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Because of this ability to alter ecosystems, exotic invaders are considered major agents in driving native species to extinction and are thought to be responsible for an estimated 40 per

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exists The most attractive for

indus-trial applications is H2production by

photosynthetic microbes These

mi-croorganisms, such as microscopic

algae, cyanobacteria, and

photosyn-thetic bacteria, use sunlight as an

en-ergy source and water to generate

hy-drogen

Butanol can be produced by the

fermentation of sugars similar to the

ethanol production The most

well-known pathway of butanol

genera-tion is fermentagenera-tion by bacterium

Clostridium acetobutylicum Substrates

utilized for butanol production—

starch, molasses, cheese whey, and

lignocellulosic materials—are exactly

the same as for ethanol

fermenta-tion The biological production by fermentation is

not economically attractive because of low levels of

product concentrations and high cost of product

re-covery compared to the chemical process

Uses of Biofuels

With increasing energy demands and oil prices,

etha-nol has become a valuable option as an alternative

transportation fuel The Energy Policy Act of 2005

in-cluded a requirement to increase the production of

ethanol from 15 to 28 billion liters by 2012 Beginning

in 2008, a majority of fuel stations in the United States

were selling gasoline with 10 percent ethanol in it

Nearly all cars can use E10, fuel that is 10 percent

etha-nol Blending ethanol with gasoline oxygenates the

fuel mixture, which burns more completely and

pro-duces fewer harmful CO emissions Another

environ-mental benefit of ethanol is that it degrades in the

soil, whereas petroleum-based fuels are more

resis-tant to degradation and have many damaging effects

when accidentally discharged into the environment

However, a liter of ethanol has significantly less

en-ergy content than a liter of gasoline, so vehicles must

be refueled more often Ethanol is also more

expen-sive than gasoline, although rising prices of gasoline

could cancel that disadvantage In addition,

carcino-genic aldehydes, such as formaldehyde, are produced

when ethanol is burned in internal combustion

en-gines Carbon dioxide, a major greenhouse gas, forms

as well Moreover, the widely used fuel mix that is 85

percent ethanol and 15 percent gasoline (the E85

blend) requires specially equipped “flexible fuel”

en-gines In the United States, only a fraction of all cars are considered “flex fuel” vehicles By comparison, however, most cars in Brazil have flex engines Begin-ning in 1977, the Brazilian government made using ethanol as a fuel for cars mandatory Brazil has the largest and most successful “ethanol for fuel” gram in the world As a result of this successful pro-gram, the country reached complete self-sufficiency

in energy supply in 2006

Biodiesel performs similarly to diesel and can be used in unmodified diesel engines of trucks, tractors, and other vehicles, and it is better for the environ-ment Burning biodiesel produces fewer emissions than petroleum-based diesel; it is essentially free of sulfur and aromatics and emits less CO Additionally, biodiesel is less toxic to the soil Biodiesel is often blended with petroleum diesel in different ratios of 2,

5, or 20 percent The most common blend is B20,

or 20 percent biodiesel to 80 percent diesel fuel Biodiesel can be used as a pure fuel (100 percent or B100), but pure fuel is not suitable for winter because

it thickens in cold temperatures In addition, B100 is a solvent that degrades engines’ rubber hoses and gas-kets Moreover, biodiesel energy content is less than

in diesel In general, biodiesel is not used as widely as ethanol However, biodiesel users include the United States Postal Service; the U.S Departments of De-fense, Energy, and Agriculture; national parks; school districts; transit authorities; and public-utilities, waste-management, and recycling companies across the United States In January, 2009, Continental Airlines successfully demonstrated the use of a biodiesel

This Volvo car runs on bioethanol, a biofuel manufactured from common household trash (AP/Wide World Photos)

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ture from plants and algae (50 percent to 50 percent)

to fly its Boeing 737-800

In the 1985 Mel Gibson movie Mad Max Beyond

Thunderdome, a futuristic city was run on methane that

was generated by pig manure In reality, methane can

be a very good alternative fuel It has a number of

ad-vantages over other fuels produced by

microorgan-isms First, it is easy to make and can be generated

lo-cally, which does not require distribution Extensive

natural gas infrastructure is already in place to be

uti-lized Second, the utilization of methane as a fuel is

an attractive way to reduce wastes such as manure,

wastewater, or municipal and industrial wastes In

lo-cal farms, manure is fed into digesters (bioreactors)

