Bioms

Bioms
Contents
Articles
Biome 1
Tropical and subtropical moist broadleaf forests 10
Tropical and subtropical dry broadleaf forests 11
Tropical and subtropical coniferous forests 13
Temperate broadleaf and mixed forest 14
Temperate coniferous forest 18
Taiga 20
Tropical and subtropical grasslands, savannas, and shrublands 30
Temperate grasslands, savannas, and shrublands 33
Flooded grasslands and savannas 36
Montane grasslands and shrublands 36
Tundra 39
Mediterranean forests, woodlands, and scrub 43
Sclerophyll 46
Deserts and xeric shrublands 48
Mangrove 52
River delta 64
Upland and lowland (freshwater ecology) 69
Wetland 70
Endorheic basin 76
Palearctic ecozone 82
Cold seep 90
Benthic zone 104
Kelp forest 105
Intertidal zone 113
References
Article Sources and Contributors 117
Image Sources, Licenses and Contributors 121
Article Licenses
License 124
Biome 1
Biome
The planet Earth.
Biomes are climatically and geographically defined as similar
climatic conditions on the Earth, such as communities of plants,
animals, and soil organisms,[1] and are often referred to as
ecosystems. Some parts of the earth have more or less the same
kind of abiotic and biotic factors spread over a large area creating
a typical ecosystem over that area. Such major ecosystems are
termed as biomes. Biomes are defined by factors such as plant
structures (such as trees, shrubs, and grasses), leaf types (such as
broadleaf and needleleaf), plant spacing (forest, woodland,
savanna), and climate. Unlike ecozones, biomes are not defined by
genetic, taxonomic, or historical similarities. Biomes are often
identified with particular patterns of ecological succession and
climax vegetation (quasi-equilibrium state of the local ecosystem).
An ecosystem has many biotopes and a biome is a major habitat
type. A major habitat type, however, is a compromise, as it has an
intrinsic inhomogeneity.
The biodiversity characteristic of each biome, especially the diversity of fauna and subdominant plant forms, is a
function of abiotic factors and the biomass productivity of the dominant vegetation. In terrestrial biomes, species
diversity tends to correlate positively with net primary productivity, moisture availability, and temperature.[2]
Ecoregions are grouped into both biomes and ecozones.
A fundamental classification of biomes is:
1. Terrestrial (land) biomes
2. Aquatic biomes (including Freshwater biomes and Marine biomes)
Biomes are often known in English by local names. For example, a temperate grassland or shrubland biome is
known commonly as steppe in central Asia, prairie in North America, and pampas in South America.
Tropical
grasslands are known as savanna in Australia, whereas in Southern Africa it is known as certain kinds of
veld (from
Afrikaans).
Sometimes an entire biome may be targeted for protection, especially under an individual nation's Biodiversity
Action Plan.
Climate is a major factor determining the distribution of terrestrial biomes. Among the important climatic factors
are:
• latitude: Arctic, boreal, temperate, subtropical, tropical.
• humidity: humid, semi-humid, semi-arid, and arid.
• seasonal variation: Rainfall may be distributed evenly throughout the year or be marked by seasonal variations.
• dry summer, wet winter: Most regions of the earth receive most of their rainfall during the summer months;
Mediterranean climate regions receive their rainfall during the winter months.
• elevation: Increasing elevation causes a distribution of habitat types similar to that of increasing latitude.
The most widely used systems of classifying biomes correspond to latitude (or temperature zoning) and
humidity.
Biodiversity generally increases away from the poles towards the equator and increases with humidity.
Biome 2
Biome Classification Schemes
Biome classification schemes seek to define biomes using climatic measurements. Particularly in the 1970s and
1980s there was a significant push to understand the relationships between these measurements and properties of
ecosystem energetics because such discoveries would enable the prediction of rates of energy capture and
transfer
among components within ecosystems. Such a study was conducted by Sims et al. (1978) on North American
grasslands. The study found a positive logistic correlation between evapotranspiration in mm/yr and above
ground
net primary production in g/m^2/yr. More general results from the study were that precipitation and water use
lead to
aboveground primary production, solar radiation and temperature lead to belowground primary production
(roots),
and temperature and water lead to cool and warm season growth habit.[3] These findings help explain the
categories
used in Holdridge’s bioclassification scheme, which were then later simplified in Whittaker’s. The number of
classification schemes and the variety of determinants used in those schemes, however, should be taken as a
strong
indicator that biomes do not all fit perfectly into the classification schemes created.
Holdridge Scheme
The Holdridge classification scheme was developed by L. R. Holdridge, a botanist. It maps climates based on
four
categories:
• Average total precipitation (cm) on a logarithmic scale
• Potential evapotranspiration ratio: the potential evapotranspiration divided by the precipitation; the ratio
increases
from humid to arid regions.
• Potential evapotranspiration
• Mean annual biotemperature (°C): calculated from monthly mean temperatures after converting any mean
temperature to 0°C, based on the assumption that temperatures at or below freezing all have the same effect on
plants, and delineating between -10°C and -30°C would yield unrealistic results.
In this scheme, climates are classified based on the biological effects of temperature and rainfall on vegetation
under
the assumption that these two abiotic factors are the largest determinants of the type of vegetation found in an
area.
Holdridge uses the 4 axis to define 30 so called "humidity provinces," which are clearly visible in the Holdridge
diagram. While the scheme largely ignores soil and sun exposure, Holdridge did acknowledge that these, too,
were
important factors in biome determination.
Whittaker's Biome-type Classification Scheme
Whittaker appreciated biome-types as a representation of the great diversity of the living world, and saw the
need to
establish a simple way to classify these biome-types. Whittaker based his classification scheme on two abiotic
factors: Precipitation and Temperature. His scheme can be seen as a simplification of Holdridge's, one more
readily
accessible, but perhaps missing the greater specificity that Holdrige's provides.
Whittaker based his representation of global biomes on both previous theoretical assertions as well as an ever
increasing empirical sampling of global ecosystems. Whittaker was in a unique position to make such a holistic
assertion as he had previously compiled a review of biome classification.[4]
The Whittaker Classification Scheme can be viewed at the following address: here [5]
Biome 3
Key definitions for understanding Whittaker's Scheme
• physiognomy: The apparent characteristics, outward features, or appearance of ecological communities or
species.
• biome: a grouping terrestrial ecosystems on a given continent that are similar in vegetation structure,
physiognomy, features of the environment and characteristics of their animal communities
• formation: a major kind of community of plants on a given continent
• biome-type: grouping of convergent biomes or formations of different continents; defined by physiognomy
• formation-type: grouping of convergent formations
Whittaker's distinction between biome and formation can be simplified: formation is used when applied to plant
communities only, while biome is used when concerned with both plants and animals. Whittaker's convention of
biome-type or formation-type is simply a broader method to categorize similar communities.[6] The world
biome-types, as displayed on a world map, can be viewed at the following link: here [7]
Whittaker's parameters for classifying biome-types
Whittaker, seeing the need for a simpler way to express the relationship of community structure to the
environment,
used what he called “gradient analysis” of ecocline patterns to relate communities to climate on a worldwide
scale.
Whittaker considered four main ecoclines in the terrestrial realm.[8]
1. Intertidal levels: The wetness gradient of areas that are exposed to alternating water and dryness with
intensities
that vary by location from high to low tide
2. climatic moisture gradient
3. temperature gradient by altitude
4. temperature gradient by latitude
Along these gradients, Whittaker noted several trends that allow him to qualitatively establish biome-types.
• The gradient runs from favorable to extreme with corresponding changes in productivity.
• Changes in physiognomic complexity vary with the favorability of the environment (decreasing community
structure and reduction of stratal differentiation as the environment becomes less favorable).
• Trends in diversity of structure follow trends in species diversity; alpha and beta species diversities decrease
from
favorable to extreme environments.
• Each growth-form (i.e. grasses, shrubs, etc.) has its characteristic place of maximum importance along the
ecoclines.
• The same growth forms may be dominant in similar environments in widely different parts of the world.
Whittaker summed the effects of gradients (3) and (4), to get an overall temperature gradient and combined this
with
gradient (2), the moisture gradient, to express the above conclusions in what is known as the Whittaker
Classification
Scheme. The scheme graphs average annual precipitation (x-axis) versus average annual temperature (y-axis) to
classify biome-types.
Walter System
The Heinrich Walter classification scheme was developed by Heinrich Walter, a German ecologist. It differs
from
both the Whittaker and Holdridge schemes because it takes into account the seasonality of temperature and
precipitation. The system, also based on precipitation and temperature, finds 9 major biomes, with the important
climate traits and vegetation types summarized in the accompanying table. The boundaries of each biome
correlate to
the conditions of moisture and cold stress that are strong determinants of plant form, and therefore the vegetation
that
defines the region.
• I: Equatorial
• Always moist and lacking temperature seasonality
Biome 4
• Evergreen tropical rain forest
• II: Tropical
• Summer rainy season and cooler “winter” dry season
• Seasonal forest, scrub, or savanna
• III: Subtropical
• Highly seasonal, arid climate
• Desert vegetation with considerable exposed surface
• IV: Mediterranean
• Winter rainy season and summer drought
• Sclerophyllous (drought-adapted), frost-sensitive shrublands and woodlands
• V: Warm temperate
• Occasional frost, often with summer rainfall maximum
• Temperate evergreen forest, somewhat frost-sensitive
• VI: Nemoral
• Moderate climate with winter freezing
• Frost-resistant, deciduous, temperate forest
• VII: Continental
• Arid, with warm or hot summers and cold winters
• Grasslands and temperate deserts
• VIII: Boreal
• Cold temperate with cool summers and long winters
• Evergreen, frost-hardy needle-leaved forest (taiga)
• IX: Polar
• Very short, cool summers and long, very cold winters
• Low, evergreen vegetation, without trees, growing over permanently frozen soils
Bailey System
Robert G. Bailey almost developed a biogeographical classification system for the United States in a map
published
in 1976. Bailey subsequently expanded the system to include the rest of South America in 1981 and the world in
1989. The Bailey system is based on climate and is divided into seven domains (Polar, Humid Temperate, Dry,
Human, and Humid Tropical), with further divisions based on other climate characteristics (subarctic, warm
temperate, hot temperate, and subtropical; marine and continental; lowland and mountain).[9]
• 100 Polar Domain
• 120 Tundra Division
• M120 Tundra Division - Mountain Provinces
• 130 Subarctic Division
• M130 Subarctic Division - Mountain Provinces
• 200 Humid Temperate Domain
• 210 Warm Continental Division
• M210 Warm Continental Division - Mountain Provinces
• 220 Hot Continental Division
• M220 Hot Continental Division - Mountain Provinces
• 230 Subtropical Division
• M230 Subtropical Division - Mountain Provinces
• 240 Marine Division
Biome 5
• M240 Marine Division - Mountain Provinces
• 250 Prairie Division
• 260 Mediterranean Division
• M260 Mediterranean Division - Mountain Provinces
• 300 Dry Domain
• 310 Tropical/Subtropical Steppe Division
• M310 Tropical/Subtropical Steppe Division - Mountain Provinces
WWF system
A team of biologists convened by the World Wide Fund for Nature (WWF) developed an ecological land
classification system that identified fourteen biomes,[10] called major habitat types, and further divided
the world's
land area into 867 terrestrial ecoregions. Each terrestrial Ecoregion has a specific EcoID, fomat XXnnNN (XX is
the
Ecozone, nn is the Biome number, NN is the individual number). This classification is used to define the Global
200
list of ecoregions identified by the WWF as priorities for conservation. The WWF major habitat types are:
• 01 Tropical and subtropical moist broadleaf forests (tropical and subtropical, humid)
• 02 Tropical and subtropical dry broadleaf forests (tropical and subtropical, semi-humid)
• 03 Tropical and subtropical coniferous forests (tropical and subtropical, semi-humid)
• 04 Temperate broadleaf and mixed forests (temperate, humid)
• 05 Temperate coniferous forests (temperate, humid to semi-humid)
• 06 Boreal forests/taiga (subarctic, humid)
• 07 Tropical and subtropical grasslands, savannas, and shrublands (tropical and subtropical, semi-arid)
• 08 Temperate grasslands, savannas, and shrublands (temperate, semi-arid)
• 09 Flooded grasslands and savannas (temperate to tropical, fresh or brackish water inundated)
• 10 Montane grasslands and shrublands (alpine or montane climate)
• 11 Tundra (Arctic)
• 12 Mediterranean forests, woodlands, and scrub or Sclerophyll forests (temperate warm, semi-humid to semi-
arid
with winter rainfall)
• 13 Deserts and xeric shrublands (temperate to tropical, arid)
• 14 Mangrove (subtropical and tropical, salt water inundated)
Freshwater biomes
According to the World Wildlife Fund, the following are classified as freshwater biomes:[11]
• Large lakes • Temperate upland rivers
• Large river deltas • Tropical and subtropical coastal rivers
• Polar freshwaters • Tropical and subtropical floodplain rivers and wetlands
• Montane freshwaters • Tropical and subtropical upland rivers
• Temperate coastal rivers • Xeric freshwaters and endorheic basins
• Temperate floodplain rivers and wetlands • Oceanic islands
Biome 6
Realms or Ecozones (terrestrial and freshwater, WWF)
• NA Nearctic
• PA Palearctic
• AT Afrotropic
• IM Indomalaya
• AA Australasia
• NT Neotropic
• OC Oceania
• AN Antarctic
Marine biomes
Marine biomes (H) (major habitat types), Global 200 (WWF)
Biomes of the coastal & continental shelf areas (Neritic zone - List of ecoregions (WWF))
• Polar
• Temperate shelves and sea
• Temperate upwelling
• Tropical upwelling
• Tropical coral[12]
Realms or Ecozones (marine, WWF)
• North Temperate Atlantic • Western Indo-Pacific
• Eastern Tropical Atlantic • South Temperate Indo-Pacific
• Western Tropical Atlantic • Southern Ocean
• South Temperate Atlantic • Antarctic
• North Temperate Indo-Pacific • Arctic
• Central Indo-Pacific • Mediterranean
• Eastern Indo-Pacific
Other marine habitat types
• Hydrothermal vents
• Cold seeps
• Benthic zone
• Pelagic zone (trades and westerlies)
• Abyssal
• Hadal (ocean trench)
Major Habitats, Non Global 200 (WWF)
• Littoral/Intertidal zone
• Kelp forest
• Pack ice
Summary - Ecological taxonomy (WWF)
• Biosphere (List of ecoregions)
• Ecozones or Realms (8)
• Terrestrial Biomes (Major Habitat Types, 14)
• Ecoregions (867)
• Ecosystems (Biotopes)
• Freshwater Biomes (Major Habitat Types, 12)
Biome 7
• Marine Ecozones or Realms (13)
• Continental Shelf Biomes (Major Habitat Types, 5)
• (Marine Provinces) (62)
• Open & Deep Sea Biomes (Major Habitat Types)
• Endolithic Biome
Example
• Biosphere
• Ecozone: Palearctic ecozone
• Terrestrial Biome: Temperate Broadleaf and Mixed Forests
• Ecoregion: Dinaric Mountains mixed forests (PA0418)
• Ecosystem: Orjen, vegetation belt between 1,100- 1,450 m, Oromediterranean zone, Nemoral zone
(temperate zone)
• Biotope: Oreoherzogio-Abietetum illyricae Fuk. (Plant list)
• Plant: Silver fir (Abies alba)
Anthropogenic biomes
Humans have fundamentally altered global patterns of biodiversity and ecosystem processes. As a result,
vegetation
forms predicted by conventional biome systems are rarely observed across most of Earth's land surface.
Anthropogenic biomes provide an alternative view of the terrestrial biosphere based on global patterns of
sustained
direct human interaction with ecosystems, including agriculture, human settlements, urbanization, forestry and
other
uses of land. Anthropogenic biomes offer a new way forward in ecology and conservation by recognizing the
irreversible coupling of human and ecological systems at global scales and moving us toward an understanding
how
best to live in and manage our biosphere and the anthropogenic biosphere we live in. The main biomes in the
world
are freshwater, marine, coniferous, deciduous, ice, mountains, boreal, grasslands, tundra, and rainforests.
Major Anthropogenic Biomes
• Dense Settlements
• Villages
• Croplands
• Rangelands
• Forested
Biome 8
Other biomes
The Endolithic biome, consisting entirely of microscopic life in rock pores and cracks, kilometers beneath the
surface, has only recently been discovered and does not fit well into most classification schemes.
Map of Biomes
Freshwater Biomes
Drainage basins of the principal oceans and seas of the world. Grey areas are endorheic basins that do not drain
to
the ocean.
Biome 9
References
[1] The World's Biomes (http:/ / www. ucmp. berkeley. edu/ exhibits/ biomes/ index. php), Retrieved August 19, 2008, from University of
California Museum of Paleontology (http:/ / www. ucmp. berkeley. edu/ index. php)
[2] Pidwirny, Michael (2006-10-16). "Biomes" (http:/ / www. eoearth. org/ article/ Biomes). In Sidney Draggan. Encyclopedia of Earth.
Washington, D.C.: Environmental Information Coalition, National Council for Science and the Environment. . Retrieved 2006-11-16.
[3] Pomeroy, Lawrence R. and James J. Alberts, editors. Concepts of Ecosystem Ecology. New York: Springer-Verlag, 1988.
[4] Whittaker, Robert H., Botanical Review, Classification of Natural Communities, Vol. 28, No. 1 (Jan-Mar 1962), pp. 1-239.
[5] http:/ / www. marietta. edu/ ~biol/ biomes/ images/ woodland/ whittaker_v2. jpg
[6] Whittaker, Robert H. Communities and Ecosystems New York: MacMillan Publishing Company, Inc., 1975.
[7] http:/ / www. marietta. edu/ ~biol/ biomes/ biomemap. html
[8] Whittaker, Robert H. Communities and Ecosystems New York: MacMillan Publishing Company, Inc., 1975.
[9] http:/ / www. fs. fed. us/ land/ ecosysmgmt/ index. html Bailey System, US Forest Service
[10] Olson, David M. et al. (2001); Terrestrial Ecoregions of the World (http:/ / www. csrc. sr. unh. edu/ ~palace/ cerrado/ olson et al. (world
veg
map). pdf): A New Map of Life on Earth, BioScience, Vol. 51, No. 11., pp. 933-938.
[11] "Freshwater Ecoregions of the World: Major Habitat Types" (http:/ / www. feow. org/ mht. php). Accessed May 12, 2008.
[12] WWF: Marine Ecoregions of the World (http:/ / www. worldwildlife. org/ science/ ecoregions/ marine/ item1266. html)
External links
• Biomes of the world (Missouri Botanic Garden) (http:/ / mbgnet. mobot. org)
• Global Currents and Terrestrial Biomes Map (http:/ / www. theglobaleducationproject. org/ earth/ global-
ecology.
php#3)
• WorldBiomes.com (http:/ / www. worldbiomes. com/ ) is a site covering the 5 principal world biome types:
aquatic, desert, forest, grasslands, and tundra.
• UWSP's online textbook The Physical Environment: - Earth Biomes (http:/ / www. uwsp. edu/ geo/
faculty/ ritter/
geog101/ textbook/ biomes/ outline. html)
• Panda.org's Habitats (http:/ / wwf. panda. org/ about_our_earth/ ecoregions/ about/ habitat_types/ ) - describes
the
14 major terrestrial habitats, 7 major freshwater habitats, and 5 major marine habitats.
• Panda.org's Habitats Simplified (http:/ / wwf. panda. org/ about_our_earth/ ecoregions/ about/ habitat_types/
habitats/ ) - provides simplified explanations for 10 major terrestrial and aquatic habitat types.
• UCMP Berkeley's The World's Biomes (http:/ / www. ucmp. berkeley. edu/ glossary/ gloss5/ biome/ ) -
provides
lists of characteristics for some biomes and measurements of climate statistics.
• Gale/Cengage has an excellent Biome Overview (http:/ / www. galeschools. com/ environment/ biomes/
overview. htm) of terrestrial, aquatic, and man-made biomes with a particular focus on trees native to each, and
has detailed descriptions of desert, rain forest, and wetland biomes.
• NASA's Earth Observatory Mission: Biomes (http:/ / earthobservatory. nasa. gov/ Experiments/ Biome/ ) gives
an
exemplar of each biome that is described in great detail and provides scientific measurements of the climate
statistics that define each biome.
Tropical and subtropical moist

broadleaf forests
Tropical and subtropical moist broadleaf forests (TSMF), also known as tropical moist
forests, are a tropical
and subtropical forest biome.
Tropical and subtropical forest regions with lower rainfall are home to tropical and subtropical dry broadleaf
forests
and tropical and subtropical coniferous forests. Temperate rain forests also occur in certain humid temperate
coastal
regions.
The biome includes several types of forests:
• Lowland equatorial evergreen rain forests, commonly known as tropical rainforests
or jungles, are forests
which receive high rainfall (more than 2000 mm, or 80 inches, annually) throughout the year. These forests
occur
in a belt around the equator, with the largest areas in the Amazon basin of South America, the Congo basin of
central Africa, Indonesia, and New Guinea.
• Moist deciduous and semi-evergreen seasonal forests, receive high overall rainfall with a
warm summer wet
season and a cooler winter dry season. Some trees in these forests drop some or all of their leaves during the
winter dry season. These forests are found in parts of South America, in Central America and around the
Caribbean, in coastal West Africa, parts of the Indian subcontinent, and across much of Indochina.
• Montane rain forests, some of which are known as cloud forests, are found in cooler-climate
mountainous areas.
• Flooded forests, including freshwater swamp forests and peat swamp forests.
Tropical and subtropical moist broadleaf forests are common in several terrestrial ecozones, including parts of
the
Afrotropic (equatorial Africa), Indomalaya (parts of the Indian subcontinent and Southeast Asia), the
Neotropic
(northern South America and Central America), Australasia (eastern Indonesia, New Guinea, northern and
eastern
Australia), and Oceania (the tropical islands of the Pacific Ocean). About half of the world's tropical
rainforests are
in the South American countries of Brazil and Peru. Rain forests now cover less than 6% of Earth's land surface.
Scientists estimate that more than half of all the world's plant and animal species live in tropical rain forests.
Tropical and subtropical moist forests within the Holdridge Life Zone
classification scheme.
External links
• Facts about the world's tropical rainforests [1] from The Nature Conservancy
• Worldwide Fund for Nature [2]
• Types of rainforests [3]
• Rainforest - Classroom activities [4]
• NASA picture of the afforest of the earth in the year 2002 [5]
• BBC video clips and details of the species found here [6]
References
[1] http:/ / www. nature. org/ rainforests/ explore/ facts. html
[2] http:/ / www. panda. org/ about_wwf/ where_we_work/ ecoregions/ about/ habitat_types/ selecting_terrestrial_ecoregions/ habitat01. cfm
[3] http:/ / rainforests. mongabay. com/ 0103. htm
[4] http:/ / www. phoenixzoo. org/ learn/ educator_resources/ rainforest_classroom_activities. pdf
[5] http:/ / www. spiegel. de/ img/ 0,1020,737991,00. jpg
[6] http:/ / www. bbc. co. uk/ nature/ habitats/ Tropical_and_subtropical_moist_broadleaf_forests
Tropical and subtropical dry

broadleaf forests
Trinidad and Tobago dry forest on Chacachacare showing the
dry-season deciduous nature of the vegetation
The tropical and subtropical dry broadleaf forest
biome, also known as tropical dry forest, is located at
tropical and subtropical latitudes. Though these forests
occur in climates that are warm year-round, and may
receive several hundred centimeters of rain per year,
they have long dry seasons which last several months
and vary with geographic location. These seasonal
droughts have great impact on all living things in the
forest.
Deciduous trees predominate in most of these forests,
and during the drought a leafless period occurs, which
varies with species type. Because trees lose moisture
through their leaves, the shedding of leaves allows trees
such as teak and mountain ebony to conserve water
during dry periods. The newly bare trees open up the canopy layer, enabling sunlight to reach ground level and
facilitate the growth of thick underbrush. Trees on moister sites and those with access to ground water tend to be
evergreen. Infertile sites also tend to support evergreen trees. Three tropical dry broadleaf forest ecoregions, the
East
Deccan dry evergreen forests, the Sri Lanka dry-zone dry evergreen forests, and the Southeastern Indochina dry
evergreen forests, are characterized by evergreen trees.
Though less biologically diverse than rainforests, tropical dry forests are home to a wide variety of wildlife
including
monkeys, deer, large cats, parrots, various rodents, and ground dwelling birds. Mammalian biomass tends to be
higher in dry forests than in rain forests, especially in Asian and African dry forests. Many of these species
display
extraordinary adaptations to the difficult climate.
