponentially after hunting was banned

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Figure 40.18

6,000

Year

Elep

hant

pop

ulat

ion

0

2,000

8,000

4,000

1900 1930 1940 1950 1960 19701910 1920

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Figure 40.18a

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Carrying Capacity

Exponential growth cannot be sustained for long in any population

A more realistic population model limits growth by incorporating carrying capacity

Carrying capacity (K) is the maximum population size the environment can support

Carrying capacity varies with the abundance of limiting resources

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The Logistic Growth Model

In the logistic population growth model, the per capita rate of increase declines as carrying capacity is reached

The logistic model starts with the exponential model and adds an expression that reduces per capita rate of increase as N approaches K

dNdt rmaxN

(K −N)K

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Table 40.2

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The logistic model of population growth produces a sigmoid (S-shaped) curve

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Figure 40.19

1,000

Number of generations

Popu

latio

n si

ze (N

) 1.0N

100

0 155

2,000

1,500

500

1.0N

dt dN

Exponentialgrowth

Population growthbegins slowing here.

K 1,500

dt dN (1,500 N)

1,500

Logistic growth

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The Logistic Model and Real Populations

The growth of many laboratory populations, including paramecia, fits an S-shaped curve when resources are limited

These organisms are grown in a constant environment lacking predators and competitors

Animation: Population Ecology

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Figure 40.20

1,000

Time (days)

Num

ber o

f Par

amec

ium

/mL

100

0 155

(a) A Paramecium populationin the lab

800

200

400

600

Time (days)140

00 160

30

(b) A Daphnia (water flea) population in the lab

40 6020 100 12080N

umbe

r of D

aphn

ia/5

0 m

L

180

60

150

120

90

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Figure 40.20a

1,000

Time (days)

Num

ber o

fPa

ram

eciu

m/m

L

100

0 155

(a) A Paramecium populationin the lab

800

200

400

600

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Figure 40.20b

Time (days)140

00 160

30

(b) A Daphnia (water flea) population in the lab

40 6020 100 12080

Num

ber o

fD

aphn

ia/5

0 m

L

180

60

150120

90

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Some populations overshoot K before settling down to a relatively stable density

Some populations fluctuate greatly and make it difficult to define K

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The logistic model fits few real populations but is useful for estimating possible growth

Conservation biologists can use the model to estimate the critical size below which populations may become extinct

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Concept 40.6: Population dynamics are influenced strongly by life history traits and population density

An organism’s life history comprises the traits that affect its schedule of reproduction and survival The age at which reproduction begins How often the organism reproduces How many offspring are produced during each

reproductive cycle

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“Trade-offs” and Life Histories

Organisms have finite resources, which may lead to trade-offs between survival and reproduction

Selective pressures influence the trade-off between the number and size of offspring

Some plants produce a large number of small seeds, ensuring that at least some of them will grow and eventually reproduce

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Figure 40.21

Dandelions growquickly and releasea large number oftiny fruits.

The Brazil nut tree(above), produces a moderate number oflarge seeds in pods (left).

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Figure 40.21a

Dandelions grow quickly and release a large number of tiny fruits.

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Figure 40.21ba

The Brazil nut tree produces a moderate number oflarge seeds in pods.

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Figure 40.21bb

The Brazil nut tree produces a moderate number of large seeds in pods.

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Other types of plants produce a moderate number of large seeds that provide a large store of energy that will help seedlings become established

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K-selection, or density-dependent selection, selects for life history traits that are sensitive to population density

r-selection, or density-independent selection, selects for life history traits that maximize reproduction

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Population Change and Population Density

In density-independent populations, birth rate and death rate do not change with population density

In density-dependent populations, birth rates fall and death rates rise with population density

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Figure 40.22

When populationdensity is low, b m. Asa result, the populationgrows until the densityreaches Q.

When populationdensity is high, m b, and the populationshrinks until the density reaches Q.

Equilibrium density (Q)

Density-dependentbirth rate (b)

Density-independentdeath rate (m)

Birt

h or

dea

th ra

tepe

r cap

ita

Population density

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Mechanisms of Density-Dependent Population Regulation

Density-dependent birth and death rates are an example of negative feedback that regulates population growth

Density-dependent birth and death rates are affected by many factors, such as competition for resources, territoriality, disease, predation, toxic wastes, and intrinsic factors

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Competition for resources occurs in crowded populations; increasing population density intensifies competition for resources and results in a lower birth rate

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Figure 40.23a

Competition for resources

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Toxic wastes produced by a population can accumulate in the environment, contributing to density-dependent regulation of population size

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Figure 40.23d

Toxic wastes 5 m

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Predation may increase with increasing population size due to predator preference for abundant prey species

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Figure 40.23b

Predation

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Territoriality can limit population density when space becomes a limited resource

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Figure 40.23e

Territoriality

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Disease transmission rates may increase with increasing population density

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Figure 40.23c

Disease

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Intrinsic factors (for example, physiological factors like hormonal changes) appear to regulate population size

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Figure 40.23f

Intrinsic factors

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Population Dynamics

The study of population dynamics focuses on the complex interactions between biotic and abiotic factors that cause variation in population size

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Stability and Fluctuation

Long-term population studies have challenged the hypothesis that populations of large mammals are relatively stable over time

Both weather and predator population can affect population size over time For example, the moose population on Isle Royale

collapsed during a harsh winter and when wolf numbers peaked

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Figure 40.24

Num

ber o

f wol

ves

Num

ber o

f moo

se

Year

Wolves Moose

0

10

20

30

1,000

0

2,500

1,500

500

1955 1985 1995 20051965 1975

2,000

50

40

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Immigration, Emigration, and Metapopulations

Metapopulations are groups of populations linked by immigration and emigration

Local populations within a metapopulation occupy patches of suitable habitat surrounded by unsuitable habitat

Populations can be replaced in patches after extinction or established in new unoccupied habitats through migration

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Figure 40.25

Unoccupied patch

EUROPE

AlandIslands

Occupied patch5 km

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Figure 40.25a

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Figure 40.UN01a

Number of generations

Popu

latio

n si

ze (N

) rmax N dt

dN

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Figure 40.UN01b

rmax N dt dN (K − N)

K

K carrying capacity

Number of generations

Popu

latio

n si

ze (N

)

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Figure 40.UN02

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Figure 40.UN03

Mea

n he

ight

(cm

)

Seed collection sites

1,000

0

3,000

100

50

2,000

Alti

tude

(m)

0

Sierra NevadaGreat Basin

Plateau