where microorganisms metabolize it into methane

Methane can be used to fuel electrical generators to

produce electricity In China, millions of small farms

have simple small underground digesters near the

farm houses There are several landfill gas facilities in

the United States that generate electricity using

meth-ane San Francisco has extended its recycling

pro-gram to include conversion of dog waste into

meth-ane to produce electricity and to heat homes With a

dog population of 120,000 this initiative promises to

generate a significant amount of fuel with a huge

re-duction of waste at the same time Methane was used

as a fuel for vehicles for a number of years Several

Volvo car models with bi-fuel engines were made to

run on compressed methane with gasoline as a backup

Biogas can also be compressed, like methane, and

used to power motor vehicles

In many countries, millions of small farms

main-tain a simple digester for biogas production to

gener-ate energy Currently, there are more than five million

household digesters in China, used by people mainly

for cooking and lighting, and there are more than

one million biogas plants of various capacities in

India

Utilization of methane and biogas as an energy

source in place of fossil fuels is providing significant

environmental and economic benefits Biofuels are

essentially nonpolluting, although their utilization

re-sults in production of CO2and contributes to global

warming, though with less impact on Earth’s climate

than methane itself as a greenhouse gas Even though

the use of methane and biogas as energy sources

re-leases CO2, the process as a whole can be considered

“CO2neutral” in that the released CO2can be

assimi-lated by their producers, archaea and bacteria

Some examples of biomass use as an alternative

energy source include burning wood or agricultural residue to heat homes This is an inefficient use of energy—typically only 5-15 percent of the biomass en-ergy is actually utilized Using biomass that way pro-duces harmful indoor air pollutants such as carbon monoxide Yet biomass is an almost “free” resource costing only labor to collect Biomass supplies more than 15 percent of the world’s energy consumption Biomass is the top source of energy in developing countries; in some countries it provides more than 90 percent of the energy used

Hydrogen powered U.S rockets for many years To-day, a growing number of automobile manufacturers around the world are making prototype hydrogen-powered vehicles Only water is emitted from the tail-pipe—no greenhouse gases The car is moved by a motor that runs on electricity generated in the fuel cell via a chemical reaction between H2and O2 Hy-drogen vehicles offer quiet operation, rapid accelera-tion, and low maintenance costs During peak time, when electricity is expensive, fuel-cell hydrogen cars could provide power for homes and offices Hydro-gen for these applications is obtained mainly from natural gas (methane and propane) via steam reform-ing Biohydrogen is used in experimental applica-tions only Many problems need to be overcome be-fore biohydrogen can be easily available One of the reasons for the delayed acceptance of biohydrogen

is the difficulty of its production on a cost-effective basis For biohydrogen power to become a reality, tre-mendous research and investment efforts are neces-sary

Butanol can be used as transportation fuel It con-tains almost as much energy as gasoline and more en-ergy than ethanol for a particular volume Unlike 85 percent ethanol, a butanol/gasoline mix (E85 blend) can be used in cars designed for gasoline without mak-ing any changes to the engine

Sergei A Markov

Further Reading

Chisti, Yusuf “Biodiesel from Microalgae.” Biotechnol-ogy Advances 25, no 3 (2007): 294-306.

Glazer, Alexander N., and Hiroshi Nikaido Microbial Biotechnology: Fundamentals of Applied Microbiology.

New York: W H Freeman, 2007

Service, Robert F “The Hydrogen Backlash.” Science

305, no 5686 (August 13, 2004): 958-961

Wald, Matthew L “Is Ethanol for the Long Haul?” Sci-entific American 296, no 1 (January, 2007): 42-49.

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Wright, Richard T Environmental Science: Towards a

Sustainable Future 9th ed Englewood Cliffs, N.J.:

Prentice Hall, 2004

Web Site

AE Biofuels

http://www.alternative-energy-news.info/

technology/biofuels/

See also: Brazil; Energy economics; Ethanol;

Meth-ane; Sugars; Sustainable development

Biogeochemical cycles See Carbon

cycle; Geochemical cycles;

Hydrology and the hydrologic

cycle; Nitrogen cycle; Phosphorus

cycle; Sulfur cycle

Biogeographic realms See Ecozones

and biogeographic realms

Biological invasions

Categories: Environment, conservation, and

resource management; pollution and waste

disposal

A biological invasion is an enormous increase in the

numbers of a type of organism entering an ecosystem

that the organism previously was not inhabiting The

“invading” organism may be an infectious virus, a

bacterium, a plant, or an animal.