This biome is alternately known as the tropical and subtropical dry forest biome or the tropical
and subtropical
deciduous forest biome. Locally some of these forests are also called monsoon forests, and they
tend to merge into
savannas.
Tropical and subtropical dry broadleaf forests 12
Geographical variation
Dry forests tend to exist north and south of the equator rain forest belt, south or north of the subtropical deserts,
generally in two bands, one between 10° and 20°N latitude and the other between 20° and 30°S latitude. The
most
diverse dry forests in the world occur in southern Mexico and in the Bolivian lowlands. The dry forests of the
Pacific
Coast of northwestern South America support a wealth of unique species due to their dry climate. The
subtropical
forests of Maputo land-Ponderousin the sub-tropical regions of the united states of America and in southeastern
Africa are diverse and support many endemic species. The dry forests of central India and Indochina are notable
for
their diverse large vertebrate faunas. Madagascar dry deciduous forests and New Caledonia dry forests are also
highly distinctive (pronounced extremism and a large number of reliquary taxa) for a wide range of taxa and at
higher taxonomic levels. Trees use underground water during the dry seasons.
Biodiversity patterns and requirements
Species tend to have wider ranges than moist forest species, although in some regions many species do display
highly restricted ranges; most dry forest species are restricted to tropical dry forests, particularly in plants; beta
diversity and alpha diversity high but typically lower than adjacent moist forests.
Effective conservation of dry broadleaf forests requires the preservation of large and continuous areas of forest.
Large natural areas are required to maintain larger predators and other vertebrates, and to buffer sensitive species
from hunting pressure. The persistence of riparian forests and water sources is critical for many dry forest
species.
Large swathes of intact forest are required to allow species to recover from occasional large events, like forest
fires.
Dry forests are highly sensitive to excessive burning and deforestation; overgrazing and exotic species can also
quickly alter natural communities; restoration is possible but challenging, particularly if degradation has been
intense
and persistent. Degrading dry broadleaf often leaves thorny shrublands, thickets, or dry grasslands in their place.
References
• 2006-01-13, Sciencedaily: Deep-rooted Plants Have Much Greater Impact On Climate Than Experts Thought
[1]
Citat: "...The tap roots transfer rainwater from the surface to reservoirs deep underground and redistribute
water...increases photosynthesis and the evaporation of water...by 40 percent in the dry season...During the wet
season, these plants can store as much as 10 percent of the annual precipitation as deep as 13 meters (43 feet)
underground, to be tapped during the dry months...tree roots acting like pipes to allow water to shift around
much
faster than it could otherwise percolate through the soil..."
Much material in this article has been reworked from Tropical and Subtropical Dry
Broadleaf Forests [2], by WWF
[3] with their permission.
External links
• The Tropical Dry Forest of Costa Rica [4]
References
[1] http:/ / www. sciencedaily. com/ releases/ 2006/ 01/ 060112035906. htm
[3] http:/ / www. panda. org
[4] http:/ / www. costarica21. com/ Tropical-Dry-Forest. html
Tropical and subtropical coniferous forests 13
Tropical and subtropical coniferous

forests
Tropical and subtropical coniferous forests are a forest biome. They are located in regions of
semi-humid climate
at tropical and subtropical latitudes. Most tropical and subtropical coniferous forest ecoregions are found in the
Nearctic and Neotropic ecozones, from Mexico to Nicaragua and on the Greater Antilles, Bahamas, and
Bermuda.
Other tropical and subtropical coniferous forests ecoregions occur in Asia.
Tropical and subtropical coniferous forests ecoregions
Indomalaya Tropical and subtropical coniferous forests
Himalayan subtropical pine forests Bhutan, India, Nepal, Pakistan
Luzon tropical pine forests Philippines
Northeast India-Myanmar pine forests Burma, India
Sumatran tropical pine forests Indonesia
Neotropic Tropical and subtropical coniferous forests
Bahamian pineyards The Bahamas
Belizian pine forests Belize
Central American pine-oak forests El Salvador, Guatemala, Honduras, Mexico, Nicaragua
Cuban pine forests Cuba
Hispaniolan pine forests Haiti, Dominican Republic
Miskito pine forests Honduras, Nicaragua
Sierra de la Laguna pine-oak forests Mexico
Sierra Madre de Oaxaca pine-oak forests Mexico
Sierra Madre del Sur pine-oak forests Mexico
Trans-Mexican Volcanic Belt pine-oak forests Mexico
Nearctic Tropical and subtropical coniferous forests
Bermuda subtropical conifer forests Bermuda
Sierra Madre Occidental pine-oak forests Mexico, United States
Sierra Madre Oriental pine-oak forests Mexico, United States
External links
Tropical and subtropical coniferous forests [1]
References
Temperate broadleaf and mixed

forest
Temperate mixed forest in Yunnan, southwest China.
Mixed forests are a temperate and humid biome. The
typical structure of these forests include four layers.
The upper most layer is the canopy which is composed
of tall mature trees ranging from 33 to 66 m (100 to
200 feet) high. Below the canopy is the three-layered,
shade tolerant understory which is roughly 9 to 15 m
(30 to 50 feet) shorter than the canopy. The top layer of
the understory is the sub-canopy which is composed of
smaller mature trees, saplings, and suppressed juvenile
canopy layer trees awaiting an opening in the canopy.
Below the sub-canopy is the shrub layer, composed of
low growing woody plants. Typically the lowest
growing (and most diverse) layer is the ground cover or
herbaceous layer.
Trees
Characteristic dominant broadleaf trees in this biome include oaks (Quercus spp.), beeches (Fagus spp.),
maples
(Acer spp.), and birches (Betula spp.).[1] The term "mixed forest" comes from the inclusion of coniferous trees
as a
canopy component of these forests. Typical coniferous trees include: Pines (Pinus spp.), firs (Abies spp.), and
spruces (Picea spp.). In some areas of this biome the conifers may be a more important canopy species than the
broadleaf species.
Climate
Temperate broadleaf and mixed forests occur in areas with distinct warm and cool season, which give it a
moderate
annual average temperature (3 to 15.6 °C). These forests occur in relatively warm and rainy climates, sometimes
also
with a distinct dry season. A dry season occurs in the winter in East Asia and in summer on the wet fringe of the
Mediterranean climate zones. Other areas have a fairly even distribution of rainfall; annual rainfall is typically
over
600 millimetres (24 inches) and often over 1500 millimetres (60 inches). Temperatures are typically moderate
except
in parts of Asia such as Ussuriland where temperate forests can occur despite very harsh conditions with very
cold
winters.
Temperate broadleaf and mixed forest ecoregions
Oceania
Oceania Temperate broadleaf and mixed forests
Chatham Islands temperate forests New Zealand
Eastern Australian temperate forests Australia
Fiordland temperate forests New Zealand
Nelson Coast temperate forests New Zealand
Northland temperate forests New Zealand
Northland temperate kauri forests New Zealand
Stewart Island/Rakiura temperate forests New Zealand
Richmond temperate forests New Zealand
Southeast Australia temperate forests Australia
Southland temperate forests New Zealand
Tasmanian Central Highland forests Australia
Tasmanian temperate forests Australia
Tasmanian temperate rain forests Australia
Westland temperate forests New Zealand
Eurasia
Indomalaya Temperate broadleaf and mixed forests
Eastern Himalayan broadleaf forests Bhutan, India, Nepal
Northern Triangle temperate forests Burma
Western Himalayan broadleaf forests India, Nepal, Pakistan
Palearctic Temperate broadleaf and mixed forests
Appenine deciduous montane forests Italy
Atlantic mixed forests Belgium, Denmark, France, Germany. Netherlands
Azores temperate mixed forests Portugal
Balkan mixed forests Bulgaria, Greece, Macedonia, Romania, Serbia (Kosovo), Turkey
Baltic mixed forests Denmark, Germany, Poland, Sweden
Cantabrian mixed forests Portugal, Spain
Caspian Hyrcanian mixed forests Azerbaijan, Iran
Caucasus mixed forests Armenia, Azerbaijan, Georgia, Russia, Turkey
Celtic broadleaf forests Ireland, United Kingdom
Central Anatolian deciduous forests Turkey
Central China loess plateau mixed
forests
China
Central European mixed forests Austria, Belarus, Czech Republic, Germany, Lithuania, Moldova, Poland
Central Korean deciduous forests North Korea, South Korea
Changbai Mountains mixed forests China, North Korea
Changjiang Plain evergreen forests China
Crimean Submediterranean forest
complex
Russia, Ukraine
Daba Mountains evergreen forests China
Dinaric Mountains mixed forests Albania, Bosnia and Herzegovina, Croatia, Italy, Montenegro, Serbia, Slovenia
East European forest steppe Bulgaria, Moldova, Romania, Russia, Ukraine
Eastern Anatolian deciduous forests Turkey
English Lowlands beech forests United Kingdom
Euxine-Colchic deciduous forests Georgia, Turkey
Hokkaido deciduous forests Japan
Huang He Plain mixed forests China
Madeira evergreen forests Portugal
Manchurian mixed forests China, North Korea, Russia, South Korea
Nihonkai evergreen forests Japan
Nihonkai montane deciduous forests Japan
North Atlantic moist mixed forests Ireland, United Kingdom
Northeast China Plain deciduous forests China
Pannonian mixed forests Austria, Bosnia and Herzegovina, Croatia, Czech Republic, Hungary, Romania, Serbia, Slovakia,
Slovenia, Ukraine
Po Basin mixed forests Italy
Pyrenees conifer and mixed forests Andorra, France, Spain
Qin Ling Mountains deciduous forests China
Rodope montane mixed forests Bulgaria, Greece, Macedonia, Serbia
Sarmatic mixed forests Belarus, Estonia, Finland, Latvia, Lithuania, Norway, Russia, Sweden
Sichuan Basin evergreen broadleaf
forests
China
South Sakhalin-Kurile mixed forests Russia
Southern Korea evergreen forests South Korea
Taiheiyo evergreen forests Japan
Taiheiyo montane deciduous forests Japan
Tarim Basin deciduous forests and
steppe
Xinjiang, China
Ussuri broadleaf and mixed forests Russia
West Siberian broadleaf and mixed
forests
Russia
Western European broadleaf forests Austria, Czech Republic, France, Germany, Switzerland
Zagros Mountains forest steppe Iran
Americas
Nearctic Temperate broadleaf and mixed forests
Allegheny Highlands forests United States
Appalachian mixed mesophytic forests United States
Appalachian-Blue Ridge forests United States
California mixed evergreen forest United States
Central U.S. hardwood forests United States
East Central Texas forests United States
Eastern forest-boreal transition Canada, United States
Eastern Great Lakes lowland forests Canada, United States
Gulf of St. Lawrence lowland forests Canada
Lac Saint-Jean and Saguenay valley forests Canada
Mississippi lowland forests United States
New England-Acadian forests Canada, United States
Northeastern coastal forests United States
Ozark Mountain forests United States
Southeastern mixed forests United States
Southern Great Lakes forests United States
Upper Midwest forest-savanna transition United States
Western Great Lakes forests Canada, United States
Willamette Valley forests United States
Neotropic Temperate broadleaf and mixed forests
Juan Fernandez Islands temperate forests Chile
Magellanic subpolar forests Argentina, Chile
Polylepis forests Bolivia, Peru
San Felix-San Ambrosio Islands temperate forests (Desventuradas Islands) Chile
Valdivian temperate rain forests Argentina, Chile
External links
• Temperate forest [2]
• Index of North American Temperate Broadleaf & Mixed Forests ecoregions at bioimages.vanderbilt.edu [3]
• Terraformers Canadian Forest Conservation Foundation [4]
References
[1] World Wildlife Fund: Temperate Broadleaf and Mixed Forest Ecoregions (http:/ / www. panda. org/ about_wwf/ where_we_work/
ecoregions/ about/ habitat_types/ selecting_terrestrial_ecoregions/ habitat04. cfm)
[2] http:/ / www. barrameda. com. ar/ ecology/ temperate-forest. htm
[3] http:/ / www. cas. vanderbilt. edu/ bioimages/ ecoframe-list. htm
[4] http:/ / forest. terraformers. ca/
Temperate coniferous forest

A temperate coniferous forest
ecosystem in Petroglyphs Provincial
Park, Ontario, Canada.
Temperate coniferous forest is a terrestrial biome found in temperate regions of
the world with warm summers and cool winters and adequate rainfall to sustain a
forest. In most temperate coniferous forests, evergreen conifers predominate,
while some are a mix of conifers and broadleaf evergreen trees and/or broadleaf
deciduous trees. Temperate evergreen forests are common in the coastal areas of
regions that have mild winters and heavy rainfall, or inland in drier climates or
mountain areas. Many species of trees inhabit these forests including cedar,
cypress, douglas-fir, fir, juniper, kauri, pine, podocarpus, spruce, redwood and
yew. The understory also contains a wide variety of herbaceous and shrub
species.
A Pine forest is an example of a temperate
coniferous forest
Structurally, these forests are rather simple, generally consisting of two
layers: an overstory and understory. Some forests may support an
intermediate layer of shrubs. Pine forests support an herbaceous
understory that is generally dominated by grasses and herbaceous
perennials, and are often subject to ecologically important wildfires.
Temperate coniferous forest 19
Carpathian montane conifer forest, Slovakia
Temperate rain forests occur only in seven regions around the world:
the Pacific temperate rain forests of the Pacific Northwest, the
Valdivian temperate rain forests of southwestern South America, the
rain forests of New Zealand and Tasmania, northwest Europe (small
pockets in Ireland, Scotland, Wales, Iceland and a somewhat larger
area in Norway), southern Japan, and the eastern Black Sea-Caspian
Sea region of Turkey and Georgia to northern Iran. The moist
conditions of temperate rain forests generally support an understory of
mosses, ferns and some shrubs. Temperate rain forests can be
Temperate coniferous forests or Temperate broadleaf and mixed
forests.
The temperate coniferous rain forests sustain the highest levels of biomass in any terrestrial ecosystem and are
notable for trees of massive proportions, including Giant Sequoia (Sequoiadendron gigantea), Coast
Redwood
(Sequoia sempervirens), Douglas-fir (Pseudotsuga menziesii), Sitka Spruce (Picea sitchensis),
Alerce (Fitzroya
cupressoides) and Kauri (Agathis australis). These forests are quite rare, occurring in small areas of
North America,
southwestern South America and northern New Zealand. The Klamath-Siskiyou forests of northwestern
California
and southwestern Oregon is known for its rich variety of plant and animal species, including many endemic
species.
External links
• Temperate forest [2]
• WWF - Temperate Coniferous Forest Ecoregions [1]
• Temperate Coniferous Forest of Northwest California [2]
References
[1] http:/ / www. panda. org/ about_our_earth/ ecoregions/ about/ habitat_types/ selecting_terrestrial_ecoregions/ habitat05. cfm
[2] http:/ / conifercountry. com/
Taiga 20
Taiga
The taiga is found throughout the high northern
latitudes, between the tundra, and the temperate
forest, from about 50°N to 70°N, but with
considerable regional variation.
Taiga (pronounced English pronunciation: /ˈtaɪɡə/, Russian: тайга́; from
Turkic[1] or Mongolian), also known as the boreal forest, is a biome
characterized by coniferous forests.
Taiga is the world's largest terrestrial biome and covers: in North
America most of inland Canada and Alaska as well as parts of the
extreme northern continental United States (especially northern
Minnesota, Michigan's Upper Peninsula, northern Wisconsin, Upstate
New York, Vermont, New Hampshire, and Maine); and in Eurasia
most of Sweden, Finland, inland and northern Norway, much of Russia
(especially Siberia), northern Kazakhstan, northern Mongolia, and
northern Japan (on the island of Hokkaidō).
The term boreal forest is sometimes, particularly in Canada, used to refer to the more southerly part of the
biome,
while the term taiga is often used to describe the more barren areas of the northernmost part of the taiga
approaching
the tree line.
White Spruce taiga, Denali Highway, Alaska
Range, Alaska
Climate and geography
Taiga is the world's largest land biome, and makes up 27% of the
world's forest cover;[2] the largest areas are located in Russia and
Canada. The taiga is the terrestrial biome with the lowest annual
average temperatures after the tundra and permanent ice caps.
However, extreme minimums in the taiga are typically lower than
those of the tundra. The lowest reliably recorded temperatures in the
Northern Hemisphere were recorded in the taiga of northeastern
Russia. The taiga or boreal forest has a subarctic climate with very
large temperature range between seasons, but the long and cold winter
is the dominant feature. This climate is classified as Dfc, Dwc, Dsc, Dfd, Dwd and Dsd in the Köppen
climate
classification scheme,[3] meaning that the short summer (24-hr average 10°C or more) lasts 1–3 months and
always
less than 4 months. There are also some much smaller areas grading towards the oceanic Cfc climate with
milder
winters. The mean annual temperature generally varies from -5°C to 5°C,[4] but there are taiga areas in both
eastern
Siberia and interior Alaska-Yukon where the mean annual reaches down to -10°C.[5] [6] According to some
sources,
the boreal forest grades into a temperate mixed forest when mean annual temperature reaches about 3 °C.[7]
Permafrost is common in areas with mean annual temperature below 0 °C. The winters last 5 – 7 months, with
average temperatures below freezing. Temperatures vary from −54 °C to 30 °C (-65 °F to 86 °F) throughout the
whole year.The summers, while short, are generally warm and humid. In much of the taiga, -20 °C would be a
typical winter day temperature and 18 °C an average summer day.
Taiga 21
The taiga in the river valley near Verkhoyansk,
Russia, at 67°N, must deal with the coldest winter
temperatures in the northern hemisphere, but the
extreme continentality of the climate gives an
average daily high of 22 °C in July.
The growing season, when the vegetation in the taiga comes alive, is
usually slightly longer than the climatic definition of summer as the
plants of the boreal biome have a lower threshold to trigger growth. In
Canada, Scandinavia and Finland, the growing season is often
estimated by using the period of the year when the 24-hr average
temperature is 5 °C or more.[8] For the Taiga Plains in Canada,
growing season varies from 80 to 150 days, and in the Taiga Shield
from 100 to 140 days.[9] Some sources claim 130 days growing season
as typical for the taiga.[10] Other sources mention that 50 - 100
frost-free days are characteristic.[11] Data for locations in southwest
Yukon gives 80 - 120 frost-free days.[12] The closed canopy boreal
forest in Kenozersky near Plesetsk, Arkhangelsk Province, Russia, on
average has 108 frost-free days.[13] The longest growing season is
found in the smaller areas with oceanic influences; in coastal areas of
Scandinavia and Finland, the growing season of the closed boreal forest can be 145 – 180 days.[14] The shortest
growing season is found at the northern taiga - tundra ecotone, where the northern taiga forest no longer can
grow
and the tundra dominates the landscape when the growing season is down to 50 – 70 days,[15] [16] and the 24-hr
average of the warmest month of the year usually is 10°C or less.[17] High latitudes mean that the sun does not
rise
far above the horizon, and less solar energy is received than further south. But the high latitude also ensures very
long summer days, as the sun stays above the horizon nearly 20 hours each day, with only around 6 hours of
daylight
occurring in the dark winters, depending on latitude. The areas of the taiga inside the Arctic circle have midnight
sun
in mid-summer and polar night in mid-winter.
Lakes and other water bodies are very common.
The Helvetinjärvi National Park, Finland, is
located in the southern boreal forest.[18]
The taiga experiences relatively low precipitation throughout the year
(generally 200–750 mm annually, 1,000 mm in some areas), primarily
as rain during the summer months, but also as fog and snow. As
evaporation is also low for most of the year, precipitation exceeds
evaporation, and is sufficient to sustain the dense vegetation growth.
Snow may remain on the ground for as long as nine months in the
northernmost extensions of the taiga ecozone.[19]
In general, taiga grows to the south of the 10 °C July isotherm, but
occasionally as far north as the 9 °C July isotherm.[20] The southern
limit is more variable, depending on rainfall; taiga may be replaced by
forest steppe south of the 15 °C July isotherm where rainfall is very
low, but more typically extends south to the 18 °C July isotherm, and locally where rainfall is higher (notably in
eastern Siberia and adjacent northern Manchuria) south to the 20 °C July isotherm. In these warmer areas the
taiga
has higher species diversity, with more warmth-loving species such as Korean Pine, Jezo Spruce, and
Manchurian
Fir, and merges gradually into mixed temperate forest or, more locally (on the Pacific Ocean coasts of North
America and Asia), into coniferous temperate rainforests.
Much of the area currently classified as taiga was recently glaciated. As the glaciers receded they left
depressions in
the topography that have since filled with water, creating lakes and bogs (especially muskeg soil) found
throughout
the taiga.
Taiga 22
Yukon, Canada. Several of the world's longest
rivers go through the taiga, including Ob,
Yenisei, Lena, and Mackenzie.
Soils
Taiga soil tends to be young and poor in nutrients. It lacks the deep,
organically-enriched profile present in temperate deciduous forests.[21]
The thinness of the soil is due largely to the cold, which hinders the
development of soil and the ease with which plants can use its
nutrients.[21] Fallen leaves and moss can remain on the forest floor for
a long time in the cool, moist climate, which limits their organic
contribution to the soil; acids from evergreen needles further leach the
soil, creating spodosol.[22] Since the soil is acidic due to the falling
pine needles, the forest floor has only lichens and some mosses
growing on it.
Flora
Boreal Forest near Lake Baikal in
Russia
Since North America and Asia used to be connected by the Bering land bridge, a
number of animal and plant species (more animals than plants) were able to
colonize both continents and are distributed throughout the taiga biome (see
Circumboreal Region). Others differ regionally, typically with each genus having
several distinct species, each occupying different regions of the taiga. Taigas also
have some small-leaved deciduous trees like birch, alder, willow, and poplar;
mostly in areas escaping the most extreme winter cold. However, the Dahurian
Larch tolerates the coldest winters in the northern hemisphere in eastern Siberia.
The very southernmost parts of the taiga may have trees such as oak, maple, elm,
and tilia scattered among the conifers, and there is usually a gradual transition
into a temperate mixed forest, such as the Eastern forest-boreal transition of
eastern Canada. In the interior of the continents with the driest climate, the boreal
forests might grade into temperate grassland.
There are two major types of taiga. The southern part is the closed canopy
forest, consisting of many closely-spaced trees with mossy ground cover. In
clearings in the forest, shrubs and wildflowers are common, such as the fireweed. The other type is the lichen
woodland or sparse taiga, with trees that are farther-spaced and lichen ground cover; the latter is
common in the
northernmost taiga.[23] In the northernmost taiga the forest cover is not only more sparse, but often stunted in
growth
form; moreover, ice pruned asymmetric Black Spruce (in North America) are often seen, with diminished foliage
on
the windward side.[24] In Canada, Scandinavia and Finland, the boreal forest is usually divided into three
subzones:
The high boreal (north boreal) or taiga zone; the middle boreal (closed forest); and the southern
boreal, a closed
canopy boreal forest with some scattered temperate deciduous trees among the conifers,[25] such as maple, elm
and
oak. This southern boreal forest has the longest and warmest growing season of the biome, and in some regions
(including Scandinavia, Finland and western Russia) this subzone is commonly used for agricultural purposes.
The
boreal forest is home to many types of berries; some are confined to the southern and middle closed boreal forest
(such as raspberry), others grow in most areas of the taiga (such as cranberry and cloudberry), and some can
grow in
both the taiga and the low arctic (southern part of) tundra (such as bilberry and lingonberry).
The forests of the taiga are largely coniferous, dominated by larch, spruce, fir, and pine. The woodland mix
varies
according to geography and climate so for example the Eastern Canadian forests ecoregion of the higher
elevations
of the Laurentian Mountains and the northern Appalachian Mountains in Canada is dominated by balsam fir
Abies
Taiga 23
balsamea, while further north the Eastern Canadian Shield taiga of northern Quebec and Labrador is notably
black
spruce Picea mariana and tamarack larch Larix laricina.
Evergreen species in the taiga (spruce, fir, and pine) have a number of adaptations specifically for survival in
harsh
taiga winters, although larch, the most cold-tolerant of all trees, is deciduous. Taiga trees tend to have shallow
roots
to take advantage of the thin soils, while many of them seasonally alter their biochemistry to make them more
resistant to freezing, called "hardening".[26] The narrow conical shape of northern conifers, and their
downward-drooping limbs, also help them shed snow.[26]
Because the sun is low in the horizon for most of the year, it is difficult for plants to generate energy from
photosynthesis. Pine, spruce and fir do not lose their leaves seasonally and are able to photosynthesize with their
older leaves in late winter and spring when light is good but temperatures are still too low for new growth to
commence. The adaptation of evergreen needles limits the water lost due to transpiration and their dark green
color
increases their absorption of sunlight. Although precipitation is not a limiting factor, the ground freezes during
the
winter months and plant roots are unable to absorb water, so desiccation can be a severe problem in late winter
for
evergreens.
Moss (Ptilium crista-castrensis) cover on the
floor of taiga
Although the taiga is dominated by coniferous forests, some broadleaf
trees also occur, notably birch, aspen, willow, and rowan. Many
smaller herbaceous plants grow closer to the ground. Periodic
stand-replacing wildfires (with return times of between 20–200 years)
clear out the tree canopies, allowing sunlight to invigorate new growth
on the forest floor. For some species, wildfires are a necessary part of
the life cycle in the taiga; some, e.g. Jack Pine have cones which only
open to release their seed after a fire, dispersing their seeds onto the
newly cleared ground. Grasses grow wherever they can find a patch of
sun, and mosses and lichens thrive on the damp ground and on the
sides of tree trunks. In comparison with other biomes, however, the
taiga has low biological diversity.
Coniferous trees are the dominant plants of the taiga biome. A very few species in four main genera are found:
the
evergreen spruce, fir, and pine, and the deciduous larch. In North America, one or two species of fir and one or
two
species of spruce are dominant. Across Scandinavia and western Russia, the Scots pine is a common component
of
the taiga, while taiga of the Russian Far East and Mongolia is dominated by larch.
Fauna
Brown bear, Kamchatka peninsula. Brown bears
are among the largest and most widespread taiga
predators.
The boreal forest, or taiga, supports a large range of animals. Canada's
boreal forest includes 85 species of mammals, 130 species of fish, and
an estimated 32,000 species of insects.[27] Insects play a critical role as
pollinators, decomposers, and as a part of the food chain. Many nesting
birds rely on them for food. The cold winters and short summers make
the taiga a challenging biome for reptiles and amphibians, which
depend on environmental conditions to regulate their body
temperatures, and there are only a few species in the boreal forest.
Some hibernate underground in winter.
The taiga is home to a number of large herbivorous mammals, such as
moose and reindeer/caribou. Some areas of the more southern closed
Taiga 24
boreal forest also have populations of other deer species such as the elk (wapiti) and roe deer.[28] [29] There is
also a
range of rodent species including beaver, squirrel, mountain hare, snowshoe hare, and vole. These species have
evolved to survive the harsh winters in their native ranges. Some larger mammals, such as bears, eat heartily
during
the summer in order to gain weight, and then go into hibernation during the winter. Other animals have adapted
layers of fur or feathers to insulate them from the cold.
A number of wildlife species threatened or endangered with extinction can be found in the Canadian boreal
forest,
including woodland caribou, American black bear, grizzly bear, and wolverine. Habitat loss, mainly due to
logging,
is the primary cause of decline for these species.
Due to the climate, carnivorous diets are an inefficient means of obtaining energy; energy is limited, and most
energy is lost between trophic levels. Predatory birds (owls and eagles) and other smaller carnivores, including
foxes
and weasels, feed on the rodents. Larger carnivores, such as lynx and wolves, prey on the larger animals.
Omnivores,
such as bears and raccoons are fairly common, sometimes picking through human garbage.
More than 300 species of birds have their nesting grounds in the taiga.[30] Siberian Thrush, White-throated
Sparrow,
and Black-throated Green Warbler migrate to this habitat to take advantage of the long summer days and
abundance
of insects found around the numerous bogs and lakes. Of the 300 species of birds that summer in the taiga only
30
stay for the winter.[31] These are either carrion-feeding or large raptors that can take live mammal prey,
including
Golden Eagle, Rough-legged Buzzard (also known as the Rough-legged Hawk), and Raven, or else seed-eating
birds, including several species of grouse and crossbills.
Threats
Human activities
Plesetsk Cosmodrome is situated in the taiga
Large areas of Siberia’s taiga have been harvested for lumber since the
collapse of the Soviet Union.[32] In Canada, eight percent of the boreal
forest is protected from development, the provincial government
allows forest management to occur on Crown land under rigorous
constraints. The main forestry practice in the boreal forest of Canada is
clearcutting, which involves cutting down most of the trees in a given
area, then replanting the forest as a monocrop (one species of tree) the
following season. Industry officials claim that this process emulates the
natural effects of a forest fire, which they claim clearcutting
suppresses, protecting infrastructure, communities and roads.
However, from an ecological perspective, this is a falsehood, for
several reasons, including: a) Removing most of the trees in a given area is usually done using large machines
which
disrupt the soil greatly, and the dramatic diminution of ground cover permits large-scale erosion and avalanches,
which further damage the habitat and sometimes engangers infrastructure, roads and communities. b)
Clearcutting
removes most of the biomass from an area, and the various macro and micro-nutrients it contains. This sudden
decrease in nutrients in an area, contrasts with a forest fire, which returns most of the nutrients to the soil. c)
Forest
fires leave standing snags, and leave patches of unburned trees. This helps preserve structure and micro-habitats
within the area, whereas clearcutting destroys most of these habitats. In the past, clearcuts upwards of 110 km˛
have
been recorded in the Canadian boreal forest. However, today 80% of clearcuts are less than 260 hectares(2.6
square
km). Some of the products from logged boreal forests include toilet paper, copy paper, newsprint, and lumber.
More
than 90% of boreal forest products from Canada are exported for consumption and processing in the United
States,
however with the recession and fewer US homes being built that has changed. Some of the larger cities situated
in
this biome are Murmansk,[33] Arkhangelsk, Yakutsk, Anchorage,[34] Yellowknife, Tromsř, Luleĺ, and Oulu.
Taiga 25
Most companies that harvest in Canadian forests are certified by an independent third party agency such as the
Forest Stewardship Council (FSC), Sustainable Forests Initiative (SFI), or the Canadian Standards Association
(CSA). While the certification process differs between these groups, all of them include forest stewardship,
respect
for aboriginal peoples, compliance with local, provincial or national environmental laws, forest worker safety,
education and training, and other environmental, business, and social requirements. The prompt renewal of all
harvest sites by planting or natural renewal is also required.
Climate change
The zone of latitude occupied by the boreal forest has experienced some of the greatest temperature increases on
Earth, especially during the last quarter of the twentieth century. Winter temperatures have increased more than
summer temperatures. The number of days with extremely cold temperatures (e.g., −20 to −40°C) has decreased
irregularly but systematically in nearly all the boreal region, allowing better survival for tree-damaging insects.
In
summer, the daily low temperature has increased more than the daily high temperature.[35] In Fairbanks, Alaska,
the
length of the frost-free season has increased from 60 – 90 days in the early twentieth century to about 120 days a
century later. Summer warming has been shown to increased water stress and reduce tree growth in dry areas of
the
southern boreal forest in central Alaska, western Canada and portions of far eastern Russia. Precipitation is
relatively
abundant in Scandinavia, Finland, northwest Russia and eastern Canada, where warmer summers accelerate tree
growth. As a consequence of this warming trend, the warmer parts of the boreal forests are susceptible to
replacement by grassland, parkland or temperate forest.[36] In Siberia, the taiga is converting from predominantly
needle-shedding larch trees to evergreen conifers in response to a warming climate. This is likely to further
accelerate warming, as the evergreen trees will absorp more of the suns rays. Given the vast size of the area,
such a
change has the potential to affect areas well outside of the region.[37] In much of the boreal forest in Alaska, the
growth of white spruce trees are stunted by unusually warm summers, while trees on some of the coldest fringes
of
the forest are experiencing faster growth than previously.[38] Lack of moisture in the warmer summers are also
stressing the birch trees of central Alaska.[39]
Taiga 26
Insects
Recent years have seen outbreaks of insect pests in forest-destroying plagues: the spruce-bark beetle
(Dendroctonus
rufipennis) in the Yukon Territory, Canada, and Alaska;[40] the aspen-leaf miner; the larch sawfly; the spruce
budworm (Choristoneura fumiferana);[41] the spruce coneworm.[42]
Protection
Peat bog in Dalarna, Sweden. Bogs and peatland
are widespread in the taiga. They are home to a
unique flora, and store vast amounts of carbon. In
western Eurasia, the pine is common in the boreal
forest.
Many nations are taking direct steps to protect the ecology of the taiga
by prohibiting logging, mining, oil and gas production, and other forms
of development. In February 2010 the Canadian government
established protection for 13,000 square kilometres of boreal forest by
creating a new 10,700 square kilometre park reserve in the Mealy
Mountains area of eastern Canada and a 3,000 square kilometre
waterway provincial park that follows alongside the Eagle River from
headwaters to sea.[43] The taiga stores enormous quantities of carbon,
possibly more than the temperate and tropical forests combined, much
of it in peatland.[44]
Natural disturbance
One of the biggest areas of research and a topic still full of unsolved
questions is the recurring disturbance of fire and the role it plays in
propagating the lichen woodland.[45] The phenomenon of wildfire by
lighting strike is the primary determinant of understory vegetation and
because of this, it is considered to be predominate driving force behind
community and ecosystem properties in the lichen woodland.[46] The
significance of fire is clearly evident when one considers that
understory vegetation influences tree seedling germination in the short term and decomposition of biomass and
nutrient availability in the long term.[46] The recurrent cycle of large, damaging fire occurs approximately every
70
to 100 years.[47] Understanding the dynamics of this ecosystem is entangled with discovering the successional
paths
that the vegetation exhibits after a fire. Trees, shrubs and lichens all recover from fire induced damage through
vegetative reproduction as well as invasion by propagules.[48] Seeds that have fallen and become buried provide
little
help in re-establishment of a species. The reappearance of lichens is reasoned to occur because of varying
conditions
and light/nutrient availability in each different microstate.[48] Several different studies have been done that have
led
to the formation of the theory that post-fire development can be propagated by any of four pathways: self
replacement, species-dominance relay, species replacement, or gap-phase self replacement.[45] Self replacement
is
simply the re-establishment of the pre-fire dominant species. Species-dominance relay is a sequential attempt of
tree
species to establish dominance in the canopy. Species replacement is when fires occur in sufficient frequency to
interrupt species dominance relay. Gap-Phase Self-Replacement is the least common and so far has only been
documented in Western Canada. It is a self replacement of the surviving species into the canopy gaps after a fire
kills
another species. The particular pathway taken after a fire disturbance depends on how the landscape is able to
support trees as well as fire frequency.[49] Fire frequency has a large role in shaping the original inception of the
lower forest line of the lichen woodland taiga.
Centuries ago, the southern limits of lichen woodland taiga were only being formed.[50] It has been hypothesized
and
subsequently proved by Serge Payette that the Spruce-Moss forest ecosystem was changed into the lichen
woodland
biome due to the initiation of two compounded strong disturbances.[50] The two disturbances were large fire and
the
Taiga 27
appearance and attack of the spruce budworm. The spruce budworm is a deadly insect to the spruce populations
in
the southern regions of the taiga. J.P. Jasinski confirmed this theory five years later stating “Their [lichen
woodlands]
persistence , along with their previous moss forest histories and current occurrence adjacent to closed moss
forests,
indicate that they are an alternative stable state to the spruce–moss forests”.[51]
Taiga ecoregions
Palearctic Boreal forests/taiga
East Siberian taiga Russia
Iceland boreal birch forests and alpine tundra Iceland
Kamchatka-Kurile meadows and sparse forests Russia
Kamchatka-Kurile taiga Russia
Northeast Siberian taiguhh Russia
Okhotsk-Manchurian taiga Russia
Sakhalin Island taiga Russia
Scandinavian and Russian taiga Finland, Norway, Russia, Sweden
Trans-Baikal conifer forests Mongolia, Russia
Urals montane tundra and taiga Russia
West Siberian taiga Russia
Puszcza Romnicka Poland, Russia
Nearctic Boreal forests/taiga
Alaska Peninsula montane taiga United States
Central Canadian Shield forests Canada, United States
Cook Inlet taiga United States
Copper Plateau taiga United States
Eastern Canadian forests Canada
Eastern Canadian Shield taiga Canada
Interior Alaska-Yukon lowland taiga Canada, United States
Mid-Continental Canadian forests Canada
Midwestern Canadian Shield forests Canada, United States
Muskwa-Slave Lake forests Canada
Newfoundland Highland forests Canada
Northern Canadian Shield taiga Canada
Northern Cordillera forests Canada
Northwest Territories taiga Canada
South Avalon-Burin oceanic barrens Canada
Southern Hudson Bay taiga Canada
Yukon Interior dry forests Canada
Taiga 28
References
[1] "taiga." Dictionary.com Unabridged (v 1.1). Random House, Inc. 12 Mar. 2008. web link (http:/ / dictionary. reference. com/
browse/ taiga)
[2] "Taiga biological station: FAQ" (http:/ / www. wilds. mb. ca/ taiga/ tbsfaq. html). Wilds.mb.ca. . Retrieved 2011-02-21.
[3] "radford:Taiga climate" (http:/ / www. radford. edu/ ~swoodwar/ CLASSES/ GEOG235/ biomes/ taiga/ taiga. html). Radford.edu. .
Retrieved
2011-02-21.
[4] "Marietta the Taiga and Boreal forest" (http:/ / www. marietta. edu/ ~biol/ biomes/ boreal. htm). Marietta.edu. . Retrieved 2011-02-21.
[5] "Yakutsk climate" (http:/ / www. worldclimate. com/ cgi-bin/ data. pl?ref=N62E129+ 1102+ 24959W). Worldclimate.com. 2007-02-04. .
Retrieved 2011-02-21.
[6] "WWF: Interior Alaska-Yukon lowland taiga" (http:/ / www. worldwildlife. org/ wildworld/ profiles/ terrestrial/ na/ na0607_full. html).
Worldwildlife.org. . Retrieved 2011-02-21.
[7] "WWF: The eastern forest - boreal transition" (http:/ / www. worldwildlife. org/ wildworld/ profiles/ terrestrial/ na/ na0406_full. html).
Worldwildlife.org. . Retrieved 2011-02-21.
[8] Canada: Taiga Shield reference (http:/ / www. enr. gov. nt. ca/ _live/ documents/ documentManagerUpload/ Taiga_Shield_References.
pdf)
[9] "Climate of Canadian ecozones" (http:/ / geography. ridley. on. ca/ CGC1D/ Students/ NORTH/ Geography Project/ Ecozones. htm).
Geography.ridley.on.ca. . Retrieved 2011-02-21.
[10] "Berkley: about biomes" (http:/ / www. ucmp. berkeley. edu/ exhibits/ biomes/ forests. php#boreal). Ucmp.berkeley.edu. . Retrieved
2011-02-21.
[11] "Taiga" (http:/ / www. blueplanetbiomes. org/ taiga. htm). Blueplanetbiomes. . Retrieved 2011-02-21.
[12] "Southwest Yukon:Frost-free days" (http:/ / yukon. taiga. net/ swyukon/ frost. cfm). Yukon.taiga.net. . Retrieved 2011-02-21.
[13] "Kenozersky National Park" (http:/ / www. wild-russia. org/ bioregion2/ 2-KenozerskyNP/ 2_kenoz. htm). Wild-russia.org. . Retrieved
2011-02-21.
[14] "University of Helsinki: Carabid diversity in Finnish taiga" (http:/ / www. sekj. org/ PDF/ anzf31/ anz31-123-129. pdf) (PDF). .
Retrieved
2011-02-21.
[15] "Tundra" (http:/ / www. blueplanetbiomes. org/ tundra. htm). Blueplanetbiomes. . Retrieved 2011-02-21.
[16] "NatureWorks:Tundra" (http:/ / www. nhptv. org/ NatureWorks/ nwep8a. htm). Nhptv.org. . Retrieved 2011-02-21.
[17] "The Arctic" (http:/ / www. saskschools. ca/ ~gregory/ arctic/ Aintro. html). saskschools.ca. . Retrieved 2011-02-21.
[18] "Tampere, Finland - southern boreal climate" (http:/ / www. worldclimate. com/ cgi-bin/ data. pl?ref=N61E023+ 1102+ 02944W).
Worldclimate.com. 2007-02-04. . Retrieved 2011-02-21.
[19] A.P. Sayre, Taiga, (New York: Twenty-First Century Books, 1994) 16.
[20] Arno & Hammerly 1984, Arno et al. 1995
[21] Sayre, 19.
[22] Sayre, 19-20.
[23] Sayre, 12-3.
[24] C. Michael Hogan, Black Spruce: Picea mariana, GlobalTwitcher.com, ed. Nicklas Stromberg, November, 2008 (http:/ /
globaltwitcher.
auderis. se/ artspec_information. asp?thingid=44751)
[25] George H. La Roi. "Boreal forest" (http:/ / thecanadianencyclopedia. com/ index. cfm?PgNm=TCE& Params=A1ARTA0000888). The
Canadian Encyclopedia. . Retrieved 2011-02-21.
[26] Sayre, 23.
[27] "hww:Nature in the boreal forest biome" (http:/ / www. hww. ca/ hww2. asp?id=354). Hww.ca. . Retrieved 2011-02-21.
[28] "Wapiti facts and range" (http:/ / www. hww. ca/ hww2. asp?id=98). Hww.ca. . Retrieved 2011-02-21.
[29] "western roe deer: facts and range" (http:/ / www. borealforest. org/ world/ mammals/ western_roe_deer. htm). Borealforest.org. .
Retrieved
2011-02-21.
[30] "Boreal songbird initiative" (http:/ / www. borealbirds. org/ forest. shtml). Borealbirds.org. . Retrieved 2011-02-21.
[31] Sayre, 28.
[32] "Taiga Deforestation" (http:/ / www. american. edu/ TED/ TAIGA. HTM). American.edu. . Retrieved 2011-02-21.
[33] "Murmansk climate" (http:/ / www. worldclimate. com/ cgi-bin/ data. pl?ref=N68E033+ 1202+ 0006410G2). Worldclimate.com.
2007-02-04. . Retrieved 2011-02-21.
[34] "Anchorage climate" (http:/ / www. worldclimate. com/ cgi-bin/ data. pl?ref=N61W149+ 1302+ 502820C). Worldclimate.com. 2007-
02-04.
. Retrieved 2011-02-21.
[35] http:/ / www. arcus. org/ witness-the-arctic/ 2009/ 3/ article/ 507
[36] http:/ / www. libraryindex. com/ pages/ 3196/ Boreal-Forests-Climate-Change. html
[37] http:/ / www. sciencedaily. com/ releases/ 2011/ 03/ 110325022352. htm
[38] http:/ / www. newsminer. com/ view/ full_story/ 12094427/ article-New-study-states-boreal-forests-shifting-as-Alaska-warms
[39] http:/ / www. scientificamerican. com/ article. cfm?id=forest-changes-in-alaska-reveal-changing-climate
[40] "A New Method to Reconstruct Bark Beetle Outbreaks" (http:/ / www. colorado. edu/ INSTAAR/ AW2004/ get_abstr. html?id=88).
Colorado.edu. . Retrieved 2011-02-21.
[41] "Spruce budworm and sustainable management of the boreal forest" (http:/ / cfs. nrcan. gc. ca/ subsite/ budworm). Cfs.nrcan.gc.ca.
2007-12-05. . Retrieved 2011-02-21.
Taiga 29
[42] http:/ / www. fs. fed. us/ pnw/ pubs/ journals/ pnw_2006_chapin001. pdf
[43] Braun, David (February 7, 2010). "Boreal landscapes added to Canada's parks Boreal landscapes added to Canada's parks" (http:/ /
blogs.
nationalgeographic. com/ blogs/ news/ chiefeditor/ 2010/ 02/ boreal-landscapes-added-to-canada-parks. html). NatGeo News Watch:
News
Editor David Braun's Eye on the World. National Geographic Society. . Retrieved 17 February 2010.
[44] USA. "Boreal forest and global change" (http:/ / www. ncbi. nlm. nih. gov/ pmc/ articles/ PMC2387060/ ). Ncbi.nlm.nih.gov. .
Retrieved
2011-02-21.
[45] Kurkowski, 1911.
[46] Nilsson, 421.
[47] Johnson, 212.
[48] Johnson, 200
[49] Kurkowski, 1912.
[50] Payette, 289.
[51] Jasinski, 561.
General references
• Arno, S. F. & Hammerly, R. P. 1984. Timberline. Mountain and Arctic Forest Frontiers. The
Mountaineers,
Seattle. ISBN 0-89886-085-7
• Arno, S. F., Worral, J., & Carlson, C. E. (1995). Larix lyallii: Colonist of tree line and talus sites. Pp. 72–78 in
Schmidt, W. C. & McDonald, K. J., eds., Ecology and Management of Larix Forests: A Look
Ahead. USDA
Forest Service General Technical Report GTR-INT-319.
• Nilsson, M.C. "Understory vegetation as a forest ecosystem driver, evidence from the northern Swedish boreal
forest." Frontiers in Ecology and the Environment. 3.8 (2005): 421-428.
• Kurkowski, Thomas. "Relative Importance of Different Secondary Successional Pathways in an Alaskan
Boreal
Forest." Canadian Journal of Forest Research. 38. (2008): 1911-1923.
• Payette, Serge. "Origin of the lichen woodland at its southern range limit in eastern Canada: the catastrophic
impact of insect defoliators and fire on the spruce-moss forest." Canadian journal of forest research. 30.2 (2000):
288-305.
• Johnson, E.A. "Vegetation Organization and Dynamics of Lichen Woodland Communities in the Northwest
Territories." Ecology. 62.1 (1981): 200-215.
• Jasinski, J.P. "The Creation of Alternative Stable States in Southern Boreal Forest: Quebec, Canada."
Ecological
Monographs. 75.4 (2005): 561-583.
Further reading
• Sayre, April Pulley (1994), Taiga (http:/ / books. google. ca/ books?id=yvGkDWBoXHsC& lpg=PP1&
dq=Taiga& pg=PP1#v=onepage& q& f=true), Twenty-First Century Books, ISBN 0805028307
• Day, Trevor; Richard Garratt (2006), Taiga (http:/ / books. google. ca/ books?id=n3sdmk_SCXsC&
lpg=PP1&
dq=Taiga& pg=PP1#v=onepage& q& f=true), Facts On File, ISBN 0816053294
External links
• The Conservation Value of the North American Boreal Forest from an Ethnobotanical perspective (http:/ /
www.
borealbirds. org/ ethnobotany. shtml) a report by the Boreal Songbird Initiative (http:/ / borealbirds. org)
• Boreal Canadian Initiative (http:/ / www. borealcanada. ca)
• International Boreal Conservation campaign (http:/ / www. interboreal. org)
• Tundra and Taiga (http:/ / www. barrameda. com. ar/ ecology/ the-tundra-and-taiga. htm)
• Threats to Boreal Forests (http:/ / www. greenpeace. org/ canada/ en/ campaigns/ boreal) Greenpeace
• Campaign against lumber giant Weyerhaeuser's logging practices in the Canadian boreal forest (http:/ / ran.
org/
what_we_do/ old_growth/ campaigns/ wake_up_weyerhaeuser/ ) Rainforest Action Network
• Arctic and Taiga (http:/ / www. canadiangeographic. ca/ atlas/ themes. aspx?id=artic& lang=) Canadian
Geographic
Taiga 30
• Terraformers (http:/ / taiga. terraformers. ca/ ) Canadian Taiga Conservation Foundation
• Coniferous Forest, Earth Observatory (http:/ / earthobservatory. nasa. gov/ Laboratory/ Biome/ bioconiferous.
html) NASA
• Taiga Rescue Network (TRN) (http:/ / www. taigarescue. org) A network of NGOs, indigenous peoples or
individuals that works to protect the boreal forests.
• Index of Boreal Forests/Taiga ecoregions (http:/ / www. cas. vanderbilt. edu/ bioimages/ ecoframe-list. htm) at
bioimages.vanderbilt.edu
• The Canadian Boreal Forest (http:/ / www. nature. org/ wherewework/ northamerica/ canada/ work/ art12507.
html) The Nature Conservancy and its partners
• Slater museum of natural history: Taiga (http:/ / www. ups. edu/ x6111. xml)
• Taiga Biological Station (http:/ / www. wilds. mb. ca/ taiga/ index. html) founded by Dr. William (Bill) Pruitt,
Jr.,
University of Manitoba.
Tropical and subtropical grasslands,

savannas,
and shrublands
Tropical and subtropical grasslands, savannas, and shrublands are a grassland
terrestrial biome located in
semi-arid to semi-humid climate regions of subtropical and tropical latitudes. Grasslands are dominated by grass
and
other herbaceous plants. Savannas are grasslands with scattered trees. Shrublands are dominated by woody or
herbaceous shrubs.