Background

Species introduced to an area from somewhere

out-side that area are referred to as alien or exotic species

or as invaders Because the exotic species is not native

to the new area, it is often unsuccessful in establishing

a viable population and disappears The fossil record,

as well as historical documentation, indicates that this

is the fate of many exotic species as they move from

their native habitats to invade new environments

Oc-casionally, however, an invading species finds the new environment to its liking; in this case the invader may become so successful in exploiting its new habitat that

it can completely alter the ecological balance of an ecosystem, destroying biodiversity and altering the lo-cal biologilo-cal hierarchy Because of this ability to alter ecosystems, exotic invaders are considered major agents in driving native species to extinction and are thought to be responsible for an estimated 40 percent

of all known extinctions of land animals beginning in the year 1600

Biological invasions by notorious species consti-tute a significant component of Earth’s history In general, large-scale climatic changes and geological crises are at the origin of massive exchanges of flora and fauna On a geologic timescale, invasions of spe-cies from one continent to another are true evolution-ary processes, just as speciation and extinction are

On a smaller scale, physical barriers such as oceans, mountains, and deserts can be overcome by many or-ganisms as their populations expand Oror-ganisms can

be carried by water in rivers or ocean currents, trans-ported by wind, or carried by other species as they mi-grate seasonally or to escape environmental pres-sures However, the geological and historical records

of the Earth also show that specific biological inva-sions by exotic species have altered the course of world history The extinction of genetically distinct populations is the least reversible of all global changes, and evidence suggests that biological invasions con-tribute substantially to an increase in the rate of ex-tinction within ecosystems

Humans have transplanted species throughout his-tory, to the point where most people are not aware of the distinction between native and exotic species liv-ing in their region Recent increases in interconti-nental invasion rates by exotic species, brought about primarily by human activity, create important ecologi-cal problems for the recipient lands Among animals, the most notorious recent invaders of North America have been the house mouse and the Norway rat; others include the wild boar, donkey, horse, nutria, Pierid butterfly, house sparrow, starling, Africanized (“killer”) bee, tiger mosquito, and red fox One of the most destructive invaders is the house cat More than seventy million domestic and feral cats live in the United States, and they are efficient at hunting small mammals and birds Domestic cats are credited with killing twenty million birds annually in Great Britain

It would seem logical to assume that invading

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cies might add to the biodiversity of a region, but

many invaders have the opposite effect The new

spe-cies are often opportunistic and successful predators

that eliminate native species not adapted to their

pres-ence For example, the brown tree snake was

acciden-tally introduced to Guam during World War II as a

stowaway on military cargo ships, and the snakes have

eliminated most of the island’s birds The snakes are

credited with the extinction of one-third of the

is-land’s native bird species, and the surviving bird

pop-ulation is so decimated that birds are rarely seen or

heard The invasion of the brown tree snake has

unal-terably reduced the biological diversity of Guam

Ecosystem Alteration

The invasion of an ecosystem by an exotic species

can effectively alter ecosystem processes An invading

species does not simply consume or compete with

na-tive species but can actually change the rules of

exis-tence within the ecosystem by altering processes such

as primary productivity, decomposition, hydrology,

geomorphology, nutrient cycling, and natural

distur-bance regimes Invading exotic species may also drive

out native species by competing with them for re-sources One of the exotic invaders of the North American continent is the zebra mussel, which came

to the United States in 1986 in the ballast water of oceangoing vessels; it was carried from the Elbe or Rhine River in Europe and released into the water of the St Clair River near Detroit, Michigan The mussel larvae found biological conditions in the Great Lakes ideal The mussel now exists in all the Great Lakes, and after the catastrophic flood of 1993 the mussels were sighted in the Mississippi River Basin Mussel density in certain locations of the Great Lakes is known to be astonishing—greater than 94,000 indi-viduals per square meter In 1990, the Detroit Edison power plant discovered a water intake pipe blocked by

a mussel population density of 700,000 mussels per square meter When they reach high population den-sities, the mussels are able to filter virtually all the larger plankton from the water The planktonic food chain of the Great Lakes, which supports Great Lakes fisheries, may decline so much that higher trophic species will be deprived of their vital plankton food sources The mussels also cause a demise of native bi-valves through competition for food and because they attach themselves to the shells of other bivalves Forests

The invasion of native forests by non-native insects and microorganisms has been devastating on many continents The white pine blister rust and the balsam woolly adelgid have invaded both commercial and preserved forestlands in North America Both exotics were brought to North America in the late 1800’s

on nursery stock from Europe The balsam woolly adelgid attacks fir trees and causes death within two to seven years by causing chemical damage and by feed-ing on the tree’s vascular tissue The adelgid has killed nearly every adult cone-bearing fir tree in the south-ern Appalachian Mountains The white pine blister rust attacks five-needle pines; in the western United States fewer than 10 pine trees in 100,000 are resis-tant, and since white pine seeds are an essential food source for bears and other animals, the loss of the trees has had severe consequences across the forest food chain