Rainfall in tropical and subtropical grasslands, savannas, and shrublands is between 450 and 1500 millimeters
(18 to
60 inches) a year, and can be highly seasonal, with the entire year's rainfall sometimes occurring within a couple
of
weeks.
Tropical and subtropical grasslands, savannas, and shrublands occur on all continents but Antarctica. They are
widespread on Africa, and are also found all throughout South Asia, the northern parts of South America and
Australia, and the southern United States.
African Savannas occur between forest or woodland regions and grassland regions. The climate varies,
with an
average temperature of 27 °C with peaks of 30 °C in April and October, and between 300 and 1500 millimetres
of
rain per year. Flora includes:
• Acacia (Tree)
• Loses leaves in dry season to conserve moisture
• Leafy canopy in rainy season (flattened by winds)
• Baobab (Tree)
• Stores water in trunk
• Thick bark to protect it from fires in dry season
• Long tap roots to reach underground moisture
• Few leaves to reduce water loss through transpiration
• Grass
• Grow quickly to 3 or 4 meters in clumps
• Shoots die in dry season leaving only roots
• Low shrubs
• Drought resistant
• Small
• Thorns not leaves
Tropical and subtropical grasslands, savannas, and
shrublands ecoregions
Afrotropic Tropical and subtropical grasslands, savannas, and shrublands
Angolan Miombo woodlands Angola
Angolan Mopane woodlands Angola, Namibia
Ascension scrub and grasslands Ascension Island
Central Zambezian Miombo
woodlands
Angola, Burundi, Democratic Republic of the Congo, Malawi, Tanzania, Zambia
East Sudanian savanna Cameroon, Central African Republic, Chad, Democratic Republic of the Congo, Eritrea, Ethiopia Sudan,
Uganda
Eastern Miombo woodlands Mozambique, Tanzania
Guinean forest-savanna mosaic Benin, Burkina Faso, Cameroon, Gambia, Ghana, Guinea, Guinea Bissau, Côte d'Ivoire, Nigeria, Senegal,
Togo
Itigi-Sumbu thicket Tanzania, Zambia
Kalahari Acacia-Baikiaea
woodlands
Botswana, Namibia, South Africa, Zimbabwe
Mandara Plateau mosaic Cameroon, Nigeria
Northern Acacia-Commiphora
bushlands and thickets
Ethiopia, Kenya, Sudan, Uganda
Northern Congolian forest-savanna
mosaic
Cameroon, Central African Republic, Democratic Republic of the Congo, Sudan, Uganda
Sahel Burkina Faso, Cameroon, Chad, Eritrea, Ethiopia, Mali, Mauritania, Niger, Nigeria, Senegal, Sudan
Serengeti Kenya, Tanzania
Somali Acacia-Commiphora
Eritrea, Ethiopia, Kenya, Somalia, Sudan
Southern Acacia-Commiphora
Kenya, Tanzania
Southern Africa bushveld Botswana, South Africa, Zimbabwe
Southern Congolian forest-savanna
mosaic
Angola, Democratic Republic of the Congo
Southern Miombo woodlands Malawi, Mozambique, Zambia, Zimbabwe
Sudan grass-belt southern Senegal, southern Mali, Guinea, northern Cote d'Ivoire, Burkina Faso, northern Ghana, northern
Benin, northern Togo, central Nigeria, northern Cameroon, southern Chad, southern Sudan, western Ethiopia
Saint Helena scrub and woodlands Saint Helena
Victoria Basin forest-savanna
mosaic
Burundi, Kenya, Rwanda, Tanzania, Uganda
West Sudanian savanna Benin, Burkina Faso, Gambia, Ghana, Guinea, Côte d'Ivoire, Niger, Nigeria, Senegal
Western Congolian forest-savanna
mosaic
Angola, Democratic Republic of the Congo, Republic of the Congo
Western Zambezian grasslands Angola, Zambia
Zambezian and Mopane woodlands Botswana, Malawi, Mozambique, Namibia, South Africa, Swaziland, Zambia, Zimbabwe
Zambezian Baikiaea woodlands Angola, Botswana, Namibia, Zambia, Zimbabwe
Australasia Tropical and subtropical grasslands, savannas, and shrublands
Arnhem Land tropical savanna Australia
Brigalow tropical savanna Australia
Cape York tropical savanna Australia
Carpentaria tropical savanna Australia
Einasleigh upland savanna Australia
Kimberly tropical savanna Australia
Mitchell grass downs Australia
Trans Fly savanna and grasslands Indonesia, Papua New Guinea
Victoria Plains tropical savanna Australia
Indomalaya Tropical and subtropical grasslands, savannas, and shrublands
Terai-Duar savanna and grasslands Bhutan, India, Nepal, Philippines
Nearctic Tropical and subtropical grasslands, savannas, and shrublands
Pre-Columbian savannas of North America (greatly declined) Canada, United States
Eastern savannas of the United States (greatly declined) United States
Central Hardwood Region (greatly declined) United States
Western Gulf coastal grasslands Mexico, United States
Neotropic Tropical and subtropical grasslands, savannas, and shrublands
Beni savanna Bolivia
Campos Rupestres montane savanna Brazil
Cerrado Bolivia, Brazil, Paraguay
Clipperton Island shrub and grasslands Clipperton Island is an overseas territory of France
Córdoba montane savanna Argentina
Guyanan savanna Brazil, Guyana, Venezuela
Gran Chaco Argentina, Brazil, Paraguay
Llanos Colombia, Venezuela
Uruguayan savanna Argentina, Brazil, Uruguay
Oceania Tropical and subtropical grasslands, savannas, and shrublands
Hawaiian tropical high shrublands Hawaiʻi
Hawaiian tropical low shrublands Hawaiʻi
Northwestern Hawaii scrub Hawaiʻi
External links
• (World Wildlife Fund) [1]
References
[1] http:/ / www. panda. org/ news_facts/ education/ university/ habitats/ index. cfm
Temperate grasslands, savannas,

and shrublands
A restored Illinois grassland ecosystem at Morton Arboretum.
Temperate grasslands, savannas, and shrublands is
a terrestrial biome whose predominant vegetation
consists of grasses and/or shrubs. The climate is
temperate and semi-arid to semi-humid.
• Temperature: warm to hot season (often with a cold
to freezing season in winter)
• Soil: fertile with rich nutrients and minerals
• Plants: grass; trees or shrubs in savanna and
shrubland
• Animals: large, grazing mammals; birds
Steppes/shortgrass prairies are short grasslands that
occur in semi-arid climates. Tallgrass prairies are tall
grasslands in areas of higher rainfall. Heaths and pastures are, respectively, low shrublands and grasslands where
forest growth is hindered by human activity, not climate.
Tall grasslands, including the tallgrass prairie of North America and the Humid Pampas of Argentina, have
moderate
rainfall and rich soils which make them ideally suited to extensive agriculture, and tall grassland ecoregions
include
some of the most productive grain-growing regions in the world.
Savannas are areas with both grass and trees, but the trees do not form a canopy as they would in a forest.
Temperate grasslands, savannas, and shrublands
ecoregions
Afrotropic Temperate grasslands, savannas, and shrublands
Al Hajar Al Gharbi montane woodlands Oman
Amsterdam and Saint-Paul Islands temperate grasslands Amsterdam Island, Saint-Paul Island
Tristan da Cunha-Gough Islands shrub and grasslands Tristan da Cunha, Gough Island
Australasia Temperate grasslands, savannas, and shrublands
Canterbury-Otago tussock grasslands New Zealand
Eastern Australia mulga shrublands Australia
Southeast Australia temperate savanna Australia
Nearctic Temperate grasslands, savannas, and shrublands
California Central Valley grasslands United States
Canadian aspen forests and parklands Canada, United States
Central and Southern mixed grasslands United States
Central forest-grasslands transition United States
Central tall grasslands United States
Columbia Plateau United States
Edwards Plateau savanna United States
Flint Hills tall grasslands United States
Montana valley and foothill grasslands United States
Nebraska Sand Hills mixed grasslands United States
Northern mixed grasslands Canada, United States
Northern short grasslands Canada, United States
Northern tall grasslands Canada, United States
Palouse grasslands United States
Texas blackland prairies United States
Western short grasslands United States
Neotropic Temperate grasslands, savannas, and shrublands
Argentine Espinal Argentina
Argentine Monte Argentina
Humid Pampas Argentina
Patagonian grasslands Argentina, Chile
Patagonian steppe Argentina, Chile
Semi-arid Pampas Argentina
Palearctic Temperate grasslands, savannas, and shrublands
Alai-Western Tian Shan steppe Kazakhstan, Tajikistan, Uzbekistan
Altai steppe and semi-desert Kazakhstan
Central Anatolian steppe Turkey
Daurian forest steppe China, Mongolia, Russia
Eastern Anatolian montane steppe Armenia, Iran, Turkey
Emin Valley steppe China, Kazakhstan
Faroe Islands boreal grasslands Faroe Islands, part of Denmark
Gissaro-Alai open woodlands Kyrgyzstan, Tajikistan, Uzbekistan
Kazakh forest steppe Kazakhstan, Russia
Kazakh steppe Kazakhstan, Russia
Kazakh upland Kazakhstan
Middle East steppe Iraq, Syria
Mongolian-Manchurian grassland China, Mongolia, Russia
Pontic steppe Kazakhstan, Moldova, Romania, Russia, Ukraine
Sayan Intermontane steppe Russia
Selenge-Orkhon forest steppe Mongolia, Russia
South Siberian forest steppe Russia
Tian Shan foothill arid steppe China, Kazakhstan, Kyrgyzstan
External links
• Temperate Grassland [1]
References
[1] http:/ / www. barrameda. com. ar/ ecology/ temperate-grassland. htm
Flooded grasslands and savannas 36
Flooded grasslands and savannas

Flooded grasslands and savannas is a terrestrial biome. Its component ecoregions are generally
located at
subtropical and tropical latitudes, which are flooded seasonally or year-round.
Characteristics
Flooded grasslands and savannas is characterized by:
• water : very wet
• temperature : warm
• soils : nutrient rich soil
External links
• More on the Flooded grasslands and savannas biome [1]
References
Montane grasslands and shrublands

Montane grasslands and shrublands is a biome defined by the World Wildlife Fund. The biome
includes high
altitude (montane, subalpine, and alpine) grasslands and shrublands around the world.
Montane grasslands and shrublands located above the tree line are commonly known as alpine tundra, which
occurs
in mountain regions around the world. Below the tree line are subalpine and montane grasslands and shrublands.
Stunted subalpine forests are known as krummholz, and occur just below the tree line, where harsh, windy
conditions and poor soils create dwarfed and twisted forests of slow-growing trees.
Montane grasslands and shrublands, particularly in subtropical and tropical regions, often evolved as virtual
islands,
separated from other montane regions by warmer, lower elevation regions, and are frequently home to many
distinctive and endemic plants which evolved in response to the cool, wet climate and abundant tropical sunlight.
Characteristic plants of these habitats display adaptations such as rosette structures, waxy surfaces, and hairy
leaves.
A unique feature of many wet tropical montane regions is the presence of giant rosette plants from a variety of
plant
families, such as Lobelia (Afrotropic), Puya (Neotropic), Cyathea (New Guinea), and Argyroxiphium
(Hawaii).
The most extensive Montane grasslands and shrublands occur in the Neotropic Paramo of the Andes Mountains.
This
biome also occurs in the mountains of east and central Africa, Mount Kinabalu of Borneo, highest elevations of
the
Western Ghats in South India and the Central Highlands of New Guinea.
Where conditions are drier, one finds montane grasslands, savannas, and woodlands, like the Ethiopian
Highlands,
and montane steppes, like the steppes of the Tibetan Plateau.
Montane grassland and shrubland ecoregions
Afrotropic Montane grasslands and shrublands
Angolan montane forest-grassland mosaic Angola
Angolan Scarp savanna and woodlands Angola
Drakensberg alti-montane grasslands and woodlands Lesotho, South Africa
Drakensberg montane grasslands, woodlands and forests Lesotho, South Africa, Swaziland
East African montane moorlands Kenya, Sudan, Tanzania, Uganda
Eastern Zimbabwe montane forest-grassland mosaic Mozambique, Zimbabwe
Ethiopian montane grasslands and woodlands Ethiopia
Ethiopian montane moorlands Ethiopia
Highveld grasslands Lesotho, South Africa
Jos Plateau forest-grassland mosaic Nigeria
Madagascar ericoid thickets Madagascar
Maputaland-Pondoland bushland and thickets Mozambique, South Africa, Swaziland
Ruwenzori-Virunga montane moorlands Democratic Republic of the Congo, Rwanda, Uganda
South Malawi montane forest-grassland mosaic Malawi, Mozambique
Southern Rift montane forest-grassland mosaic Malawi, Tanzania
Australasia Montane grasslands and shrublands
Australian Alps montane grasslands Australia
Central Range sub-alpine grasslands Indonesia, Papua New Guinea
Southland montane grasslands New Zealand
Indomalaya Montane grasslands and shrublands
Kinabalu montane alpine meadows Malaysia
Neotropic Montane grasslands and shrublands
Central Andean dry puna Argentina, Bolivia, Chile
Central Andean puna Argentina, Bolivia, Peru
Central Andean wet puna Bolivia, Peru
Cordillera Central páramo Ecuador, Peru
Cordillera de Merida páramo Venezuela
Northern Andean páramo Colombia, Ecuador
Santa Marta páramo Colombia
Talamanca Paramo Costa Rica, Panama
Southern Andean steppe Argentina, Chile
Zacatonal Mexico, Guatemala
Palearctic Montane grasslands and shrublands
Altai alpine meadow and tundra People's Republic of China, Kazakhstan, Mongolia, Russia
Central Tibetan Plateau alpine steppe China
Eastern Himalayan alpine shrub and meadows Bhutan, Burma, People's Republic of China, India, Nepal
Ghorat-Hazarajat alpine meadow Afghanistan
Hindu Kush alpine meadow Afghanistan, Pakistan
Karakoram-West Tibetan Plateau alpine steppe Afghanistan, People's Republic of China, India, Pakistan
Khangai Mountains alpine meadow Mongolia
Kopet Dag woodlands and forest steppe Iran, Turkmenistan
Kuhrud-Kohbanan Mountains forest steppe Iran
Mediterranean High Atlas juniper steppe Morocco
North Tibetan Plateau-Kunlun Mountains alpine desert China
Northwestern Himalayan alpine shrub and meadows People's Republic of China, India, Pakistan
Ordos Plateau steppe China
Pamir alpine desert and tundra Afghanistan, China, Kyrgyzstan, Tajikistan
Qilian Mountains subalpine meadows China
Sayan Alpine meadows and tundra Mongolia, Russia
Southeast Tibet shrub and meadows China
Sulaiman Range alpine meadows Afghanistan, Pakistan
Tian Shan montane steppe and meadows China, Kazakhstan, Kyrgyzstan
Tibetan Plateau alpine shrub and meadows People's Republic of China
Western Himalayan alpine shrub and meadows India, Nepal
Yarlung Zambo arid steppe China
External links
• more on this biome [1]
References
[1] http:/ / www. panda. org/ about_our_earth/ ecoregions/ about/ habitat_types/ selecting_terrestrial_ecoregions/ habitat10. cfm
Tundra 39
Tundra
Tundra
Map of arctic tundra shown in orange
Geography
Climate type ET
In physical geography, tundra is a biome where the tree growth is hindered by low temperatures and short
growing
seasons. The term tundra comes through Russian тундра from the Kildin Sami word tūndâr "uplands,"
"treeless
mountain tract."[1] There are three types of tundra: Arctic tundra,[2] alpine tundra,[2] and Antarctic
tundra.[3] In
tundra, the vegetation is composed of dwarf shrubs, sedges and grasses, mosses, and lichens. Scattered trees
grow in
some tundra. The ecotone (or ecological boundary region) between the tundra and the forest is known as the tree
line
or timberline.
Arctic
Arctic tundra occurs in the far Northern Hemisphere, north of the taiga belt. The word "tundra" usually refers
only to
the areas where the subsoil is permafrost, or permanently frozen soil. (It may also refer to the treeless plain in
general, so that northern Sápmi would be included.) Permafrost tundra includes vast areas of northern Russia and
Canada.[2] The polar tundra is home to several peoples who are mostly nomadic reindeer herders, such as the
Nganasan and Nenets in the permafrost area (and the Sami in Sápmi).
Arctic tundra contains areas of stark landscape and is frozen for much of the year. The soil there is frozen from
25–90 cm (9.8–35.4 inches) down, and it is impossible for trees to grow. Instead, bare and sometimes rocky land
can
only support low growing plants such as moss, heath (Ericaceae varieties such as crowberry and black
bearberry),
and lichen. There are two main seasons, winter and summer, in the polar tundra areas. During the winter it is
very
cold and dark, with the average temperature around −28 °C (−18 °F), sometimes dipping as low as −50 °C (−58
°F).
However, extreme cold temperatures on the tundra do not drop as low as those experienced in taiga areas further
south (for example, Russia's and Canada's lowest temperatures were recorded in locations south of the tree line).
During the summer, temperatures rise somewhat, and the top layer of the permafrost melts, leaving the ground
very
soggy. The tundra is covered in marshes, lakes, bogs and streams during the warm months. Generally daytime
temperatures during the summer rise to about 12 °C (54 °F) but can often drop to 3 °C (37 °F) or even below
freezing. Arctic tundras are sometimes the subject of habitat conservation programs. In Canada and Russia,
many of
these areas are protected through a national Biodiversity Action Plan.
Tundra 40
Tundra in Alaska
The tundra is a very windy area, with winds often blowing upwards of
48–97 km/h (30–60 miles an hour). However, in terms of precipitation,
it is desert-like, with only about 15–25 cm (6–10 inches) falling per
year (the summer is typically the season of maximum precipitation).
During the summer, the permafrost thaws just enough to let plants
grow and reproduce, but because the ground below this is frozen, the
water cannot sink any lower, and so the water forms the lakes and
marshes found during the summer months. Although precipitation is
light, evaporation is also relatively minimal.
The biodiversity of the tundras is low: 1,700 species of vascular plants and only 48 land mammals can be found,
although millions of birds migrate there each year for the marshes.[4] There are also a few fish species such as
the
flatfish. There are few species with large populations. Notable animals in the Arctic tundra include caribou
(reindeer), musk ox, arctic hare, arctic fox, snowy owl, lemmings, and polar bears (only the extreme north).[5]
Due to the harsh climate of the Arctic tundra, regions of this kind have seen little human activity, even though
they
are sometimes rich in natural resources such as oil and uranium. In recent times this has begun to change in
Alaska,
Russia, and some other parts of the world.
A severe threat to the tundras, specifically to the permafrost, is global warming. The melting of the permafrost in
a
given area on human time scales (decades or centuries) could radically change which species can survive there.
[6]
Another concern is that about one third of the world's soil-bound carbon is in taiga and tundra areas. When the
permafrost melts, it releases carbon in the form of carbon dioxide and methane,[7] both of which are greenhouse
gases. The effect has been observed in Alaska. In the 1970s the tundra was a carbon sink, but today, it is a
carbon
source.[8]
Antarctic
Tundra on the Péninsule Rallier du Baty,
Kerguelen Islands.
Antarctic tundra occurs on Antarctica and on several Antarctic and
subantarctic islands, including South Georgia and the South Sandwich
Islands and the Kerguelen Islands. Most of Antarctica is too cold and
dry to support vegetation, and most of the continent is covered by ice
fields. However, some portions of the continent, particularly the
Antarctic Peninsula, have areas of rocky soil that support plant life.
The flora presently consists of around 300–400 lichens, 100 mosses,
25 liverworts, and around 700 terrestrial and aquatic algae species,
which live on the areas of exposed rock and soil around the shore of
the continent. Antarctica's two flowering plant species, the Antarctic
hair grass (Deschampsia antarctica) and Antarctic pearlwort
(Colobanthus quitensis), are found on the northern and western parts of the Antarctic Peninsula.[9]
In contrast with the Arctic tundra, the Antarctic tundra lacks a large mammal fauna, mostly due to its physical
isolation from the other continents. Sea mammals and sea birds, including seals and penguins, inhabit areas near
the
shore, and some small mammals, like rabbits and cats, have been introduced by humans to some of the
subantarctic
islands. The Antipodes Subantarctic Islands tundra ecoregion includes the Bounty Islands, Auckland Islands,
Antipodes Islands, the Campbell Island group, and Macquarie Island.[10] Species endemic to this ecoregion
include
Nematoceras dienemum and Nematoceras sulcatum, the only Subantarctic orchids; the royal
penguin; and the
Antipodean albatross.[10]
The flora and fauna of Antarctica and the Antarctic Islands (south of 60° south latitude) are protected by the
Antarctic Treaty.[11]
Tundra 41
Alpine
Hikers traversing the Franconia Ridge in the
White Mountains, New Hampshire, United
States, much of which is in the alpine zone.
Alpine tundra does not contain trees because it has high altitude.
Alpine tundra is distinguished from arctic tundra, because alpine
tundra typically does not have permafrost, and alpine soils are
generally better drained than arctic soils. Alpine tundra transitions to
subalpine forests below the tree line; stunted forests occurring at the
forest-tundra ecotone are known as Krummholz.
Alpine tundra occurs in mountains worldwide. The flora of the alpine
tundra is characterized by dwarf shrubs close to the ground. The cold
climate of the alpine tundra is caused by the low air pressure, and is
similar to polar climate.
Climatic classification
Tundra region with fjords, glaciers and
mountains. Kongsfjorden, Spitsbergen.
Tundra climates ordinarily fit the Köppen climate classification ET,
signifying a local climate in which at least one month has an average
temperature high enough to melt snow (0°C or 32°F), but no month
with an average temperature in excess of (10°C/50°F). The cold limit
generally meets the EF climates of permanent ice and snows; the
warm-summer limit generally corresponds with the poleward or
altitudinal limit of trees, where they grade into the subarctic climates
designated Dfd and Dwd (extreme winters as in parts of Siberia), Dfc
typical in Alaska, Canada, European Russia, and Western Siberia (cold
winters with months of freezing), or even Cfc (no month colder than
-3°C as in parts of Iceland and southernmost South America). Tundra
climates as a rule are hostile to woody vegetation even where the
winters are comparatively mild by polar standards, as in Iceland.
Despite the potential diversity of climates in the ET category involving precipitation, extreme temperatures, and
relative wet and dry seasons, this category is rarely subdivided. Rainfall and snowfall are generally slight due to
the
low vapor pressure of water in the chilly atmosphere, but as a rule potential evapotranspiration is extremely low,
allowing soggy terrain of swamps and bogs even in places that get precipitation typical of deserts of lower and
middle latitudes. The amount of native tundra biomass depends more on the local temperature than the amount
of
precipitation.
Tundra 42
References
[1] Aapala, Kirsti. "Tunturista jängälle" (http:/ / web. archive. org/ web/ 20061001211854/ http:/ / www. kotus. fi/ julkaisut/ ikkunat/ 1999/
kielii1999_19. shtml). Kieli-ikkunat. Archived from the original (http:/ / www. kotus. fi/ julkaisut/ ikkunat/ 1999/ kielii1999_19. shtml) on
2006-10-01. . Retrieved 2009-01-19.
[2] "The Tundra Biome" (http:/ / www. ucmp. berkeley. edu/ glossary/ gloss5/ biome/ tundra. html). The World's Biomes. . Retrieved
2006-03-05.
[3] "Terrestrial Ecoregions: Antarctica"" (http:/ / www. nationalgeographic. com/ wildworld/ profiles/ terrestrial_an. html). Wild World.
National
Geographic. . Retrieved 2009-11-02.
[4] "Great Plain of the Koukdjuak" (http:/ / www. ibacanada. com/ site. jsp?siteID=NU078& seedet=N). Ibacanada.com. . Retrieved 2011-
02-16.
[5] "Tundra" (http:/ / www. blueplanetbiomes. org/ tundra. htm). Blue Planet Biomes. . Retrieved 2006-03-05.
[6] "Tundra Threats" (http:/ / science. nationalgeographic. com/ science/ earth/ habitats/ tundra-threats. html). National Geographic. .
Retrieved
2008-04-03.
[7] "Global Warming: Methane Could Be Far Worse Than Carbon Dioxide" (http:/ / www. communicationagents. com/ sepp/ 2005/ 02/ 01/
global_warming_methane_could_be_far_worse_than_carbon_dioxide. htm). Sepp Hasslberger. 2005-02-01. .
[8] Oechel, Walter C.; et al. (1993). "Recent change of Arctic tundra ecosystems from a net carbon dioxide sink to a source". Nature
361 (6412):
520–523. doi:10.1038/361520a0.
[9] "Terrestrial Plants" (http:/ / www. antarctica. ac. uk/ About_Antarctica/ Wildlife/ Plants/ index. html). British Antarctic Survey:
About
Antarctica. . Retrieved 2006-03-05.
[10] "Antipodes Subantarctic Islands tundra" (http:/ / www. worldwildlife. org/ wildworld/ profiles/ terrestrial/ aa/ aa1101_full. html). Wild
World Full Report. World Wildlife Fund. . Retrieved 2009-11-02.
[11] "Protocol on Environmental Protection to the Antarctic Treaty" (http:/ / www. antarctica. ac. uk/ About_Antarctica/ Treaty/ protocol.
html).
British Antarctic Survey: About Antarctica. . Retrieved 2006-03-05.
Further reading
• Allaby, Michael; Peter D Moore; Trevor Day; Richard Garratt (2008), Tundra (http:/ / books. google. ca/
books?id=VdA8qT5KZvkC& lpg=PP1& dq=Tundra& pg=PP1#v=onepage& q& f=true), Facts On File,,
ISBN 0816059349
• Bliss, L. C; O. W. Heal, J. J. Moore (1981), Tundra Ecosystems: A Comparative Analysis (http:/ /
books. google.
ca/ books?id=mK48AAAAIAAJ& lpg=PP1& dq=Tundra& pg=PP1#v=onepage& q& f=true), International
Biological Programme Synthesis Series (No. 25), ISBN 9780521227766
• Warhol, Tom (2007), Tundra (http:/ / books. google. ca/ books?id=oYUc7o43cuAC& lpg=PP1&
dq=Tundra&
pg=PP1#v=onepage& q& f=true), Marshall Cavendish Benchmark, ISBN 9780761421931
• Yu I, Chernov (1998), The Living Tundra;Studies in Polar Research (http:/ / books. google. ca/
books?id=agc5AAAAIAAJ& lpg=PP1& dq=Tundra& pg=PP1#v=onepage& q& f=true), Cambridge University
Press, ISBN 9780521357548
External links
• WWF Tundra Ecoregions (http:/ / www. panda. org/ about_our_earth/ ecoregions/ about/ habitat_types/
selecting_terrestrial_ecoregions/ habitat11. cfm)
• The Arctic biome at Classroom of the Future (http:/ / www. cotf. edu/ ete/ modules/ msese/ earthsysflr/ tundra.
html)
• Arctic Feedbacks to Global Warming: Tundra Degradation in the Russian Arctic (http:/ / kaares. ulapland. fi/
home/ arktinen/ tundra/ tundra. htm)
• British Antarctica Survey (http:/ / www. antarctica. ac. uk/ about_antarctica/ wildlife/ plants/ )
Mediterranean forests, woodlands,
and scrub
Mediterranean forests, woodlands, and scrub
Eucalyptus forest.
A Mediterranean forest, the Upper Galilee
Mediterranean forests, woodlands, and
scrub is a temperate biome, characterized
by dry summers and rainy winters. Summers
are typically hot in low-lying inland
locations but can be cool near some seas, as
near San Francisco, which have a sea of
cool waters. Winters are typically mild to
cool in low-lying locations but can be cold
in inland and higher locations.
Mediterranean forests, woodlands, and scrub
eco-regions occur in the world's five
Mediterranean climate zones, on the west
coast of continents in the mid-latitudes.[1]
the 5 Mediterranean climate regions
are:
• the Mediterranean Basin
• the Chilean Matorral
• the California chaparral and woodlands
ecoregion of California and the Baja
California Peninsula
• the Cape Province-Western Cape of
South Africa
• the Southwest Australia corner area
Diversity
These regions are home to a tremendous
diversity of habitats and species. Vegetation
types can range from forests to woodlands,
savannas, shrublands, and grasslands;
"mosaic habitat" landscapes are common,
where differing vegetation types are
interleaved with one another in complex
patterns created by variations in soil,
topography, exposure to wind and sun, and
fire history. Much of the woody vegetation
in Mediterranean-climate regions is
sclerophyll, which means 'hard-leaved' in
Greek. Sclerophyll vegetation generally has
small, dark leaves covered with a waxy
outer layer to retain moisture in the dry
summer months.
All these ecoregions are highly distinctive, collectively harboring 10% of the Earth’s plant species.[2]
Phytogeographers consider the fynbos as a separate floral kingdom because 68% of the 8,600 vascular plant
species
crowded into its 90000 square kilometres (35000 sq mi) are endemic and highly distinctive at several taxonomic
levels).[1] [3]
In terms of species densities, this is equivalent to about 40% of the plant species of the United States and Canada
combined, found within an area the size of the state of Maine. The Fynbos and Southwest Australia shrublands
have
flora that are significantly more diverse than the other ecoregions, although any Mediterranean shrubland is still
rich
in species and endemics relative to other non-forest ecoregions.[1] [3]
Biome Plant Groups
Major plant communities in this biome include:
• Forest: Mediterranean forests are generally composed of broadleaf trees, such as the oak and mixed sclerophyll
forests of California and the Mediterranean region, the Eucalyptus forests of Southwest Australia, and the
Nothofagus forests of central Chile. Forests are often found in riparian areas, where they receive more
summer
water. Coniferous forests also occur. Pine and Deciduous Oak forest are widespread across California
• Woodland: Oak woodlands are characteristic of the Mediterranean Basin and in California, along with pine
woodlands and, in California, walnut woodlands.
• Savanna and grassland: The California Central Valley grasslands are the largest Mediterranean grassland
eco-region, although these grasslands have mostly been converted to agriculture. Small Woodland area occur
mainly oak, walnut, or pine woodlands.
• Shrubland: Shrublands are dense thickets of evergreen sclerophyll shrubs and small trees, called chaparral
(California), matorral (Chile and southern Spain), maquis (France and elsewhere around the Mediterranean),
macchia (Italy), fynbos (South Africa), or kwongan (Southwest Australia). In some places shrublands are the
mature vegetation type, and in other places the result of degradation of former forest or woodland by logging or
overgrazing, or disturbance by major fires.
• Scrubland: Scrublands are most common near the seacoast, and are often adapted to wind and salt air from the
ocean. Low, soft-leaved scrublands around the Mediterranean are known as garrigue in France, gariga in
Italy,
phrygana in Greece, tomillares in Spain, and batha in Israel. Northern coastal scrub and coastal sage
scrub, also
known as soft chaparral, occur near the California coast; strandveld in the Western Cape of South Africa; coastal
matorral in the central Chile, and sand-heath and kwongan in Southwest Australia.
Scrubland in Aliso Canyon, Santa Susana Mountains, Southern
California.
Fire as a medium of change
Fire, both natural and human-caused, has played a large
role in shaping the ecology of Mediterranean
ecoregions. The hot, dry summers make much of the
region prone to fires, and lightning-caused fires occur
with some frequency. Many of the plants are
pyrophytes, or fire-loving, adapted or even depending
on fire for reproduction, recycling of nutrients, and the
removal of dead or senescent vegetation. In both the
Australian and Californian Mediterranean-climate
eco-regions, native peoples used fire extensively to
clear brush and trees, making way for the grasses and
herbaceous vegetation that supported game animals and
useful plants. The plant communities in these areas adapted to the frequent human-caused fires, and pyrophyte
species grew more common and more fire-loving, while plants that were poorly adapted to fire retreated. After
European colonization of these regions, fires were suppressed, which has caused some unintended consequences
in
these ecoregions; fuel builds up, so that when fires do come they are much more devastating, and some species
dependent on fire for their reproduction are now threatened. The European shrublands have also been shaped by
anthropogenic fire, historically associated with transhumance herding of sheep and goats.
Geography
Mediterranean eco-regions are semi-arid, and often have poor soils, so they are vulnerable to degradation by
human
activities such as logging, overgrazing, and the introduction of exotic species. These regions are also some of the
most endangered on the planet, and many eco-regions have suffered tremendous degradation and habitat loss
through
logging, overgrazing, conversion to agriculture, urbanization, and introduction of exotic and invasive species.
The
eco-regions around the Mediterranean basin and in California have been particularly affected by degradation due
to
human activity, suffering extensive loss of forests and soil erosion, and many native plants and animals have
become
extinct or endangered.
References
[1] "Mediterranean Forests, Woodlands, and Scrub Ecoregions" (http:/ / www. panda. org/ about_our_earth/ ecoregions/ about/
habitat_types/
selecting_terrestrial_ecoregions/ habitat12. cfm). WWF. . Retrieved 2010-05-27. (included verbatim material: licensed under CC-BY-SA-3.0
(http:/ / creativecommons. org/ licenses/ by-sa/ 3. 0/ ))
[2] Cody, M.L. (1986). "Diversity, rarity, and conservation in Mediterranean-climate regions". In Soulé, M.E.. Conservation biology:
the science
of scarcity and diversity. Sunderland, Massachusetts, USA.: Sinauer. pp. 122–152.
[3] Cowling, R.M.; MacDonald, I.A.W.; Simmons, M.T. (1996). "The Cape Peninsula, South Africa: Physiographical, biological and
historical
background to an extraordinary hot-spot of biodiversity". Biodiversity and Conservation 5: 527–550.
External links
• National Geographic Global 200 Ecoregions: Mediterranean Forest, Woodland and Scrub. (http:/ / www.
nationalgeographic. com/ wildworld/ profiles/ g200_index. html#mediter) Part of National Geographic's Wild
World Terrestrial Ecoregions of the World (http:/ / www. nationalgeographic. com/ wildworld/ terrestrial. html)
mapping project.
Sclerophyll 46
Sclerophyll
Fynbos in South Africa
Sclerophyll is the term for a type of vegetation that has hard leaves
and short internodes (the distance between leaves along the stem). The
word comes from the Greek sclero (hard) and phyllon (leaf).
Sclerophyllous plants occur in many parts of the world,[1] but are most
typical in the chaparral biomes. They are prominent throughout
western and southern Australia, in the Mediterranean forests,
woodlands, and scrub biome that cover the Mediterranean Basin,
Californian woodlands, Chilean Matorral, and the Cape Province of
South Africa.
Australian bush
Bush around Eagle Bay, Western Australia
Most areas of the Australian continent able to support woody plants are
occupied by sclerophyll communities as forests, savannas or
heathlands. Common plants include the Proteaceae (Grevilleas,
Banksias and Hakeas), tea-trees, Acacias, Boronias, and the Eucalypts.
The most common sclerophyll communities in Australia are savannas
dominated by grasses with an overstorey of Eucalypts and Acacias.
Acacia (particularly mulga) shrublands also cover extensive areas. All
the dominant overstorey Acacia species and a majority of the
understorey Acacias have a scleromorphic adaptation in which the
leaves have been reduced to phyllodes consisting entirely of the
petiole.[2] Many plants of the sclerophyllous woodlands and shrublands
also produce leaves unpalatable to herbivores by the inclusion of toxic
and indigestible compounds in an attempt to maintain these long-lived
leaves. This trait is particularly noticeable in the eucalypt and
Melaleuca species which possess oil glands within their leaves that
produce a pungent volatile oil that makes them unpalatable to most browsers.[3] These traits make the majority of
woody plants in these woodlands largely unpalatable to domestic livestock.[4] It is therefore important from a
grazing
perspective that these woodlands support a more or less continuous layer of herbaceous ground cover dominated
by
grasses.
Sclerophyll forests cover a much smaller area of the continent, being restricted to relatively high rainfall
locations.
They have a eucalyptus overstory (10 to 30 metres) with the understory also being hard-leaved. Dry
sclerophyll
forests are the most common forest type on the continent, and although it may seem barren dry sclerophyll forest
is
highly diverse. For example, a study of sclerophyll vegetation in Seal Creek, Victoria, found 138 species.[5]
Even less extensive are wet sclerophyll forests. They have a taller eucalyptus overstory than dry sclerophyll
forests,
30 metres or more (typically Mountain Ash, Alpine Ash, Messmate Stringybark or Manna Gum), and a soft-
leaved,
fairly dense understory (tree ferns are common). They require ample rainfall — at least 1000mm (40 inches).
Sclerophyll 47
History
Sclerophyllous plants are all part of a specific environment and are anything but newcomers. By the time of
European settlement, sclerophyll forest accounted for the vast bulk of the forested areas.
Most of the wooded parts of present-day Australia have become sclerophyll dominated as a result of the extreme
age
of the continent combined with Aboriginal fire use. Deep weathering of the crust over many millions of years
leached chemicals out of the rock, leaving Australian soils deficient in nutrients, particularly phosphorus. Such
nutrient deficient soils support non-sclerophyllous plant communities elsewhere in the world and did so over
most of
Australia prior to human arrival. However such deficient soils cannot support the nutrient losses associated with
frequent fires and are rapidly replaced with sclerophyllous species under traditional Aboriginal burning regimes.
With the cessation of traditional burning non-sclerophyllous species have re-colonised sclerophyll habitat in
many
parts of Australia. The presence of toxic compounds combined with a low carbon : nitrogen ratio make the
leaves
and branches of scleromorphic species long-lived in the litter, and can lead to a large build-up of litter in
woodlands.[6] [7] The toxic compounds of many species, notably Eucalyptus species, are volatile and flammable
and
the presence of large amounts of flammable litter, coupled with an herbaceous understorey, encourages fire.[8]
All
the Australian sclerophyllous communities are liable to be burnt with varying frequencies and many of the
woody
plants of these woodlands have developed adaptations to survive and minimise the effects of fire.[9]
Sclerophyllous plants generally resist dry conditions well, making them successful in areas of seasonally
variable
rainfall. In Australia, however, they evolved in response to the low level of phosphorus in the soil — indeed,
many
Australian native plants cannot tolerate higher levels of phosphorus and will die if fertilised incorrectly. The
leaves
are hard due to lignin, which prevents wilting and allows plants to grow even when there isn't enough
phosphorus for
substantial new cell growth.[10]
References
[1] C. Michael Hogan. 2010. Leather Oak, Quercus durata. Encyclopedia of Earth. National Council for Science and Environment
(http:/ / www.
eoearth. org/ article/ Leather_Oak). Washington DC
[2] Simmons, M. 1982. Acacias of Australia. Thomas Nelson, Melbourne
[3] Florence, R. G. 1996. Ecology and silviculture of eucalypt forests. CSIRO Publishing, Collingwood.
[4] Mott, J. J., and J. C. Tothill. 1984. Tropical and subtropical woodlands. in G. N. Harrington and A. D. Wilson, editors. Management of
Australia's Rangelands. CSIRO Publishing, Melbourne
[5] Parsons R. F.; Cameron D .G. (1974). "Maximum Plant Species Diversity in Terrestrial Communities". Biotropica (The Association for
Tropical Biology and Conservation) 6 (3): 202. doi:10.2307/2989653. JSTOR 2989653.
[6] White, M. E. 1986. The Greening of Gondwana. Reed Books, Frenchs Forest, Australia.
[7] Flannery, T. 1994. The future eaters. Reed New Holland, Frenchs Forest, Australia.
[8] Burrows, W. H., J. O. Carter, J. C. Scanlan, and E. R. Anderson. 1990. Management of savannas for livestock production in north-east
Australia: contrasts across the tree-grass continuum. Journal of Biogeography 17:503-512.
[9] Harrington, G. N., M. H. Friedel, K. C. Hodgkinson, and J. C. Noble. 1984. Vegetation ecology and management. in G. N. Harrington
and A.
D. Wilson, editors. Management of Australia's Rangelands. CSIRO Publishing, Melbourne.
[10] R. Major (2003). "Sclerophyll forests" (http:/ / www. amonline. net. au/ factsheets/ sclerophyll_forests. htm). Australian Museum. .
Retrieved 2005-02-14.
Deserts and xeric shrublands

Deserts and xeric shrublands is a biome characterized by, relating to, or requiring only a small
amount of
moisture.[1]
Definition and occurrence
Deserts and xeric shrublands receive an annual average rainfall of ten inches or less, and have an arid or
hyperarid
climate, characterized by a strong moisture deficit, where annual potential loss of moisture from
evapotranspiration
well exceeds the moisture received as rainfall. Deserts and xeric shrublands occur in tropical, subtropical, and
temperate climate regions. Desert soils tend to be sandy or rocky, and low in organic materials. Saline or alkaline
soils are common. Plants and animals in deserts and xeric shrublands are adapted to low moisture conditions.
Hyperarid regions are mostly devoid of vegetation and animal life, and include rocky deserts and sand dunes.
Vegetation in arid climate regions can include sparse grasslands, shrublands, and woodlands. Plants adapted to
arid
climates are called xerophytes, and include succulent plants, geophytes, sclerophyll, and annual plants. Animals,
including insects, reptiles, arachnids, birds and mammals, are frequently nocturnal to avoid moisture loss.
In isolation, Hawai'i's Silverswords have adapted
to xeric microclimates within volcanic craters,
trapping and channeling dew and protecting
leaves with reflective hairs.
Desertification
The conversion of productive drylands to desert conditions is known as
desertification, and can occur from a variety of causes. One factor is
human intervention in imposing intensive agricultural tillage or
overgrazing[2] in areas which cannot support such exploitation.
Climatic shifts such as global warming or the Milankovitch cycle
(which drives glacials and interglacials) also affect the pattern of
deserts on Earth.
Desert and xeric shrublands ecoregions
Afrotropic Deserts and xeric shrublands
Aldabra Island xeric scrub Seychelles
Arabian Peninsula coastal fog desert Oman, Saudi Arabia, Yemen
East Saharan montane xeric woodlands Chad, Sudan
Eritrean coastal desert Djibouti, Eritrea
Ethiopian xeric grasslands and shrublands Djibouti, Ethiopia, Somalia, Sudan
Gulf of Oman desert and semi-desert Oman, United Arab Emirates
Hobyo grasslands and shrublands Somalia
Ile Europa and Bassas da India xeric scrub Bassas da India, Europa
Kalahari xeric savanna Botswana, Namibia, South Africa
Kaokoveld desert Angola, Namibia
Madagascar spiny thickets Madagascar
Madagascar succulent woodlands Madagascar
Masai xeric grasslands and shrublands Ethiopia, Kenya
Nama Karoo Namibia, South Africa
Namib desert Namibia
Namibian savanna woodlands Namibia
Red Sea coastal desert Egypt, Sudan
Socotra Island xeric shrublands Yemen
Somali montane xeric woodlands Somalia
Southwestern Arabian foothills savanna Oman, Saudi Arabia, Yemen
Southwestern Arabian montane woodlands Saudi Arabia, Yemen
Succulent Karoo South Africa
Australasia Deserts and xeric shrublands
Carnarvon xeric shrublands Australia
Central Ranges xeric scrub Australia
Gibson Desert Australia
Great Sandy-Tanami Desert Australia
Great Victoria Desert Australia
Nullarbor Plain xeric shrublands Australia
Pilbara shrublands Australia
Simpson Desert Australia
Tirari-Sturt's Stony Desert Australia
Western Australian mulga shrublands Australia
Indomalaya Deserts and xeric shrublands
Deccan thorn scrub forests India, Sri Lanka
Indus Valley desert India, Pakistan
Northwestern thorn scrub forests India, Pakistan
Thar desert India, Pakistan
Nearctic Deserts and xeric shrublands
Baja California desert Mexico
Central Mexican matorral Mexico
Chihuahuan desert Mexico, United States
Colorado Plateau shrublands United States
Great Basin shrub steppe United States
Gulf of California xeric scrub Mexico
Meseta Central matorral Mexico
Mojave desert United States
Okanagan (South) shrub steppe Canada
Snake-Columbia shrub steppe United States
Sonoran desert Mexico, United States
Tamaulipan matorral Mexico
Tamaulipan mezquital Mexico
Wyoming Basin shrub steppe United States
Neotropic Deserts and xeric shrublands
Araya and Paria xeric scrub Venezuela
Aruba-Curaçao-Bonaire cactus
scrub
Aruba, Bonaire, Curaçao
Atacama desert Chile, Peru
Caatinga Brazil
Cayman Islands xeric scrub Cayman Islands
Cuban cactus scrub Cuba
Galápagos Islands xeric scrub Ecuador
Guajira-Barranquilla xeric scrub Colombia, Venezuela
La Costa xeric shrublands Venezuela
Leeward Islands xeric scrub Anguilla, Antigua and Barbuda, British Virgin Islands, Guadeloupe, Saint Martin, Saint Barthélemy, Saba, US
Virgin Islands
Malpelo Island xeric scrub Colombia
Motagua Valley thornscrub Guatemala
Paraguana xeric scrub Venezuela
San Lucan xeric scrub Mexico
Sechura desert Peru
Tehuacán Valley matorral Mexico
Windward Islands xeric scrub Barbados, Dominica, Grenada, Martinique, Saint Lucia, Saint Vincent and the Grenadines
Saint Peter and Saint Paul rocks Brazil
• this is because it gets only ten inches of precipitation a year
Palearctic Deserts and xeric shrublands
Afghan Mountains semi-desert Afghanistan
Alashan Plateau semi-desert China, Mongolia
Arabian Desert and East Sahero-Arabian xeric shrublands Egypt, Jordan, Oman, Saudi Arabia, Syria, United Arab Emirates, Yemen
Atlantic coastal desert Mauritania, Western Sahara
Azerbaijan shrub desert and steppe Azerbaijan, Georgia, Iran
Badkhiz-Karabil semi-desert Afghanistan, Iran, Tajikistan, Turkmenistan, Uzbekistan
Baluchistan xeric woodlands Afghanistan, Pakistan
Caspian lowland desert Iran, Kazakhstan, Russia, Turkmenistan
Central Afghan Mountains xeric woodlands Afghanistan
Central Asian northern desert Kazakhstan, Uzbekistan
Central Asian riparian woodlands Kazakhstan
Central Asian southern desert Kazakhstan, Turkmenistan, Uzbekistan
Central Persian desert basins Afghanistan, Iran
Eastern Gobi desert steppe China, Mongolia
Gobi Lakes Valley desert steppe Mongolia
Great Lakes Basin desert steppe Mongolia, Russia
Junggar Basin semi-desert China, Mongolia
Kazakh semi-desert Kazakhstan
Kopet Dag semi-desert Iran, Turkmenistan
Mesopotamian shrub desert Iraq, Jordan, Syria
North Saharan steppe and woodlands Algeria, Egypt, Libya, Morocco, Tunisia, Western Sahara
Paropamisus xeric woodlands Afghanistan
Persian Gulf desert and semi-desert Bahrain, Kuwait, Qatar, Saudi Arabia, United Arab Emirates
Qaidam Basin semi-desert China
Red Sea Nubo-Sindian tropical desert and semi-desert Iraq, Jordan, Oman, Saudi Arabia, Yemen
Rigestan-North Pakistan sandy desert Afghanistan, Iran, Pakistan
Sahara desert Algeria, Chad, Egypt, Libya, Mali, Mauritania, Niger, Sudan
South Iran Nubo-Sindian desert and semi-desert Iran, Iraq, Pakistan
South Saharan steppe and woodlands Algeria, Chad, Mali, Mauritania, Niger, Sudan
Taklimakan desert China
Tibesti-Jebel Uweinat montane xeric woodlands Chad, Egypt, Libya, Sudan
West Saharan montane xeric woodlands Algeria, Mali, Mauritania, Niger
References
[1] Merriam-Webster.com (http:/ / www. merriam-webster. com/ dictionary/ xeric) Merriam-Webster Dictionary definition of xeric
[2] C.Michael Hogan. 2009. Overgrazing (http:/ / www. eoearth. org/ article/ Overgrazing). Encyclopedia of Earth. Sidney Draggan, topic
ed.;
Cutler J. Cleveland, ed., National council for Science and the Environment, Washington DC
External links
• Deserts and xeric shrublands (World Wildlife Fund) (http:/ / www. panda. org/ news_facts/ education/
university/
habitats/ index. cfm?habitatID=13)
• Map of the ecozones (http:/ / www. panda. org/ about_wwf/ where_we_work/ ecoregions/ maps/ index. cfm)
• Index to Deserts & Xeric Shrublands at bioimages.vanderbilt.edu (http:/ / www. cas. vanderbilt. edu/
bioimages/
ecoframe-list. htm)
• Xeric World (http:/ / www. xericworld. com/ forums/ ) Online Community focused on the study of Xeric Plant
Species.
Mangrove
World mangrove distribution
Pneumatophores penetrate the sand surrounding a
mangrove tree.