Beginning in the 1800’s the deciduous forests of eastern North America were attacked numerous times

by waves of invading exotic species and diseases One

of the most notable invaders is the gypsy moth, which consumes a variety of tree species Other invaders of

The tunicate is an invasive species that grows in the habitat of

anemones and sea cucumbers (AP/Wide World Photos)

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eastern forests have virtually eliminated the once

dominant American chestnut and the American elm

Other tree species that continue to decline because of

new invaders include the American beech, mountain

ash, white birch, butternut, sugar maple, flowering

dogwood, and eastern hemlock It is widely accepted

that the invasion of exotic species is the single greatest

threat to the diversity of deciduous forests in North

America

Effects on Humans, and Humans as Invaders

Some introduced exotic species are beneficial to

hu-manity It would be impossible to support the present

world human population entirely on species native to

their regions However, many invading species

de-grade human health and wealth, and others affect the

structure of ecosystems or the ability to maintain

na-tive biodiversity Many invading species can act as

vec-tors of disease: Examples include bubonic plague,

vectored by rats; a host of diseases transmitted

be-tween human populations during first contacts,

cluding smallpox, polio, influenza, and venereal

in-fections; and malaria, dengue fever, Ross River fever,

and eastern equine encephalitis, carried by

mosqui-toes Mosquitoes alone are thought to account for

half of all human deaths throughout history

Humans, the ultimate biological invaders, have

been responsible for the extinction of many species

and will continue to be in the future Like other

ani-mal invaders, humans tend to have a broad diet

Hu-mans are also able to adapt culturally to diverse

habi-tats, an ability that complements an ability to breed all

year round These attributes give humans a distinct

advantage over less aggressive and less destructive

spe-cies

Randall L Milstein

Further Reading

Burdick, Alan Out of Eden: An Odyssey of Ecological

Inva-sion New York: Farrar, Straus and Giroux, 2005.

Cartwright, Frederick F., and Michael Biddiss Disease

and History 2d ed Stroud, England: Sutton, 2000.

Crosby, Alfred W Ecological Imperialism: The Biological

Expansion of Europe, 900-1900 2d ed New York:

Cambridge University Press, 2004

Elton, Charles S The Ecology of Invasions by Animals and

Plants London: Methuen, 1958 Reprint Chicago:

University of Chicago Press, 2000

Hengeveld, Rob Dynamics of Biological Invasions New

York: Chapman and Hall, 1989

Lockwood, Julie L., Martha F Hoopes, and Michael P

Marchetti Invasion Ecology Malden, Mass.:

Black-well, 2007

Mooney, Harold A., and James A Drake, eds Ecology of Biological Invasions of North America and Hawaii New

York: Springer, 1986

Mooney, Harold A., and Richard J Hobbs, eds Inva-sive Species in a Changing World Washington, D.C.:

Island Press, 2000

Nentwig, Wolfgang, ed Biological Invasions New York:

Springer, 2007

Pimentel, David, ed Biological Invasions: Economic and Environmental Costs of Alien Plant, Animal, and Mi-crobe Species Boca Raton, Fla.: CRC Press, 2002.

Web Site University of Tennessee, Department of Ecology and Environmental Biology Institute for Biological Invasions

http://invasions.bio.utk.edu See also: Genetic diversity; Pesticides and pest con-trol; Species loss

Biomes

Category: Ecological resources

Biomes (terrestrial and aquatic ecosystems) are distrib-uted throughout the Earth’s surface Terrestrial biomes occupy the landmass from North Pole to South Pole Aquatic biomes occupy the bodies of water on Earth.

Background Biomes are natural habitats for bacteria, protists, fungi, plants, and animals Biomes maintain the natu-ral life cycle of these organisms and preserve the products of geological processes on Earth A biome is

a source of shelter, rocks and minerals, and food and fiber for human needs

Technical Definition

A terrestrial biome is a large ecosystem characterized

by a particular type of climate and soils with defined groups of highly adapted living organisms Biome for-mation is influenced by warm temperature and heavy precipitation in the tropics and extreme cold and low precipitation near the poles Most ecologists do not

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consider aquatic ecosystems as biomes and refer to

them as “aquatic biomes,” which are classified based

on the concentration of dissolved salts: less than 0.1

percent in freshwater biomes, 0.1 to 1.0 percent in

es-tuaries, and more than 1.0 percent in marine biomes

Climate and Biomes

Climate shapes terrestrial biomes Climate is

predom-inantly driven by the solar energy and atmospheric

circulation Air circulation is initiated at the equator,

because the equator receives the greatest solar energy

with the warmest air near the ground Because of

dif-ferent air densities, warm air in the troposphere rises

into the stratosphere and cools Cool air in the

strato-sphere descends into tropostrato-sphere and warms This

rise and fall pattern of circulating air starts at 0°

(equator) to 30° latitude, then continues at 30° to 60°

latitude, and ends at 60° to 90° latitude (poles)