Short video of a mangrove in Brasil
Mangroves are various kinds of trees up to medium height and shrubs
that grow in saline coastal sediment habitats in the tropics and
subtropics – mainly between latitudes 25° N and 25° S. Such bosks are
also part of the mangrove forest biome. The saline conditions
tolerated by various species range from brackish water, through pure
seawater (30 to 40 ppt), to water concentrated by evaporation to over
twice the salinity of ocean seawater (up to 90 ppt).[1] [2]
There are many species of trees and shrubs adapted to saline
conditions. Not all are closely related, and the term "mangrove" may
be used for all of them, or more narrowly only for the mangrove family
of plants, the Rhizophoraceae, or even more specifically just for
mangrove trees of the genus Rhizophora.
Mangroves form a distinct characteristic saline woodland or shrubland
habitat, called mangrove swamp, mangrove forest, mangrove or
mangal.[3] Mangals are found in depositional coastal environments,
where fine sediments (often with high organic content) collect in areas
protected from high-energy wave action. Mangroves dominate three
quarters of tropical coastlines.[2]
Ecology
Mangroves are found in tropical and subtropical tidal areas. Areas
where mangals occur include estuaries and marine shorelines.[1]
The intertidal existence to which these trees are adapted repesents the
major limitation to the number of species able to thrive in their habitat.
High tide brings in salt water, and when the tide recedes, solar
evaporation of the seawater in the soil leads to further increases in
Mangrove 53
salinity. The return of tide can flush out these soils, bringing them back to salinity levels comparable to that of
seawater. At low tide, organisms are also exposed to increases in temperature and desiccation, and are then
cooled
and flooded by the tide. Thus, in order for a plant to survive in this environment, it must tolerate broad ranges of
salinity, temperature, and moisture, as well as a number of other key environmental factors. It is unsurprising,
perhaps, that only a select few species make up the mangrove tree community.
About 110 species are considered mangroves.[1] However, a given mangrove typically features only a small
number
of tree species. It is not uncommon for a mangrove forest in the Caribbean to feature only three or four tree
species.
For comparison, the tropical rainforest biome contains thousands of tree species. That is not to say that
mangrove
forests lack diversity. Though the trees themselves are few in species, the ecosystem that these trees create
provides
a home for a great variety of other organisms.
Mangroves require a number of physiological adaptations to overcome the problems of anoxia, high salinity and
frequent tidal inundation. Each species has its own solutions to these problems; this may be the primary reason
why,
on some shorelines, mangrove tree species show distinct zonation. Small environmental variations within a
mangal
may lead to greatly differing methods for coping with the environment. Therefore, the mix of species is partly
determined by the tolerances of individual species to physical conditions, like tidal inundation and salinity, but
may
also be influenced by other factors such as predation of plant seedlings by crabs.
Once established, mangrove roots provide an oyster habitat and slow water flow, thereby enhancing sediment
deposition in areas where it is already occurring. The fine, anoxic sediments under mangroves act as sinks for a
variety of heavy (trace) metals which colloidal particles in the sediments scavenged from the water. Mangrove
removal disturbs these underlying sediments, often creating problems of trace metal contamination of seawater
and
biota.
Mangroves protect coastal areas from erosion, storm surge (especially during hurricanes), and tsunamis.[4] [5]
The
mangroves' massive root systems are efficient at dissipating wave energy.[6] Likewise, they slow down tidal
water
enough that its sediment is deposited as the tide comes in, leaving all except fine particles when the tide ebbs.[7]
In
this way, mangroves build their own environment.[4] Because of the uniqueness of mangrove ecosystems and the
protection against erosion they provide, they are often the object of conservation programs, including national
biodiversity action plans.[5]
However, mangroves' protective value is sometimes overstated. Wave energy is typically low in areas where
mangroves grow,[8] so their effect on erosion can only be measured over long periods.[6] Their capacity to limit
high-energy wave erosion is limited to events such as storm surges and tsunamis.[9] Erosion often occurs on the
outer
sides of bends in river channels that wind through mangroves, while new stands of mangroves are appearing on
the
inner sides where sediment is accruing.
The unique ecosystem found in the intricate mesh of mangrove roots offers a quiet marine region for young
organisms. In areas where roots are permanently submerged, the organisms they host include algae, barnacles,
oysters, sponges, and bryozoans, which all require a hard surface for anchoring while they filter feed. Shrimps
and
mud lobsters use the muddy bottoms as their home.[10] Mangrove crabs mulch the mangrove leaves, adding
nutritients to the mangal muds for other bottom feeders.[11] In at least some cases, export of carbon fixed in
mangroves is important in coastal food webs.
Mangrove plantations in Vietnam, Thailand, the Philippines and India host several commercially important
species
of fish and crustaceans. Despite restoration efforts, developers and others have removed over half of the world's
mangroves in recent times.
Mangrove 54
Biology
Of the recognized 110 mangrove species, only about 54 species in 20 genera from 16 families constitute the
"true
mangroves", species that occur almost exclusively in mangrove habitats.[3] Demonstrating convergent evolution,
many of these species found similar solutions to the tropical conditions of variable salinity, tidal range
(inundation),
anaerobic soils and intense sunlight. Plant biodiversity is generally low in a given mangal.[1] This is especially
true
in higher latitudes and in the Americas. The greatest biodiversity occurs in the mangal of New Guinea, Indonesia
and
Malaysia.[12]
Adaptations to low oxygen
A red mangrove, Rhizophora mangle
Above and below water view at the edge of the
mangal
Red mangroves, which can survive in the most inundated areas, prop
themselves above the water level with stilt roots and can then absorb
air through pores in their bark (lenticels). Black mangroves live on
higher ground and make many pneumatophores (specialised root-like
structures which stick up out of the soil like straws for breathing)
which are also covered in lenticels. These "breathing tubes" typically
reach heights of up to thirty centimeters, and in some species, over
three meters. There are four types of pneumatophore—stilt or prop
type, snorkel or peg type, knee type, and ribbon or plank type. Knee
and ribbon types may be combined with buttress roots at the base of
the tree. The roots also contain wide aerenchyma to facilitate transport
within the plant.
Mangrove 55
Salt crystals formed on grey mangrove leaf
Limiting salt intake
Red mangroves exclude salt by having significantly impermeable roots
which are highly suberised, acting as an ultrafiltration mechanism to
exclude sodium salts from the rest of the plant. Analysis of water
inside mangroves has shown 90% to 97% of salt has been excluded at
the roots. Salt which does accumulate in the shoot concentrates in old
leaves which the plant then sheds. Red mangroves can also store salt in
cell vacuoles. White (or grey) mangroves can secrete salts directly;
they have two salt glands at each leaf base (hence their name—they are
covered in white salt crystals).
Limiting water loss
Because of the limited fresh water available in salty intertidal soils,
mangroves limit the amount of water they lose through their leaves.
They can restrict the opening of their stomata (pores on the leaf
surfaces, which exchange carbon dioxide gas and water vapour during
photosynthesis). They also vary the orientation of their leaves to avoid
the harsh midday sun and so reduce evaporation from the leaves.
Anthony Calfo, a noted aquarium author, observed anecdotally a red
mangrove in captivity only grows if its leaves are misted with fresh water several times a week, simulating the
frequent tropical rainstorms.[13]
Nutrient uptake
The biggest problem that mangroves face is nutrient uptake. Because the soil is perpetually waterlogged, there is
little free oxygen. Anaerobic bacteria liberate nitrogen gas, soluble iron, inorganic phosphates, sulfides, and
methane, which makes the soil much less nutritious and contributes to mangroves' pungent odor.
Pnuematophores
(aerial roots) allow mangroves to absorb gases directly from the atmosphere, and other nutrients such as iron,
from
the inhospitable soil. Mangroves store gases directly inside the roots, processing them even when the roots are
submerged during high tide.
Increasing survival of offspring
Red mangrove seeds germinate while still on the
parent tree
In this harsh environment, mangroves have evolved a special
mechanism to help their offspring survive. Mangrove seeds are
buoyant and therefore suited to water dispersal. Unlike most plants,
whose seeds germinate in soil, many mangroves (e.g. red mangrove)
are viviparous, whose seeds germinate while still attached to the parent
tree. Once germinated, the seedling grows either within the fruit (e.g.
Aegialitis, Avicennia and Aegiceras), or out through the fruit (e.g.
Rhizophora, Ceriops, Bruguiera and Nypa) to form a propagule (a
ready-to-go seedling) which can produce its own food via
photosynthesis. The mature propagule then drops into the water, which
can transport it great distances. Propagules can survive desiccation and
Mangrove 56
remain dormant for over a year before arriving in a suitable environment. Once a propagule is ready to root, its
density changes so the elongated shape now floats vertically rather than horizontally. In this position, it is more
likely to lodge in the mud and root. If it does not root, it can alter its density and drift again in search of more
favorable conditions.
Taxonomy
The following listing (modified from Tomlinson, 1986) gives the number of species of mangroves in each listed
plant genus and family.
Major components
Family Genus, number of species Common name
Acanthaceae, Avicenniaceae or
Verbenaceae
(family allocation disputed)
Avicennia, 9 Black mangrove
Combretaceae Conocarpus, 1; Laguncularia, 11; Lumnitzera, 2 Buttonwood, white mangrove
Arecaceae Nypa, 1 Mangrove palm
Rhizophoraceae Bruguiera, 6; Ceriops, 2; Kandelia, 1; Rhizophora, 8 Red mangrove
Lythraceae Sonneratia, 5 Mangrove apple
Minor components
Family Genus, number of species
Acanthaceae Acanthus, 1; Bravaisia, 2
Bombacaceae Camptostemon, 2
Cyperaceae Fimbristylis, 1
Euphorbiaceae Excoecaria, 2
Lecythidaceae Barringtonia, 6
Lythraceae Pemphis, 1
Meliaceae Xylocarpus, 2
Myrsinaceae Aegiceras, 2
Myrtaceae Osbornia, 1
Pellicieraceae Pelliciera, 1
Plumbaginaceae Aegialitis, 2
Pteridaceae Acrostichum, 3
Rubiaceae Scyphiphora, 1
Sterculiaceae Heritiera, 3
Mangrove 57
Geographical regions
Mangroves occur in numerous areas worldwide. See List of mangrove ecoregions.
Africa
There are important mangrove swamps in Kenya, Tanzania and Madagascar, with the latter even admixing at the
coastal verge with dry deciduous forests. The delta of the Rufiji River is reportedly home to the largest
mangrove
forest in the world.
Nigeria has Africa's largest mangrove concentration, spanning 36,000 km2. Oil spills and leaks have destroyed
many
in the last fifty years, damaging the local fishing economy and water quality.[14]
Along the coast of the Red Sea, both on the Egyptian side and in the Gulf of Aqaba, mangroves composed
primarily
of Avicennia marina and Rhyzophora mucronata in about 28 stands cover about 525 hectares. Almost
all Egyption
mangrove stands are now protected..
Americas
Mangroves live in many parts of the tropical and subtropical coastal zones of North, South and Central America.
Continental United States
Because of their sensitivity to subfreezing temperatures, mangroves in the continental United States are limited
to the
Florida peninsula (see Florida mangroves) and isolated growths[15] of black mangrove (Avicennia
germinans) along
the coast of southern Louisiana[16] and south Texas[17]
Central America and Caribbean
Mangroves occur on the west coast of Costa Rica, on the Pacific and Caribbean coasts of Nicaragua, Belize,
Guatemala, Honduras, and Panama, and on many Caribbean Islands, such as Curaçao, Bonaire, Antigua,
Anguilla,
the Bahamas, Saint Kitts and Nevis and St. Lucia. Significant mangals include the Marismas Nacionales-San
Blas
mangroves in Mexico. Mangroves can also be found in Puerto Rico, Cuba, the Dominican Republic, Haiti,
Jamaica,
Trinidad, Barbados, and the Pacific coast of El Salvador.
Belize
The nation of Belize boasts the highest overall percentage of forest cover of any of the Central American
countries.[18] In terms of Belize's mangrove cover - which assumes the form not only of mangrove 'forest' but
also of
scrubs and savannas, among others[19] - a 2010 satellite-based study [20] of Belize's mangroves by the World
Wildlife
Fund (WWF) [21] and the Water Center for the Humid Tropics of Latin America and the Caribbean
(CATHALAC)
[22] found that in 2010, mangroves covered some 184,548 acres (74,684 hectares) or 3.4% of Belize's territory. In
1980, by contrast, mangrove cover stood at 188,417 acres (76,250 hectares) - also 3.4% of Belize's territory,
although based on the work of mangrove researcher Simon Zisman,[23] it is estimated Belize's mangrove cover in
1980 represented 98.7% of the precolonial extent of those ecosystems. Belize's mangrove cover in 2010 was thus
estimated to represent 96.7% of the precolonial cover.[24] Assessing changes in Belize's mangrove cover over a
30-year period was possible because of Belize's participation in the Regional Visualization & Monitoring
System [25]
(SERVIR [25], in Spanish) a regional observatory jointly implemented by CATHALAC [22], RCMRD [26],
ICIMOD
[27], NASA [28], USAID [29], and other partners.[30]
Mangrove 58
South America
Mangrove near the town of Cienaga, Magdalena,
in the Ciénaga Grande de Santa Marta swampy
marshes, Colombia.
Brazil contains approximately 26,000 km2 of mangals, 15% of the
world's total of 172,000 km2.
Ecuador and Peru have significant areas of mangroves, mainly in the
Gulf of Guayaquil-Tumbes mangroves.
Venezuela's northern Caribbean island, Margarita, possesses mangrove
forests in the Parque Nacional la Restinga.
Colombia possesses large mangrove forests on both its Caribbean and
Pacific coasts.
Asia
Indomalaya ecozone
Mangroves occur on Asia's south coast, throughout the Indian subcontinent, in all southeast Asian countries, and
on
islands in the Indian Ocean, Arabian Sea, Bay of Bengal, South China Sea and the Pacific.
The mangal is particularly prevalent in the deltas of large Asian rivers. The Sundarbans is the largest mangrove
forest in the world, located in the Ganges river delta in Bangladesh and West Bengal, India.
A view of the Pichavaram mangrove forest from the viewing tower
The Pichavaram Mangrove Forest near
Chidambaram, South India, by the Bay
of Bengal, is the world's second largest
mangrove forest.
Major mangals live on the Andaman
and Nicobar Islands and the Gulf of
Kutch in Gujarat.[31]
Other significant mangals include the
Bhitarkanika Mangroves and
Godavari-Krishna mangroves.
In Vietnam, mangrove forests grow along the southern coast, including two forests: the Can Gio Mangrove
Forest
biosphere reserve and the U Minh mangrove forest in the Sea and Coastal Region of Kien Giang, Ca Mau and
Bac
Lieu province.
The mangrove forests of Kompong Sammaki in Cambodia are of major ecological and cultural importance, as
the
human population relies heavily on the crabs and fish that live in the roots.
The three most important mangrove forests of Taiwan are: Tamsui River in Taipei, Jhonggang River in Miaoli
and
the Sihcao Wetlands in Tainan. According to research, there are four existing types of mangrove in Taiwan.
Some
places have been developed as scenic areas, such as the log raft routes in Sihcao.
In the Indonesian Archipelago, mangroves occur around much of Sumatra, Borneo, Sulawesi and the
surrounding
islands, while further north, they are found along the coast of the Malay Peninsula.
Mangrove 59
A cluster of mangroves on
the banks of the Vellikeel
River in Kannur
An old light house in the middle
of mangroves on the banks of the
Godavari River in Andhra
Pradesh
Mangroves in Mangalavanam
Bird Sanctuary of Kerala
The green tunnel of
mangrove in Sihcao, Tainan,
Taiwan.
A mangrove of the genus
Sonneratia, showing
abundant pneumatophores
growing on the landward
margin of the reef flat on
Yap.
The location and relative density
of mangroves in Southeast Asia
and Australasia.
Pakistan
Pakistani mangroves are located mainly along the Indus delta (the Indus River Delta-Arabian Sea mangroves
ecoregion). Major mangrove forests are found on the coastline of the provinces of Sindh and Balochistan. In
Karachi, land reclamation projects have led to the cutting down of mangrove forests for commercial and urban
development.
Middle East
Oman, near Muscat, supports large areas of mangroves, in particular at Shinas, Qurm Park and Mahout Island. In
Arabic, mangrove trees are known as qurm, thus the mangrove area in Oman is known as Qurm Park.
Mangroves are
also present extensively in neighboring Yemen.[32]
Iranian mangrove forests occur between 25°11′N to 27°52′N. These forests exist in the north part of the Persian
Gulf
and Oman Sea, along three maritime provinces in the south of Iran. These provinces respectively from southwest
to
southeast of Iran, include Bushehr, Hormozgan and Sistan & Balouchestan.
Australia
More than fifty species of Rhizophoraceae grow in Australasia[33] with particularly high biodiversity on the
island of
New Guinea and northern Australia.[33]
Australia has approximately 11,500 km2 of mangroves, primarily on the northern and eastern coasts of the
continent,
with occurrences as far south as Millers Landing in Wilsons Promontory, Victoria[34] (38°54′S)[35] and Barker
Inlet
in Adelaide, South Australia.[36]
Mangrove 60
New Zealand
New Zealand also has mangrove forests extending to around 38°S (similar to Australia's southernmost mangrove
incidence): the furthest geographical extent on the west coast is Raglan Harbour (37°48′S); on the east coast,
Ohiwa
Harbour (near Opotiki) is the furthest south mangroves are found (38°00′S).[37]
Pacific islands
Twenty-five species of mangrove are found on various Pacific islands, with extensive mangals on some islands.
Mangals on Guam, Palau, Kosrae and Yap have been badly affected by development.[38]
Mangroves are not native to Hawaii, but the red mangrove, Rhizophora mangle, and Oriental mangrove,
Bruguiera
sexangula, have been introduced and are now naturalized.[39] Both species are classified as pests by the
University of
Hawaii Botany Department.[40]
Cultivating mangroves
Red mangroves are the most common choice for cultivation, used particularly in marine aquariums in a sump to
reduce nitrates and other nutrients in the water. Mangroves also appear in home aquariums, and as ornamental
plants,
such as in Japan.
The Manzanar Mangrove Initiative is an ongoing experiment in Arkiko, Eritrea, part of the Manzanar Project
founded by Dr Gordon H. Sato, establishing new mangrove plantations on the coastal mudflats. Initial plantings
failed, but observation of the areas where mangroves did survive by themselves led to the conclusion that
nutrients in
water flow from inland were important to the health of the mangroves. Trials with the Eritrean Ministry of
Fisheries
followed, and a planting system was designed to introducing the nitrogen, phosphorus, and iron missing from
seawater. The propagules are planted inside a reused galvanized steel can with the bottom knocked out; a small
piece
of iron and a pierced plastic bag with fertilizer containing nitrogen and phosphorus are buried with the
propagule. As
at 2007, after six years of planting, there are 700,000 mangroves growing; providing stock feed for sheep and
habitat
for oysters, crabs, seashells and fish.[41] [42]
Exploitation and conservation
Mangroves in West Bali National Park, Indonesia
Approximately 35% of mangrove area was lost during the last several
decades of the twentieth century (in countries for which sufficient data
exist), which encompass about half of the area of mangroves.[43] The
United Nations Environment Program also estimated shrimp farming
causes a quarter of the destruction of mangrove forests.[44] Likewise,
the 2010 update of the World Mangrove Atlas [45] (WMA) indicated a
fifth of the world's mangrove ecosystems have been lost since 1980.[46]
Grassroots efforts to save mangroves from development are becoming
more popular as the benefits of mangroves become more widely
known. In the Bahamas, for example, active efforts to save mangroves are occurring on the islands of Bimini and
Great Guana Cay. In Trinidad and Tobago as well, efforts are underway to protect a mangrove threatened by the
construction of a steelmill and a port.[47] In Thailand, community management has been effective in restoring
damaged mangroves.[48]
It has been cited that mangroves can help buffer against Tsunami, cyclones, and other storms. One village in
Tamil
Nadu was protected from tsunami destruction - the villagers in Naluvedapathy planted 80,244 saplings to get
into the
Guinness Book of World Records. This created a kilometre-wide belt of trees of various varieties. When the
tsunami
struck, much of the land around the village was flooded, but the village itself suffered minimal damage.[49] [50]
Mangrove 61
National studies
In terms of local and national studies of mangrove loss, the case of Belize's mangroves is illustrative in its
contrast to
the global picture. A recent, satellite-based study[24] - funded by the World Wildlife Fund (WWF) [21] and
conducted
by the Water Center for the Humid Tropics of Latin America and the Caribbean (CATHALAC) [22] – indicates
Belize's mangrove cover declined by a mere 2% over a thirty-year period. The study was borne out of the need to
verify the popular conception that mangrove clearing in Belize was rampant.[51] Instead, the assessment showed
that
between 1980 and 2010, under 4,000 acres of mangroves had been cleared, although clearing of mangroves near
Belize's main coastal settlements (e.g. Belize City and San Pedro) was relatively high. The rate of loss of Belize's
mangroves - at 0.07% per year between 1980 and 2010 - was much lower than Belize's overall rate of forest
clearing
(0.6% per year between 1980 and 2010).[52] These findings can also be interpreted to indicate Belize's mangrove
regulations (under the nation's [53] ) have largely been effective. Nevertheless, the need to protect Belize's
mangroves
is imperative, as a 2009 study by the World Resources Institute (WRI) [54] indicates the ecosystems contribute
US$174–249 million per year to Belize's national economy.[55]
In popular media
• The mangrove is used as a symbol in Annie Dillard's essay Sojourner due to its significance as a self-
sustaining
biome.
• The manga series One Piece features a forest of giant mangroves that form the Sabaody Archipelago. The
mangroves produce a resin that combines with the oxygen exhaled by the trees to create large bubbles. The local
population uses the bubbles for everything from transport to hotels.
Notes
[1] Mangal (Mangrove). World Vegetation. Mildred E. Mathias Botanical Garden, University of California at Los Angeles (http:/ / www.
botgard. ucla. edu/ html/ botanytextbooks/ worldvegetation/ marinewetlands/ mangal/ index. html)
[2] Morphological and Physiological Adaptations: Florida mangrove website (http:/ / www. nhmi. org/ mangroves/ phy. htm)
[3] Hogarth, Peter J. (1999) The Biology of Mangroves Oxford University Press, Oxford, England, What page? ISBN 0-19-850222-2
[4] Mazda, Y.; Kobashi, D. and Okada, S. (2005) "Tidal-Scale Hydrodynamics within Mangrove Swamps" Wetlands Ecology and
Management
13(6): pp. 647-655
[5] Danielsen, F. et al. (2005) "The Asian tsunami: a protective role for coastal vegetation" Science 310: p. 643.
[6] Massel, S. R.; Furukawa, K.and Brinkman R. M. (1999) "Surface wave propagation in mangrove forests" Fluid Dynamics Research
24(4):
pp. 219–249
[7] Mazda, Yoshihiro et al. (1997) "Drag force due to vegetation in mangrove swamps" Mangroves and Salt Marshes 1: pp. 193–199
[8] Baird, Andrew (26 December 2006) "False Hopes and Natural Disasters" New York Times editorial
[9] Dahdouh-Guebas, F. et al. (2005) "How effective were mangroves as a defence against the recent tsunami?" Current Biology 15(12):
pp.
443–447
[10] Encarta Encyclopedia 2005. Article — Seashore, by Heidi Nepf.
[11] Skov, Martin W. and Hartnoll, Richard G. (March 2002). Paradoxical selective feeding on a low-nutrient diet: why do mangrove crabs
eat
leaves? Oecologia 131(1): pp. 1–7.
[12] UN Report on mangrove diversity (http:/ / maps. grida. no/ go/ graphic/ distribution_of_coral_mangrove_and_seagrass_diversity/ )
[13] Calfo, Anthony (2006). Mangroves for the Marine Aquarium. (http:/ / www. reefkeeping. com/ issues/ 2004-12/ ac/ feature/
index. php)
[14] O'Neill.T (February 2007). "Curse of the Black Gold". National Geographic: 88 to 117.
[15] "Modeling Hurricane Effects on Mangrove Ecosystems" U.S. Geological Survey, USGS FS-095-97, June 1997 (http:/ / www. nwrc.
usgs.
gov/ climate/ fs95_97. pdf)
[16] "Coastal Mangrove-Marsh Shrubland" (http:/ / www. wlf. state. la. us/ pdfs/ experience/ Coastal Mangrove-Marsh Shrubland. pdf)
(PDF).
Conservation Habitats & Species Assessments. Louisiana Department of Wildlife & Fisheries. December 2005. .
[17] Yang, Chenghai; Everitt, James; Fletcher, Reginald; Jensen, Ryan;Mausel, Paul (2008-03-15). "Mapping Black Mangrove Along the
South
Texas Gulf Coast Using AISA+ Hyperspectral Imagery" (http:/ / www. ars. usda. gov/ research/ publications/ publications.
htm?seq_no_115=213366). Biennial Workshop on Aerial Photography, Videography, and High Resolution Digital
Imagery for Resource
Assessment Proceedings (American Society for Photogrammetry and Remote Sensing). .