There are six major atmospheric circulations:

Three move from the equator to the North Pole; the

other three move from the equator to the South Pole

At 0° latitude, the ascending warm, humid air from

the troposphere cools and condenses as it reaches the

stratosphere, releasing heavy rain to or near the

equa-tor That the dominant biomes formed at the equator

are the tropical rain forests is no accident After

re-leasing rain, the cool, dry air moves poleward and

de-scends at 30° latitude The descending cool, dry air

becomes warm as it reaches the troposphere and then

absorbs all the available moisture Not surprisingly,

the dominant biomes at 30° latitude are the deserts,

where the warm, humid air splits One air mass moves

equatorward to recirculate at 0° latitude The other

moves poleward and rises at 60° latitude, releasing rain

or snow while at the stratosphere As a result, the

dom-inant biomes at 60° latitude are the temperate forests

and temperate grasslands The cool, dry air at the

stratosphere divides again 60° latitude One air mass

moves toward 30° latitude to descend and recirculate

in the desert The other moves poleward, then

de-scends and releases the remaining moisture near the

poles, where the arctic tundra biomes are formed

Terrestrial Biomes

The are nine major terrestrial biomes

Arctic Tundra Arctic tundra is located in the

Northern Hemisphere near the North Pole and

cov-ers 20 percent of Earth’s landmass It has extremely

long, freezing, and harsh winters, with very short

(six-to eight-week) summers It is considered “cold desert,”

because it receives 20 centimeters of precipitation per year Melting snow creates bogs in summer, but there are frozen layers of subsoil (permafrost) at least a meter deep that exist throughout the year Soil is nutrient-poor Only the low-growing grasses and dwarf woody shrubs adapted to extreme cold and a short growing season are found No trees survive Their roots cannot penetrate the permafrost Few ani-mal species live in tundra In winters, ptarmigans, musk oxen, snowy owls, lynxes, arctic foxes, and snow-shoe hares are found Polar bears are common in the coastal regions In summers, few migrating animals from taiga move to tundra No reptiles are found, but mosquitoes survive

Taiga Taiga, also called boreal coniferous forest, exists south of tundra and covers 11 percent of the Earth’s land surface It is found in the northern parts

of North America and Eurasia and along the Pacific coast of northern North America to Northern Califor-nia It has patchy and shallower permafrost than tun-dra, and has acidic, nutrient-poor soil It has short summers and long, cold winters and receives 50 centi-meters precipitation per year Evergreen conifers are adapted to these conditions, with low-lying mosses and lichens beneath the forest canopy Seeds of coni-fers attract birds Bears, deer, moose, beavers, musk-rats, wolves, mountain lions, and wolverines inhabit the taiga

Temperate Rain Forest Temperate rain forest, a coniferous forest, stretches along the west coast of Canada and the United States, the southeast of Aus-tralia, and the south of South America It has dense fog, mild winters, cool summers, and high annual pre-cipitation of 250 centimeters With abundant rain and nutrient-rich soil, the temperate rain forests have re-tained some of the tallest conifers (such as coastal red-woods) and oldest trees, some as old as eight hundred years Moisture-loving plants (mosses and ferns) grow

on the tree trunks of evergreen conifers Temperate rain forest is a habitat for squirrels, lynxes, and several species of amphibians, reptiles, and birds (such as the spotted owl)

Temperate Deciduous Forest Temperate decid-uous forest is located south of the taiga in eastern North America, eastern Asia, and much of Europe Temperate deciduous forests have a moderate cli-mate, with occasional hot summers and cold winters and high annual precipitation of 75 to 150 centime-ters They have long growing seasons ranging from

140 to 300 days The soil is rich in minerals The

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nant trees are deciduous (oak, beech, sycamore, and

maple), which shed their broad leaves in the fall and

grow them in the spring Under the forest’s canopy,

understory trees and shrubs are found Layers of

growth in the forest are home for several insects and

birds Ground animals include rabbits, squirrels,

wood-chucks, chipmunks, turkeys, beavers, and muskrats

Temperate Grasslands Temperate grasslands

in-clude the South American pampas, the Russian

steppes, and the North American prairies Tall-grass

prairies are found between Illinois and Indiana,

whereas short-grass prairies extend from Texas to

Montana and North Dakota They have hot and dry

summers and bitterly cold winters, with annual

pre-cipitation of 25 to 75 centimeters Grasses in these

biomes produce a deep, dark, mineral-rich soil

Her-bivore mammals (bison, pronghorn antelope, mice,

prairie dogs, and rabbits) dominate the temperate

grasslands Hawks, snakes, badgers, coyotes, and foxes

are the predators in this biome

Shrubland Shrubland, or chaparral, is composed

of thickets of small-leaf evergreen shrubs (shorter

than trees and without main trunks) Shrublands,

with frequent fires in dry summers and winters of 25

to 75 centimeters of rain annually, are found along

the cape of South Africa, the western coast of North

America, the southwestern and southern shores of

Australia, around the Mediterranean Sea, and in

cen-tral Chile The shrubland in California is called

chap-arral, because it lacks understory Shrubs are

fire-adapted and highly flammable The seeds of many

species require the scarring action of fire to induce

germination Other shrubs resprout from the roots

af-ter fire Mule deer, rodents, scrub jays, and lizards

in-habit the shrublands

Deserts Deserts exist near or at 30° north and

south latitudes and cover approximately 30 percent of

the Earth’s land surface The dry air that descends in

this region absorbs most of the available moisture,

then moves away to the equator and to 60° latitude

Deserts receive less than 25 centimeters of rain

annu-ally The Sahara Desert of Africa and the Arabian

Peninsula and the deserts of North America (Mojave,

Chihuahuan, and Sonoran) have little or no

vegeta-tion Organisms with specialized water-conserving

adaptations survive, including cactus, agave, Joshua

trees, and sagebrush plants Hawks prey on lizards,

snakes, roadrunners, and kangaroo rats

Tropical Grasslands Tropical grasslands, or

sa-vannas (such as African sasa-vannas), characterized by

widespread growth of grasses with few interspersed trees, are found in areas with seasonal low rainfall and prolonged dry periods Other savannas occur in South America and northern Australia Savanna has

an annual precipitation of 25 to 75 centimeters Sa-vanna soil is nutrient-poor Acacia trees survive the se-vere dry season Hoofed herbivore mammals (giraffes, elephants, zebras, and rhinoceroses) feed on tree veg-etation and on grasses Carnivores such as hyenas, lions, cheetahs, and leopards prey on herbivores Tropical Rain Forests Tropical rain forests are located in South America, Africa, Southeast Asia, and the Indo-Malayan region on or near the equator Wet and dry seasons are warm year-round Annual rainfall

is 200 to 450 centimeters Tropical rain forest soil is typically nutrient-poor, but plentiful rain supports the growth of diverse groups of woody and herbaceous plants Some of the rains come from recycled water re-leased by forest trees by transpiration Of all the biomes, tropical rain forest is the richest, based on species diversity, productivity, and abundance of all organisms Tropical rain forest has three levels: the canopy (the highest layer of the forest), the under-story (middle layers of small trees and shrubs), and forest floor (ground layers of herbaceous plants) Epiphyte plants (such as bromeliads, orchids, ferns, and Spanish moss) gain access to sunlight by growing

on trunks and branches of tall trees Lemurs, sloths, and monkeys are tree-dwelling primates that feed on fruits The largest carnivores in the tropical rain forest are the jaguars in South America and the leopards in Africa and Asia

Aquatic Biomes All aquatic biomes share three ecological groups of organisms: the plankton, nekton, and benthos Plank-ton are classified into microscopic phytoplankPlank-ton and large zooplankton Phytoplankton are producers and include photosynthetic cyanobacteria and free-floating algae, which provide oxygen and food for heterotrophic organisms Zooplankton are consum-ers, heterotrophic, nonphotosynthetic organisms that include protozoa, small crustaceans, and larvae of aquatic animals Nekton are larger swimming ani-mals such as turtles, fishes, and whales Benthos are bottom-dwelling animals that attach themselves to

a substratum (sponges, oysters, and barnacles), bur-row themselves into soil (clams, worms, and echi-noderms) or simply swim or walk on the bottom (cray-fish, crabs, lobsters, insect larvae, and brittle stars)