[18] Vreugdenhil, D., Meerman, J., Meyrat, A., Gómez, L.D., and D.J. Graham. 2002. "Map of the Ecosystems of Central America: Final
Report." World Bank, Washington, DC. 56 pp. http:/ / biological-diversity. info/ Downloads/ Ecosystem%20Mapping. zip
Mangrove 62
[19] Murray, M.R., Zisman, S.A., Furley, P.A., Munro, D.M., Gibson, J., Ratter, J., Bridgewater, S., Mity, C.D., and C.J. Place. 2003. "The
Mangroves of Belize: Part 1. Distribution, Composition and Classification." Forest Ecology and Management 174: 265–279
[20] http:/ / maps. cathalac. org/ Downloads/ data/ bz/ bz_mangroves_1980-2010_highres. pdf
[21] http:/ / www. wwf. org
[22] http:/ / www. cathalac. org
[23] Zisman, S.A. 1998. "Sustainability or Status Quo: Elite Influence and the Political Ecology of Mangrove Exploitation in Belize."
Doctoral
dissertation, Department of Geography, University of Edinburgh. Edinburgh, Scotland.
[24] Cherrington, E.A., Hernandez, B.E., Trejos, N.A., Smith, O.A., Anderson, E.R., Flores, A.I., and B.C. Garcia. 2010. "Identification of
Threatened and Resilient Mangroves in the Belize Barrier Reef System." Technical report to the World Wildlife Fund. Water Center for the
Humid Tropics of Latin America and the Caribbean (CATHALAC) / Regional Visualization & Monitoring System (SERVIR). 28 pp. http:/ /
maps. cathalac. org/ Downloads/ data/ bz/ bz_mangroves_1980-2010_highres. pdf
[25] http:/ / www. servir. net
[26] http:/ / www. rcmrd. org
[27] http:/ / www. icimod. org
[28] http:/ / www. nasa. gov
[29] http:/ / www. usaid. gov
[30] http:/ / www. nasa. gov/ home/ hqnews/ 2010/ oct/ HQ_10-241_SERVIR_Himalaya. html
[31] Mangroves of India (http:/ / www. mangroveindia. org/ ) - URL retrieved November 26, 2006
[32] Rouphael, Tony ;Turak, Emre and Brodie, Jon (1992) "Chapter 3: Seagrasses and Mangroves of Yemen's Red Sea" (http:/ / www. actfr.
jcu.
edu. au/ idc/ groups/ public/ documents/ journal_article/ jcudev_015628. pdf) In DouAbal, A. et al. (editors) (1992) Protection of
Marine
Ecosystems of the Red Sea Coast of Yemen Global Environment Facility, United Nations Development Programme, New York,
pp. 41-49
[33] Food and Agriculture Organization of the United Nations (FAO) (2007) The world's mangroves, 1980-2005: a thematic
study in the
framework of the Global Forest Resources Assessment 2005 (FAO forestry paper #153(FAO) Rome, page 37 (http:/ / books.
google. co. uk/
books?id=tLdlpOiuSmEC& pg=PA37), ISBN 978-92-5-105856-5
[34] "Millers Landing" (http:/ / www. ga. gov. au/ bin/ gazd01?rec=248996). Victorian Resources Online:West Gippsland.
Department of
Primary Industries.. . Retrieved 2009-03-30.
[35] "Millers Landing" (http:/ / www. ga. gov. au/ bin/ gazd01?rec=248996). Geoscience Australia Place Names Search.
Australian Government.
. Retrieved 2009-03-30.
[36] Zann, Leon P. (1996) [1995]. "Mangrove ecosystems in Australia: structure, function and status" (http:/ / www. deh. gov. au/ coasts/
publications/ somer/ annex1/ mangrove. html). State of the Marine Environment Report for Australia. Australian Government,
Dept of
Environment and Heritage. ISBN 0-642-17399-0. ISBN. . Retrieved 2006-11-25.
[37] Mangroves and Seagrasses - Treasures of the Sea (http:/ / www. treasuresofthesea. org. nz/ mangroves-and-seagrasses)
[38] Hawaii and the Pacific Islands (http:/ / biology. usgs. gov/ s+ t/ SNT/ noframe/ pi179. htm)
[39] Allen, James A. and Krauss, Ken W. (2006) "Influence of Propagule Flotation Longevity and Light Availability on Establishment of
Introduced Mangrove Species in Hawai'i". Pacific Science 60:3, July 2006. Abstract at (http:/ / muse. jhu. edu/ journals/ pacific_science/
toc/
psc60. 3. html) - URL retrieved November 28, 2006.
[40] Hawaiian Alien Plant Studies (http:/ / www. botany. hawaii. edu/ faculty/ cw_smith/ aliens. htm) - URL retrieved November 28, 2006.
[41] Warne, Kennedy; Tim Laman, photographer (2007-02). "Mangroves: Forests of the Tide" (http:/ / ngm. nationalgeographic. com/ 2007/
02/
mangroves/ warne-text/ 5). National Geographic. National Geographic Society. . Retrieved 2010-08-08.
[42] Sato, Gordon; Abraham Fisseha, Simon Gebrekiros, Hassan Abdul Karim, Samuel Negassi, Martin Fischer, Emanuel Yemane, Johannes
Teclemariam, and Robert Riley (2005-09). "A Novel Approach to Growing Mangroves on the Coastal Mud Flats of Eritrea with the Potential
for Relieving Regional Poverty and Hunger" (http:/ / springerlink. com/ content/ p251664gp74v5681/ ). Wetlands (The Society of Wetland
Scientists) 25 (3): 776–779. doi:10.1672/0277-5212(2005)025[0776:ANATGM]2.0.CO;2. . Retrieved 2010-08-08.
[43] Millennium Ecosystem Assessment (2005) Ecosystems and Human Well-being: Synthesis (p.2) Island Press, Washington,
DC. World
Resources Institute ISBN 1-59726-040-1
[44] Botkin, D. and E. Keller (2003) Environmental Science: Earth as a living planet (p.2) John Wiley & Sons. ISBN 0-471-
38914-5
[45] http:/ / www. earthscan. co. uk/ Products/ 9781844076574/ tabid/ 34104/ Default. aspx
[46] The Nature Conservancy (TNC). 2010a. ""World Atlas of Mangroves" Highlights the Importance of and Threats to Mangroves:
Mangroves
among World's Most Valuable Ecosystems." Press release. Arlington, Virginia. http:/ / www. nature. org/ pressroom/ press/ press4573. html
[47] http:/ / www. thepetitionsite. com/ petition/ 957999809
[48] http:/ / ecotippingpoints. org/ our-stories/ indepth/ thailand-mangrove-restoration-community-management. html
[49] Tree News, Spring/Summer 2005,Publisher Felix Press (http:/ / www. treecouncil. org. uk)
[50] Mangrove India website (http:/ / www. mangroveindia. org/ )
[51] http:/ / ambergriscaye. com/ pages/ town/ art/ Pelican_Review. pdf
[52] Cherrington, E.A., Ek, E., Cho, P., Howell, B.F., Hernandez, B.E., Anderson, E.R., Flores, A.I., Garcia, B.C., Sempris, E., and D.E.
Irwin.
2010. "Forest Cover and Deforestation in Belize: 1980-2010." Water Center for the Humid Tropics of Latin America and the Caribbean.
Panama City, Panama. 42 pp. http:/ / www. servir. net/ servir_bz_forest_cover_1980-2010. pdf
[53] Government of Belize (GOB). 2003. "Forests Act Subsidiary Laws." Chapter 213 in: Substantive Laws of Belize. Revised Edition 2003.
Government Printer: Belmopan, Belize. 137 pp. http:/ / www. belizelaw. org/ lawadmin/ PDF%20files/ cap213s. pdf
Mangrove 63
[54] http:/ / www. wri. org
[55] Cooper, E., Burke, L., and N. Bood. 2009. "Coastal Capital: Belize. The Contribution of Belize's Coral Reefs and Mangroves." WRI
Working Paper. World Resources Institute, Washington, DC. 53 pp. http:/ / pdf. wri. org/ working_papers/ coastal_capital_belize_wp. pdf
References
• Saenger, Peter (2002). Mangrove Ecology, Silviculture, and Conservation. Kluwer Academic
Publishers,
Dordrecht. ISBN 1-4020-0686-1.
• Hogarth, Peter J. (1999). The Biology of Mangroves. Oxford University Press, Oxford. ISBN 0-19-
850222-2.
• Thanikaimoni, Ganapathi (1986). Mangrove Palynology UNDP/UNESCO and the French Institute of
Pondicherry, ISSN 0073-8336 (E).
• Tomlinson, Philip B. (1986). The Botany of Mangroves. Cambridge University Press, Cambridge, ISBN
0-521-25567-8.
• Teas, H. J. (1983). Biology and Ecology of Mangroves. W. Junk Publishers, The Hague. ISBN 90-6193-
948-8.
• Plaziat, J.C., et al. (2001). "History and biogeography of the mangrove ecosystem, based on a critical
reassessment of the paleontological record". Wetlands Ecology and Management 9 (3): pp. 161–179.
• Sato, Gordon; Riley, Robert; et al. Growing Mangroves With The Potential For Relieving Regional Poverty
And
Hunger (http:/ / www. bioone. org/ perlserv/ ?request=get-pdf& doi=10. 1672/
0277-5212(2005)025[0776:ANATGM]2. 0. CO;2) WETLANDS, Vol. 25, No. 3 – September 2005
• Jayatissa, L. P., Dahdouh-Guebas, F. & Koedam, N. (2002). "A review of the floral composition and
distribution
of mangroves in Sri Lanka". Botanical Journal of the Linnean Society 138: 29–43.
• Warne, K. (February 2007). "Forests of the Tide". National Geographic pp. 132–151
• Aaron M. Ellison (2000) "Mangrove Restoration: Do We Know Enough?" Restoration Ecology 8 (3), 219–229
doi: 10.1046/j.1526-100x.2000.80033.x
• Agrawala, Shardul; Hagestad; Marca; Koshy, Kayathu; Ota, Tomoko; Prasad, Biman; Risbey, James; Smith,
Joel;
Van Aalst, Maarten. 2003. Development and Climate Change in Fiji: Focus on Coastal Mangroves. Organisation
of Economic Co-operation and Development, Paris, Cedex 16, France.
• Barbier, E.B., Sathirathai, S., 2001. Valuing Mangrove Conservation in Southern Thailand. Contemproary
Economic Policy. 19 (2) 109–122.
• Bosire, J.O., Dahdouh-Guebas, F., Jayatissa, L.P., Koedam, N., Lo Seen, D., Nitto, Di D. 2005. How Effective
were Mangroves as a Defense Against the Recent Tsunami? Current Biology Vol. 15 R443-R447.
• Bowen, Jennifer L., Valiela, Ivan, York, Joanna K. 2001. Mangrove Forests: One of the World's Threatened
Major Tropical Environments. Bio Science 51:10, 807–815.
• Jin-Eong, Ong. 2004. The Ecology of Mangrove Conservation and Management. Hydrobiologia. 295:1-3,
343–351.
• Glenn, C. R. 2006. "Earth's Endangered Creatures" (Online). Accessed 4/28/2008 at http:/ / earthsendangered.
com.
• Lewis, Roy R. III. 2004. Ecological Engineering for Successful Management and Restoration of Mangrove
Forest. Ecological Engineering. 24:4, 403–418.
• Lucien-Brun H. 1997. Evolution of world shrimp production: Fisheries and aquaculture. World Aquaculture.
28:21–33.
• Twilley, R. R., V.H. Rivera-Monroy, E. Medina, A. Nyman, J. Foret, T. Mallach, and L. Botero. 2000. Patterns
of
forest development in mangroves along the San Juan River estuary, Venezuela. Forest Ecology and
Management.
• Murray, M.R., Zisman, S.A., Furley, P.A., Munro, D.M., Gibson, J., Ratter, J., Bridgewater, S., Mity, C.D., and
C.J. Place. 2003. "The Mangroves of Belize: Part 1. Distribution, Composition and Classification." Forest
Ecology and Management 174: 265–279
• Cherrington, E.A., Hernandez, B.E., Trejos, N.A., Smith, O.A., Anderson, E.R., Flores, A.I., and B.C. Garcia.
2010. "Identification of Threatened and Resilient Mangroves in the Belize Barrier Reef System." Technical
report
Mangrove 64
to the World Wildlife Fund. Water Center for the Humid Tropics of Latin America and the Caribbean
(CATHALAC) / Regional Visualization & Monitoring System (SERVIR). 28 pp. http:/ / maps. cathalac. org/
Downloads/ data/ bz/ bz_mangroves_1980-2010_highres. pdf
• Vreugdenhil, D., Meerman, J., Meyrat, A., Gómez, L.D., and D.J. Graham. 2002. "Map of the Ecosystems of
Central America: Final Report." World Bank, Washington, DC. 56 pp.
External links
• Mangroves (http:/ / ocean. si. edu/ ocean-life-ecosystems/ mangroves/ )- At the Smithsonian Ocean Portal
• Rhizophoraceae (http:/ / www. dmoz. org/ Science/ Biology/ Flora_and_Fauna/ Plantae/ Magnoliophyta/
Magnoliopsida/ Rhizophoraceae/ ) at the Open Directory Project
• Mangrove forests (http:/ / www. dmoz. org/ Science/ Biology/ Ecology/ Aquatic_Ecology/ Marine/
Mangrove_Forests/ ) at the Open Directory Project
River delta
Nile River delta, as seen from Earth orbit. The
Nile is an example of a wave-dominated delta
that has the classic Greek delta (Δ) shape after
which River deltas were named. Photo courtesy
of NASA.
A delta is a landform that is formed at the mouth of a river where that
river flows into an ocean, sea, estuary, lake, reservoir, flat arid area, or
another river. Deltas are formed from the deposition of the sediment
carried by the river as the flow leaves the mouth of the river. Over long
periods of time, this deposition builds the characteristic geographic
pattern of a river delta.
The Greek historian Herodotus coined the term delta for the Nile River
delta because the sediment deposited at its mouth had the shape of the
upper-case Greek letter Delta: Δ.
Formation
River deltas form when a river carrying sediment reaches a body of
standing water,such as a lake, ocean, or reservoir. When the flow
enters the standing water, it is no longer confined to its channel and
expands in width. This flow expansion results in a decrease in the flow
velocity, which diminishes the ability of the flow to transport sediment.
As a result, sediment drops out of the flow and deposits. Over time,
this single channel will build a deltaic lobe (such as the bird's-foot of the Mississippi or Ural River deltas),
pushing
its mouth further into the standing water. As the deltaic lobe advances, the gradient of the river channel becomes
lower because the river channel is longer but has the same change in elevation (see slope). As the slope of the
river
channel decreases, it becomes unstable for two reasons. First, water under the force of gravity will tend to flow
in the
most direct course down slope. If the river could breach its natural levees (i.e., during a flood), it would spill out
onto
a new course with a shorter route to the ocean, thereby obtaining a more stable steeper slope.[1] Second, as its
slope
gets lower, the amount of shear stress on the bed will decrease, which will result in deposition of sediment
within the
channel and for the channel bed to rise relative to the floodplain. This will make it easier for the river to breach
its
levees and cut a new channel that enters the body of standing water at a steeper slope. Often when the channel
does
this, some of its flow can remain in the abandoned channel. When these channel switching events a mature delta
will
gain a distributary network.
River delta 65
Another way in which these distributary networks may form is from the deposition of mouth bars (mid-channel
sand
and/or gravel bars at the mouth of a river). When this mid-channel bar is deposited at the mouth of a river, the
flow is
routed around it. This results in additional deposition on the upstream end of the mouth-bar, which splits the
river
into two distributary channels. A good example of the result of this process is the Wax Lake Delta in Louisiana.
In both of these cases, depositional processes force redistribution of deposition from areas of high deposition to
areas
of low deposition. This results in the smoothing of the planform (or map-view) shape of the delta as the channels
move across its surface and deposit sediment. Because the sediment is laid down in this fashion, the shape of
these
deltas approximates a fan. It is closer to an ideal fan the more often the flow changes course because more rapid
changes in channel position results in more uniform deposition of sediment on the delta front. The Mississippi
and
Ural River deltas, with their bird's-feet, are examples of rivers that do not avulse often enough to form a
symmetrical
fan shape. Alluvial fan deltas, as seen in their name, avulse frequently and more closely approximate an ideal fan
shape.
Types of deltas
Lower Mississippi River landloss over time
Delta lobe switching in the Mississippi Delta,
4600 yrs BP, 3500 yrs BP, 2800 yrs BP, 1000 yrs
BP, 300 yrs BP, 500 yrs BP, current
Deltas are typically classified according to the main control on
deposition, which is usually either a river, waves, or tides.[2] These
controls have a large effect on the shape of the resulting delta.
River delta 66
Wave-dominated deltas
In wave dominated deltas, wave erosion controls the shape of the delta, although deposition still outweighs the
amount of erosion and the delta is able to advance into the sea. Deltas of this form, such as the Nile Delta, tend
to
have a characteristic Greek-capital-delta shape .
The Ganges Delta in India and Bangladesh is the
largest delta in the world and it is also one of the
most fertile regions in the world.
Tide-dominated deltas
Erosion is also an important control in tide dominated deltas, such as
the Ganges Delta, which may be mainly submarine, with prominent
sand bars and ridges. This tends to produce a "dendritic" structure.[2]
Tidal deltas behave differently from river- and wave-dominated deltas,
which tend to have a few main distributaries. Once a wave- or riverdistributary
silts up, it is abandoned, and a new channel forms
elsewhere. In a tidal delta, new distributaries are formed during times
when there's a lot of water around - such as floods or storm surges.
These distributaries slowly silt up at a pretty constant rate until they
fizzle out.[2]
Gilbert deltas
A Gilbert delta (named after Grove Karl Gilbert) is a specific type of
delta that is formed by coarse sediments, as opposed to gently-sloping muddy deltas such as that of the
Mississippi.
For example, a mountain river depositing sediment into a freshwater lake would form this kind of delta.[3] [4]
While
some authors describe both lacustrine and marine locations of Gilbert deltas,[3] others note that their formation is
more characteristic of the freshwater lakes, where it is easier for the river water to mix with the lakewater faster
(as
opposed to the case of a river falling into the sea or a salt lake, where less dense fresh water brought by the river
stays on top longer).[5]
G.K. Gilbert himself first described this type of delta on Lake Bonneville in 1885.[5] Elsewhere, similar
structures
can be found e.g. at the mouths of several creeks flowing into Okanagan Lake in British Columbia and forming
prominent peninsulas at Naramata (49°35′30″N 119°35′30″W), Summerland (49°34′23″N 119°37′45″W), or
Peachland (49°47′00″N 119°42′45″W)
Estuaries
Other rivers, particularly those located on coasts with significant tidal range, do not form a delta but enter into
the
sea in the form of an estuary. Notable examples include the Saint Lawrence River and the Tagus estuary.
Inland deltas
River delta 67
Okavango Delta
In rare cases the river delta is located inside a large valley and is called
an inverted river delta. Sometimes a river will divide into multiple
branches in an inland area, only to rejoin and continue to the sea; such
an area is known as an inland delta, and often occur on former lake
beds. The Inner Niger Delta is the most notable example. The Amazon
has also an inland delta before the island of Marajó.
In some cases a river flowing into a flat arid area splits into channels
which then evaporates as it progresses into the desert. Okavango Delta
in Botswana is one well-known example.
Sedimentary structure
The formation of a delta consists of three main forms: the topset, foreset/frontset, and bottomset.[3]
• The bottomset beds are created from the suspended sediment that settles out of the water as the river flows into
the body of water and loses energy. The suspended load is carried out the furthest into the body of water than all
other types of sediment creating a turbidite. These beds are laid down in horizontal layers and consist of smaller
grains.
• The foreset beds in turn build over the bottomset beds as the main delta form advances. The foreset beds
consist
of the bed load that the river is moving along which consists of larger sediments that roll along the main channel.
When it reaches the edge of the form, the bed load rolls over the edge, and builds up in steeply angled layers
over
the top of the bottomset beds. The angle of the outermost edge of the delta is created by the sediments angle of
repose. As the forsets build outward (which make up the majority of the delta) they pile up and miniature
landslides occur. This slope is created in this fashion as the bedload continues to be deposited and the delta
moves
outward. In cross section, one would see the foresets lying in angled, parallel bands, showing each stage of the
creation of the delta.
• The topset beds in turn overlay the foresets, and are horizontal layers of smaller sediment size that form as the
main channel of the river shifts elsewhere and the larger particles of the bed load no longer are deposited. As the
channels move across the top of the delta, the suspended load settles out in horizontal beds over the top.
Deltas and alluvial fans
Deltas are differentiated from alluvial fans in that deltas have a shallow slope, contain fine-grained sediment
(sand
and mud), and always flow into a body of water. Alluvial fans, on the other hand, are steep, have coarse-grained
sediments (including boulders), and are dominated by debris flows and large floods; these floods are often flash
floods. They can either flow onto a land surface, or into a body of water; in the latter case, they are called
alluvial fan
deltas.
Examples of notable deltas
The most famous delta is that of the Nile River, and it is this delta from which the term is derived. The
Ganges/Brahmaputra combination delta spans most of Bangladesh and West Bengal, empties into the Bay of
Bengal
and is the world's largest delta. Other rivers with notable deltas include, the Fly River, the Niger River, the
Tigris-Euphrates, the Rhine, the Po, the Rhône, the Danube, the Ebro, the Volga, the Lena, the Indus, the
Ayeyarwady (Irrawaddy), the Mekong, the Huanghe, the Yangtze, the Sacramento-San Joaquin, the Mississippi,
the
Orinoco, and the Paraná.
River delta 68
Ecological threats to deltas
Human activities, including diversion of water and the creation of dams for hydroelectric power or to create
reservoirs can radically alter delta ecosystems. Dams block sedimentation which can cause the delta to erode
away.
The use of water upstream can greatly increase salinity levels as less fresh water flows to meet the salty ocean
water.
While nearly all deltas have been impacted to some degree by humans, the Nile Delta and Colorado River Delta
are
some of the most extreme examples of the ecological devastation caused to deltas by damming and diversion of
water.
Deltas on Mars
Researchers have found a number of examples of deltas that formed in Martian lakes. Finding deltas is a major
sign
that Mars once had a lot of water. Deltas have been found over a wide geographical range. Below are pictures of
a
few.[6]
Delta in Ismenius Lacus
quadrangle, as seen by THEMIS.
Delta in Lunae Palus
quadrangle, as seen by
THEMIS.
Delta in Margaritifer Sinus
quadrangle as seen by
THEMIS.
Probable delta in a
crater to the NE of
Holden Crater, as
seen by Mars Global
Surveyor. Image in
Margaritifer Sinus
quadrangle.
References
[1] Slingerland, R. and N. D. Smith (1998), Necessary conditions for a meandering-river avulsion, Geology (Boulder), 26, 435-438.
[2] Fagherazzi, S (Dec 2008). "Self-organization of tidal deltas". Proceedings of the National Academy of Sciences of the
United States of
America 105 (48): 18692–5. doi:10.1073/pnas.0806668105. ISSN 0027-8424. PMC 2596246. PMID 19033190.
[3] Characteristics of deltas (http:/ / www. maine. gov/ doc/ nrimc/ mgs/ explore/ surficial/ facts/ dec03. htm). (Available archived at (http:/ /
web. archive. org/ web/ 20061012185632/ http:/ / www. maine. gov/ doc/ nrimc/ mgs/ explore/ surficial/ facts/ dec03. htm) - checked Dec
2008.)
[4] Bernard Biju-Duval, J. Edwin Swezey. "Sedimentary Geology". Page 183. ISBN 2710808021. Editions TECHNIP, 2002. Partial text
(http:/ /
books. google. com. au/ books?id=2txTeLt5MXgC) on Google Books.
[5] "Geological and Petrophysical Characterization of the Ferron Sandstone for 3-D Simulation of a Fluvial-deltaic Reservoir". By Thomas
C.
Chidsey, Thomas C. Chidsey, Jr (ed), Utah Geological Survey, 2002. ISBN 1557916683. Page 2-17. Partial text (http:/ / books. google. com.
au/ books?id=jacORXGQG9AC) on Google Books.