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Based on salt contents, the three major aquatic

eco-systems are the freshwater, estuary, and marine

ecosys-tems Freshwater ecosystems, which contain less than

0.1 percent dissolved salts and occupy about 2 percent

of the Earth’s surface, include flowing waters (streams

and rivers), standing waters (ponds and lakes), and

freshwater wetlands (marshes and swamps) While all

freshwater habitats provide homes for animal species,

greater vegetations are found in marshes (grasslike

plants) and in swamps (trees and shrubs) than in

flow-ing- and standflow-ing-water ecosystems Estuaries occur

where fresh water and salt water meet, with salt

con-centrations of 0.1 to 1.0 percent Temperate estuaries

called salt marshes are dominated by salt-tolerant

grasses Tropical estuaries are called mangrove

for-ests Marine ecosystems, which contain more than

1.0 percent dissolved salts, dominate, occupying about

70 percent of the Earth’s surface Marine biomes have

three zones: the intertidal, pelagic, and benthic zones

The intertidal zone is the shoreline area between low

and high tide The pelagic zone is the ocean water

(shallow or deep), where plankton and swimming

ma-rine organisms are found The benthic zone is the

ocean floor, where marine animals burrow Coral

reefs, kelp forests, and seagrass beds are part of the

benthic zone

History

The existence of aquatic and terrestrial ecosystems

was discovered through fossil records Aquatic biomes

emerged before the terrestrial biomes Approximately

542 million years ago, during the Cambrian period,

organisms in marine biomes became diversified and

included bacteria, cyanobacteria, algae, fungi,

ma-rine invertebrates, and first chordates The first

ter-restrial biome existed when the first forest and

gym-nosperm appeared about 416 million years ago, during

the Denovian period About 359 million years ago,

during the Carboniferous period, the formation of

much more diversified forest occurred, which

con-sisted of ferns, clubmosses, horsetails, and

gymno-sperms and which housed many insects, amphibians,

and first reptiles Flowering plants (angiosperms) later

evolved and became the dominant organisms of most

major biomes

Domingo M Jariel

Further Reading

Kirchner, Renee Biomes Detroit: KidHaven Press/

Thomson Gale, 2006

Roth, Richard A Freshwater Aquatic Biomes Westport,

Conn.: Greenwood Press, 2009

Solomon, Eldra Pearl, Linda R Berg, and Diana W

Martin “Ecology and the Geography of Life.” In Bi-ology 8th ed Monterey, Calif.: Brooks/Cole, 2008 Woodward, Susan L Marine Biomes Westport, Conn.:

Greenwood Press, 2008

Web Site University of California Museum of Paleontology

http://www.ucmp.berkeley.edu/exhibits/biomes/ index.php

See also: Biodiversity; Biosphere; Biosphere reserves

Biopyriboles

Category: Mineral and other nonliving resources

Biopyriboles are minerals composed of linked silicate groups Some hard biopyriboles are used as gemstones Fibrous biopyriboles are used to manufacture asbestos Micas are used in electrical components and as fillers, absorbents, and lubricants Clays are used in bricks, pottery, and fillers.

Definition Biopyriboles are a large and varied group of minerals

in which silicate groups (one silicon atom bonded

to four oxygen atoms) are linked together in one-dimensional chains (either single chains or two chains linked together) or two-dimensional sheets Those with chains are usually hard, while those with sheets are usually soft Hard biopyriboles are usually found as separate minerals within igneous and meta-morphic rocks Soft biopyriboles are usually found as flakes of mica within rocks or as particles of clay in soils and freshwater sediments

Overview There are three broad categories of biopyriboles, depending on whether the silicate groups are linked together into single chains, double chains, or sheets Single-chain biopyriboles are known as pyroxenes Double-chain biopyriboles are known as amphiboles Together these two subgroups are known as pyriboles

or inosilicates Sheet biopyriboles are known as

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phyllosilicates The word “biopyribole”

is a combination of “biotite” (a common

phyllosilicate), “pyroxene,” and

“amphi-bole.”

Pyroxenes are composed of chains of

silicate groups combined with a wide

va-riety of other atoms, including sodium,

magnesium, calcium, iron, and

alumi-num They are generally fairly hard

min-erals with a density between three and

four grams per cubic centimeter

Pyrox-enes are usually dark green or black,

but other colors also exist The most

common pyroxene is augite, a green or

black mineral sometimes used as a

gem-stone Spodumene is a white, light gray,

or light yellow pyroxene that contains

lithium It is the most important source

of that element Jadeite, a type of jade, is a green

py-roxene used as a gemstone

Amphiboles are composed of two linked chains of

silicate groups combined with the same variety of

at-oms as those found in pyroxenes They also contain

hydroxyl groups (one oxygen atom bonded to one

hy-drogen atom), which cause them to release water

when heated At high temperatures the double chains

break down into single chains to form pyroxenes Am-phiboles are fairly hard minerals with a density be-tween 2.9 and 3.6 grams per cubic centimeter The most common amphibole is hornblende, a dark green

or black mineral Nephrite, a green amphibole, is a form of jade

Phyllosilicates are composed of sheets of silicate groups combined with the same kinds of atoms as those

Augite is the most common pyroxene, a type of biopyribole (USGS)

Biopyribole Categories

Examples

Sheet silicates

Brittle mica group —

illite Serpentine Antigorite, chrysotile

asbestos

Chain silicates

Single silica tetrahedron

chains:

Monoclinic alkali

pyroxenes

Jadeite

Examples Monoclinic calcic

pyroxenes

Diopside, augite

Orthorhombic pyroxenes

Enstatite, hypersthene

Double silica tetrahedra chains:

Monoclinic alkali amphiboles

Glaucophane, riebeckite Monoclinic calcic

amphiboles

Tremolite, hornblende

Monoclinic magnesium-iron amphiboles

Cummingtonite

Orthorhobmic amphiboles

Anthophyllite

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found in pyriboles Most phyllosilicates are soft

miner-als with a density between 2 and 3 grams per cubic

cen-timeter Talc, a light-colored, very soft phyllosilicate, is

used in paint, ceramics, and talcum powder

Serpen-tine, a green, fibrous mineral, is used to make asbestos

Many phyllosilicates exist as clays, used in ceramics

and fillers, or as micas, used in electrical components

Rose Secrest

See also: Asbestos; Clays; Gems; Mica; Silicates; Talc

Biosphere

Category: Ecological resources

The biosphere is the relatively thin layer around the

Earth’s surface where life is naturally possible The

concept is important in ecology for the calculation of

energy and mineral resource budgets, in space

explora-tion for the establishment and maintenance of livable

environments for space travelers, and perhaps for

un-derstanding the possibilities for life on other planets.

Background

The first use of the term “biosphere” dates to 1875,

when geologist Eduard Suess described layers of the

Earth in his book on the origin of the Alps The

Rus-sian geologist Vladimir Vernadsky popularized the

term in his lectures, published in French in 1929 as La

Biosphere Vernadsky noted that the concept, although

not the term, had originated much earlier with the

French biologist Jean-Baptiste Lamarck (1744-1829)

Extent of the Biosphere

Although most people would think nothing of

travel-ing 50 kilometers to a nearby town, journeytravel-ing

up-ward far less than this distance would mean certain

death without a special support system As altitude

in-creases, decreases in pressure, vital gases, and

temper-ature prevent active metabolism However, dormant

bacterial and fungal spores can apparently drift

up-ward indefinitely in this “parabiosphere.” Most jet

plane passengers are aware that artificial cabin

pres-sure is required to sustain them in the thinning

atmo-sphere when they are only a few kilometers high

Chlorophyll plants cannot live above about 6,200

me-ters because all water freezes at that altitude and the

carbon dioxide available for photosynthesis is at less

than half that available at sea level The few spiders and springtails that live on top of Mount Everest sur-vive on plant and animal debris blown up there by wind currents

Life also extends downward into the deepest ocean trenches, although the density of organisms is drasti-cally less in the dark zones beneath the thin top layer, where sunlight feeds algae and the resultant food chains Most deep-ocean organisms must feed on the rain of organic matter that sinks from the surface or feed in the detritus food chain Many organisms live

on the surface of the ocean bottom, and sampling studies have shown that life extends deep into these bottom muds Not all organisms here derive their en-ergy indirectly from plant photosynthesis; some thrive

on food chains originating with sulfur bacteria Or-ganisms that have evolved to live under the tremen-dous water pressure of the lower oceans burst open

if pulled to the surface; conversely, humans would

be crushed at these depths, so exploration requires heavy protective equipment Like deep-ocean fish brought to the surface, humans decompress when ex-posed at high altitudes Therefore, much of the bio-sphere is beyond humans’ day-to-day reach

Biomes The terrestrial part of the biosphere can be subdi-vided into such categories as hot and wet tropical rain forests, frozen arctic tundra, cold mountaintop mead-ows, and prairie grasslands These natural communi-ties with similar plants and animals are called “bi-omes.” For example, conifer forests stretch around the upper latitudes of Canada, Europe, and Russia Al-though the species of conifer trees, large grazers, and predators differ, the ecology is very similar The same

is true for the grassland biome that occurs in the U.S plains states, Russia, Argentina, and South Africa, and the temperate deciduous forests of the eastern United States, Europe, and China Other biomes in-clude taiga, savanna, thornbush, chaparral, and vari-ous tropical rain forest types

The first breakdown of biotic communities was made by C Hart Merriam, working in 1890 in Califor-nia and Arizona; his “vegetative life zones” were based

on temperature and ignored rainfall Victor E Shel-ford added detailed descriptions of animal associa-tions but did not try to correlate communities with cli-mate While Shelford’s followers consider biomes to

be distinct entities, other ecologists view them as hu-man concepts that hide the fact that communities

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