[6] Irwin III, R. et al. 2005. An intense terminal epoch of widespread fluvial activity on early Mars: 2. Increased runoff and paleolake
development. Journal of Geophysical Research: 10. E12S15
External links
• Louisiana State University Geology (http:/ / www. geol. lsu. edu/ WDD/ DELTA_LISTS/ continents. htm) -
World Deltas
Upland and lowland (freshwater

ecology)
Cascadilla Creek, near Ithaca, New York in the United States, an example of an
upland river habitat.
In studies of the ecology of freshwater
rivers, habitats are classified as upland and
lowland.
Definitions
Upland habitats are cold, clear, rocky, fast
flowing rivers in mountainous areas;
lowland habitats are warm, slow flowing
rivers found in relatively flat lowland areas,
with water that is frequently coloured by
sediment and organic matter.
These classifications overlap with the
geological definitions of "upland" and
"lowland". In geology an "upland" is
generally considered to be land that is at a
higher elevation than the alluvial plain or
stream terrace, which are considered to be "lowlands". The term "bottomland" refers to low-lying alluvial land
near a
river.
Many freshwater fish and invertebrate communities around the world show a pattern of specialisation into
upland or
lowland river habitats. Classifying rivers and streams as upland or lowland is important in freshwater ecology as
the
two types of river habitat are very different, and usually support very different populations of fish and
invertebrate
species.
Upland
In freshwater ecology, upland rivers and streams are the fast flowing rivers and streams that drain elevated or
mountainous country, often onto broad alluvial plains (where they become lowland rivers). However, altitude is
not
the sole determinant of whether a river is upland or lowland. Arguably the most important determinants are that
of
stream power and course gradient. Rivers with a course that drops in altitude rapidly will have faster water flow
and
higher stream power or "force of water". This in turn produces the other characteristics of an upland river - an
incised course, a river bed dominated by bedrock and coarse sediments, a riffle and pool structure and cooler
water
temperatures. Rivers with a course that drops in altitude very slowly will have slower water flow and lower
force.
This in turn produces the other characteristics of a lowland river - a meandering course lacking rapids, a river
bed
dominated by fine sediments and higher water temperatures. Lowland rivers tend to carry more suspended
sediment
and organic matter as well, but some lowland rivers have periods of high water clarity in seasonal low flow
periods.
The generally clear, cool, fast-flowing waters and bedrock and coarse sediment beds of upland rivers encourage
fish
species with limited temperature tolerances, high oxygen needs, strong swimming ability and specialised
reproductive strategies to prevent eggs or larvae being swept away. These characteristics also encourage
invertebrate
species with limited temperature tolerances, high oxygen needs and ecologies revolving around coarse sediments
and
interstices or "gaps" between those coarse sediments.
Lowland
Amazon River near Manaus, Brazil, an example
of a lowland river habitat.
The generally more turbid, warm, slow-flowing waters and fine
sediment beds of lowland rivers encourage fish species with broad
temperature tolerances and greater tolerances to low oxygen levels, and
life history and breeding strategies adapted to these and other traits of
lowland rivers. These characteristics also encourage invertebrate
species with broad temperature tolerances and greater tolerances to low
oxygen levels and ecologies revolving around fine sediments or
alternative habitats such as submerged woody debris ("snags") or
submergent macrophytes ("water weed").
Wetland
Florida's Everglades, the largest wetland system in the United
States.[1]
A wetland is an area of land whose soil is saturated
with moisture either permanently or seasonally. Such
areas may also be covered partially or completely by
shallow pools of water.[2] Wetlands include swamps,
marshes, and bogs, among others. The water found in
wetlands can be saltwater, freshwater, or brackish. The
world's largest wetland is the Pantanal which straddles
Brazil, Bolivia and Paraguay in South America.
Wetlands are considered the most biologically diverse
of all ecosystems. Plant life found in wetlands includes
mangrove, water lilies, cattails, sedges, tamarack, black
spruce, cypress, gum, and many others. Animal life
includes many different amphibians, reptiles, birds,
insects, and mammals.[3]
In many locations, such as the United Kingdom, Iraq, South Africa and the United States, wetlands are the
subject of
conservation efforts and Biodiversity Action Plans.
Wetlands also serve as natural wastewater purification systems—e.g., in Calcutta, India[4] and Arcata,
California.[5]
The study of wetlands has recently been termed paludology in some publications.[6]
Technical definitions
Wetlands have been categorized both as biomes and ecosystems.[3] A patch of land that develops pools of water
after
a rain storm would not be considered a "wetland" though the land is wet. Wetlands have unique characteristics:
they
are generally distinguished from other water bodies or landforms based on their water level and on the types of
plants that thrive within them. Specifically, wetlands are characterized as having a water table that stands at or
near
the land surface for a long enough season each year to support aquatic plants.[3] [7] [8] Put simply, wetlands are
lands
Wetland 71
made up of hydric soil.
Wetlands have also been described as ecotones, providing a transition between dry land and water bodies.[9]
Mitsch
and Gosselink write that wetlands exist "...at the interface between truly terrestrial ecosystems and aquatic
systems,
making them inherently different from each other, yet highly dependent on both."[10]
Ramsar Convention definition
Under the Ramsar international wetland conservation treaty, wetlands are defined as follows:
• Article 1.1: "...wetlands are areas of marsh, fen, peatland or water, whether natural or artificial, permanent or
temporary, with water that is static or flowing, fresh, brackish or salt, including areas of marine water the depth
of
which at low tide does not exceed six metres."
• Article 2.1: "[Wetlands] may incorporate riparian and coastal zones adjacent to the wetlands, and islands or
bodies of marine water deeper than six metres at low tide lying within the wetlands".
Regional definitions
In the United States, wetlands are defined as "those areas that are inundated or saturated by surface or
groundwater at
a frequency and duration sufficient to support, and that under normal circumstances do support, a prevalence of
vegetation typically adapted for life in saturated soil conditions. Wetlands generally include swamps, marshes,
bogs
and similar areas".[11] Some states, such as Massachusetts and New York, have separate definitions that may
differ
from United States.
Wetlands and climate change
Wetlands perform two important functions in relation to climate change. They have mitigation effects through
their
ability to sink carbon, and adaptation effects through their ability to store and regulate water.
Mitigation
Peatswamp forests and soils are being drained, burnt, mined, and overgrazed contributing severely to climate
change.
As Peatlands form only 3% of all the world’s land area, their degradation equal 7% of all fossil fuel carbon
dioxide
(CO2) emissions. As a result of peat drainage, the organic carbon that was built up over thousands of years and is
normally under water, is suddenly exposed to the air. It decomposes and turns into carbon dioxide (CO2), which
is
released into the atmosphere. Peat fires cause the same process and in addition create enormous clouds of smoke
that
cross international borders, such as happen every year in Southeast Asia.
Through the building of dams, Wetlands International is halting the drainage of peatlands in Southeast Asia,
thereby
avoiding enormous CO2 emissions. Reforestation with native tree species as well as setting up community fire
brigades are also part of an integral, very cost effective approach shown in Central Kalimantan and Sumatra,
Indonesia. (from: www.wetlands.org)
Adaptation
Mangroves, floodplains, highland and other wetlands can reduce the impacts of increased precipitation, storms,
glacier melting and even sealevel rise. The water regulating and storage functions of these wetlands are crucial in
adapting to a changing climate. Therefore, in order to protect people living close or even far downstream from
wetlands and their source of livelihoods, we must conserve and restore wetlands. In this way, they can continue
to
play their critical role. (from www.wetlands.org)
Wetland 72
Conservation
Wetlands have historically been the victim of large-scale draining efforts for real estate development, or flooding
for
use as recreational lakes. By 1993 half the world's wetlands had been drained.[12] Since the 1970s, more focus
has
been put on preserving wetlands for their natural function — sometimes also at great expense. Wetlands provide
a
valuable flood control function. Wetlands are very effective at filtering and cleaning water pollution,[13] (often
from
agricultural runoff from the farms that replaced the wetlands in the first place). To replace these wetland
ecosystem
services enormous amounts of money have been spent on water purification plants and remediation measures,
constructing dams, leves and other artificial flood controls.
Ramsar Convention
The Convention on Wetlands of International Importance, especially as Waterfowl Habitat,
or Ramsar Convention,
is an international treaty designed to address global concerns regarding wetland loss and degradation. The
primary
purposes of the treaty are to list wetlands of international importance and to promote their wise use, with the
ultimate
goal of preserving the world's wetlands. Methods include restricting access to the majority portion of wetland
areas,
as well as educating the public to combat the misconception that wetlands are wastelands.
Conservation by country
New Zealand
Over 90% of the wetlands in New Zealand have been drained since European settlement, predominantly to
create
farmland. Wetlands now have a degree of protection under the Resource Management Act.
South Africa
The South African Department of Environmental Affairs and Tourism in conjunction with the departments of
Water
Affairs and Forestry, and of Agriculture, supports the conservation and rehabilitation of wetlands through the
Working for Wetlands program.[14] The aim of this program is to encourage the protection, rehabilitation and
sustainable use of South African wetlands through co-operative governance and partnerships. The program is
also a
poverty relief effort, providing employment in wetland maintenance.
Sweden
The Swedish national wetland inventory (VMI) is one of the world's most extensive systematic inventories of
nature
types that has ever been done. VMI has surveyed the wetlands of Sweden below the alpine region during a 25-
year
period.[15] In total 35 000 objects (sites) are included in VMI, corresponding to an area of 4.3 million hectares, or
10% of the land area of Sweden. The aim of the survey has been to increase the general knowledge of wetlands
in
Sweden, as a basis for environmental monitoring and natural resources planning. By investigating the impact of
human activities on wetlands and identifying the most valuable wetlands, their values can be preserved for future
generations. The results from the inventory were also meant to function as background data for the authorities'
decisions concerning e.g. drainage permits.
United States
The USA came to understand how biologically productive wetlands are, so the USA passed laws limiting
wetlands
destruction, and created requirements that if a wetland had to be drained, developers at least had to offset the loss
by
creating artificial wetlands. One example is the project by the U.S. Army Corps of Engineers to control flooding
and
enhance development by taming the Everglades, a project which has now been reversed to restore much of the
wetlands as a natural habitat for plant and animal life, as well as a method of flood control. Another project in
the
works to restore the Everglades is the U.S Sugar Corp Land Transaction [16]. The project entails the acquisition
of
U.S. Sugar Corp. land, which would allow for water delivery, water treatment and water storage of sufficient
Wetland 73
quantity and quality to mimic the Everglades' natural system. The Everglades Foundation, a foundation whose
mission is to protect and restore one of the world’s unique natural ecosystems, and other environment-based
organizations support the state of Florida acting to secure U.S. Sugar Corp. land.[17]
Mapping
There are many remote sensing methods that can be used to map wetlands. Remote-sensing technology permits
the
acquisition of timely digital data on a repetitive basis. This repeat coverage allows wetlands, as well as the
adjacent
land-cover and land-use types, to be monitored seasonally and/or annually. Using digital data provides a
standardized data-collection procedure and an opportunity for data integration within a geographic information
system. Traditionally, Landsat 5 Thematic Mapper (TM), Landsat 7 Enhanced Thematic Mapper Plus (ETM + ),
and
the SPOT 4 and 5 satellite systems have been used for this purpose. More recently, however, multispectral
IKONOS
and QuickBird data, with spatial resolutions of 4m by 4m and 2.44m by 2.44 m, respectively, have been shown
to be
excellent sources of data when mapping and monitoring smaller wetland habitats and vegetation communities.
A wide range of remote sensing studies has been undertaken in a variety of wetland environments. Remote
sensing
technology has permitted the acquisition of timely digital data on a repetitive basis. For example, the wetlands
and
vegetation within Detroit Lakes Wetland management District has been assessed using remote sensing. In
mapping
and monitoring large geographic areas, analysis of satellite images is less costly and time-consuming compared
to
visual interpretation of aerial photographs. Aerial photographs also require experienced interpreters to extract
information based on structure and texture while remote sensing only requires the analysis of the spectral
characteristics of data.
However, there are a number of limitations associated with image acquisition. Analysis of wetlands has proved
difficult because to obtain the data, it has to be linked with practical purposes such as the analysis of land cover
or
land use. Wetlands, in particular are difficult to monitor, are often difficult to access, especially their inner
reaches,
and are sometimes home to dangerous wildlife and endemic diseases. Developing a global inventory of wetlands
has
proven to be a large and difficult undertaking. Current efforts are based on available data, but both classification
and
spatial resolution may be inadequate for regional or site-specific management decision-making. It is difficult to
identify small, long, and narrow wetlands within the landscape. Current efforts using today’s remote sensing
satellites may not have sufficient spatial and spectral resolution to monitor wetland conditions, although
multispectral IKONOS and QuickBird data may offer improved spatial resolutions of 4m or higher. Majority of
the
pixels are just mixtures of several plant species or vegetation types and are difficult to isolate. Improved remote
sensing information, coupled with good knowledge domain on wetlands will facilitate expanded efforts in
wetland
monitoring and mapping. This will also be extremely important because we expect to see major shifts in species
composition due to both anthropogenic (land use) and natural changes in the environment caused by climate
change.
Methods to focus the classification on specific classes of interest so that identification can be done with very
high
accuracies should be looked into. The issue of the cost and expertise involved in remote sensing technology is
still a
factor hindering further advancements in image acquisition and data processing. Future improvements in wetland
vegetation mapping could include the use of more recent and better geospatial data.
Wetland 74
Climate
Temperature
Wetlands contrast the hot, arid landscape around
Middle Spring, Fish Springs National Wildlife
Refuge, Utah.
Temperatures vary greatly depending on the location of the wetland.
Many of the world's wetlands are in temperate zones (midway between
the North and South Poles and the equator). In these zones, summers
are warm and winters are cold, but temperatures are not extreme.
However, wetlands found in the tropic zone, which is around the
equator, are always warm. Temperatures in wetlands on the Arabian
Peninsula, for example, can reach 50 °C (122 °F). In northeastern
Siberia, which has a polar climate, wetland temperatures can be as cold
as −50 °C (−58 °F). And in a moderate zone, such as the Gulf of
Mexico, is usually 11 °C (51 °F).
Rainfall
The amount of rainfall a wetland receives depends upon its location. Wetlands in Wales, Scotland, and Western
Ireland receive about 150 cm (59 in) per year. Those in Southeast Asia, where heavy rains occur, can receive up
to
1000 cm (200 in) In the northern areas of North America, wetlands exist where as little as 18 centimeters (6
inches)
of rain fall each month
List of wetland types
• Swamp
• Freshwater swamp forest
• Coniferous swamp
• Marsh
• Salt marsh
• Bog
• Peat swamp forest
• Slough
• Flooded grasslands and savannas
• Constructed wetland
• Riparian zone
• Fen
• Vernal Pool
References
[1] http:/ / www. ens-newswire. com/ ens/ apr2002/ 2002-04-15-06. asp
[2] "National Geographic's Strange Days on Planet Earth: Glossary" (http:/ / www. pbs. org/ strangedays/ glossary/ W. html). Public
Broadcasting Service. . Retrieved 2009-10-02.
[3] http:/ / www. thewildclassroom. com/ biomes/ wetlands. html
[4] East Calcutta Wetlands: Wastewater, Fishponds, and Agriculture (http:/ / ecotippingpoints. org/ our-stories/ indepth/
india-calcutta-wetland-wastewater-agriculture-fishpond)
[5] Arcata, California Constructed Wetland: A Cost-Effective Alternative for Wastewater Treatment (http:/ / ecotippingpoints. org/ our-
stories/
indepth/ usa-california-arcata-constructed-wetland-wastewater)
[6] http:/ / psjc. icm. edu. pl/ psjc/ cgi-bin/ getdoc. cgi?AAAA015683
[7] http:/ / www. cvwd. net/ water_glossary. htm
[8] http:/ / mapping2. orr. noaa. gov/ portal/ calcasieu/ calc_html/ resources/ glossary. html
[9] http:/ / www. alabamapower. com/ hydro/ glossary. asp
[10] Mitsch, William J.; James G. Gosselink (2007-08-24). Wetlands (4th ed.). New York: John Wiley & Sons. ISBN 978-0471699675.
[11] EPA Regulations listed at 40 CFR 230.3(t)
[12] "unknown title". New Scientist (1894): 46. 1993-10-09.
Wetland 75
[13] Letting Nature Do the Job (http:/ / www. wild. org/ blog/ letting-nature-do-the-job)
[14] http:/ / wetlands. sanbi. org/ Working for Wetlands
[15] "Swedish national wetland inventory - The result of 25 years of inventories" (http:/ / www. naturvardsverket. se/ Documents/
publikationer/
978-91-620-5925-5. pdf). Swedish Environmental Protection Agency. . Retrieved 23 August 2010.
[16] http:/ / www. evergladesfoundation. org/ pages/ us-sugar
[17] Everglades Foundation. US Sugar. (http:/ / www. evergladesfoundation. org/ pages/ us-sugar) Retrieved 2010-07-23.
Wetlands International (http:/ / www. wetlands. org) Wetlands International Africa (http:/ / www. afrique.
wetlands.
org) Wetlands International Latin America & the Caribbean (http:/ / www. lac. wetlands. org) Wetlands
International
Indonesia (http:/ / www. wetlands. or. id) Wetlands International Japan (http:/ / www. japan. wetlands. org)
Ramsar
Convention on Wetlands (http:/ / www. ramsar. org)
Further reading
• Mitsch, W.J., J.G. Gosselink, C.J. Anderson, and L. Zhang. (2009) " Wetland Ecosystems (http:/ / www. wiley.
com/ WileyCDA/ WileyTitle/ productCd-047028630X. html)". John Wiley & Sons, Inc., New York, 295 pp.
• Ghabo, A. A. (2007) Wetlands Characterization; Use by Local Communities and Role in Supporting
Biodiversity
in the Semiarid Ijara District, Kenya (http:/ / www. terranuova. info/ file_download/ 51). Terra Nuova East
Africa. Wetlands in drylands.
• 1987 U.S. Army Corps of Engineers Wetland delineation manual (http:/ / www. cpe. rutgers. edu/ brochures/
pdfs/ 1987-Army-Corps-Wetlands-Delineation-Manual. pdf)
• Dugan, Patrick (editor) (1993) Wetlands in Danger, World Conservation Atlas Series
• Brinson, M. (1993) A Hydrogeomorphic Classification of Wetlands (http:/ / el. erdc. usace. army. mil/
wetlands/
pdfs/ wrpde4. pdf)
• Fredrikson, Leigh H. (1983) "Wetlands: A Vanishing Resource" (http:/ / naldr. nal. usda. gov/ NALWeb/
Agricola_Link. asp?Accession=IND84105117) Yearbook of Agriculture
• W.H. MacKenzie and J.R. Moran (2004) "Wetlands of British Columbia: A Guide to Identification. Ministry of
Forests, Land Management Handbook 52. (http:/ / www. for. gov. bc. ca/ hfd/ pubs/ Docs/ Lmh/ Lmh52. pdf)
External links
• Wetlands International - A non-profit global organisation to sustain and restore wetlands, their resources and
biodiversity for future generations (http:/ / www. wetlands. org)
• US Fish & Wildlife Service: National Wetlands Inventory (http:/ / www. fws. gov/ wetlands/ )
• Wetlands: The Ecological Effect of Loss (Research article) (http:/ / www. articlemyriad. com/ 37. htm)
• (French) Pôle-relais zones humides littorales de la façade atlantique, Manche et Mer du Nord (http:/ / www.
forum-marais-atl. com/ )
• Marshlands of Iberá (http:/ / www. esterosdelibera. com/ ) (in English and Spanish)
• Wetlands (http:/ / www. nal. usda. gov/ wqic/ wetl. shtml) Water Quality Information Center, U.S. Department
of
Agriculture
• Centro Studi Naturalistici (http:/ / www. centrostudinatura. it/ )
• Wetlands legal updates (http:/ / blog. sprlaw. com/ category/ wetlands/ )
• Wetland of Hong Kong (http:/ / www. wwf. org. hk/ eng/ maipo/ ?source=fp-topmenu)
• Forum des Marais Atlantiques - Pôle-relais zones humides littorales de la façade atlantique, Manche et Mer du
Nord (http:/ / www. forum-marais-atl. com/ )
Endorheic basin 76
Endorheic basin
NASA photo of the endorheic Tarim Basin
Endorheic basin showing waterflow input into
Üüreg Nuur
An endorheic basin [1] (from the Greek: ἔνδον, éndon, "within" and
ῥεῖν, rheîn, "to flow"; also terminal or closed basin) is a closed
drainage basin that retains water and allows no outflow to other bodies
of water such as rivers or oceans. Normally, water that has accrued in a
drainage basin eventually flows out through rivers or streams on
Earth's surface or by underground diffusion through permeable rock,
ultimately ending up in the oceans. However, in an endorheic basin,
rain (or other precipitation) that falls within it does not flow out but
may only leave the drainage system by evaporation and seepage. The
bottom of such a basin is typically occupied by a salt lake or salt pan.
Endorheic basins are also called internal drainage systems.
Endorheic regions, in contrast to exorheic regions which flow to the
ocean in geologically defined patterns, are closed hydrologic systems.
Their surface waters drain to inland terminal locations where the water
evaporates or seeps into the ground, having no access to discharge into
the sea.[2] Endorheic water bodies include some of the largest lakes in
the world, such as the Aral Sea and the Caspian Sea, the world’s largest
saline body of water cut off from the ocean.[3]
Endorheic lakes
Endorheic lakes are bodies of water that do not flow into the sea. Most
of the water falling on earth finds its way to the oceans through a
network of rivers, lakes and wetlands. However, there is a class of
water bodies that are located in closed or endorheic watersheds where
the topography prevents their drainage to the oceans. These endorheic watersheds (containing water in rivers or
lakes
that form a balance of surface inflows, evaporation and seepage) are often called terminal lakes or sink lakes.[4]
Endorheic lakes are usually in the interior of a body mass, far from an ocean. Their watersheds are often
confined by
natural geologic land formations such as a mountain range, cutting off water access to the ocean. The inland
water
flows into dry watersheds where the water evaporates, leaving a high concentration of minerals and other inflow
erosion products. Over time this input of erosion products can cause the endorheic lake to become relatively
saline (a
"salt lake"). Since the main outflow pathways of these lakes are chiefly through evaporation and seepage,
endorheic
lakes are usually more sensitive to environmental pollutants inputs than water bodies that have access to oceans.
[3]
Occurrence
Endorheic regions can occur in any climate but are most commonly found in hot desert locations. In areas where
rainfall is higher, riparian erosion will generally carve drainage channels (particularly in times of flood), or cause
the
water level in the terminal lake to rise until it finds an outlet, breaking the enclosed endorheic hydrological
system’s
geographical barrier and opening it to the surrounding terrain. The Black Sea was likely such a lake, having once
been an independent hydrological system before the Mediterranean Sea broke through the terrain separating the
two.
Endorheic regions tend to be far inland with their boundaries defined by mountains or other geological features
that
block their access to oceans. Since the inflowing water can evacuate only through seepage or evaporation, dried
minerals or other products collect in the basin, eventually making the water saline and also making the basin
Endorheic basin 77
vulnerable to pollution.[3] Continents vary in their concentration of endorheic regions due to conditions of
geography
and climate. Australia has the highest percentage of endorheic regions at 21 percent while North America has the
least at 5 percent.[5] Approximately 18 percent of the earth’s land drains to endorheic lakes or seas, the largest of
these land areas being the interior of Asia.
In deserts, water inflow is low and loss to solar evaporation high, drastically reducing the formation of complete
drainage systems. Closed water flow areas often lead to the concentration of salts and other minerals in the basin.
Minerals leached from the surrounding rocks are deposited in the basin, and left behind when the water
evaporates.
Thus endorheic basins often contain extensive salt pans (also called salt flats, salt lakes, alkali flats, dry lake
beds or
playas). These areas tend to be large, flat hardened surfaces and are sometimes used for aviation runways or land
speed record attempts, because of their extensive areas of perfectly level terrain.
Both permanent and seasonal endorheic lakes can form in endorheic basins. Some endorheic basins are
essentially
stable, climate change having reduced precipitation to the degree that a lake no longer forms. Even most
permanent
endorheic lakes change size and shape dramatically over time, often becoming much smaller or breaking into
several
smaller parts during the dry season. As humans have expanded into previously uninhabitable desert areas, the
river
systems that feed many endorheic lakes have been altered by the construction of dams and aqueducts. As a result
many endorheic lakes in developed or developing countries have contracted dramatically, resulting in increased
salinity, higher concentrations of pollutants, and the disruption of ecosystems.
Notable endorheic basins and lakes
Major endorheic basins of the world. Basins are shown in dark gray; major endorheic lakes are shown in black. Colored regions represent the
major
drainage patterns of the continents to the oceans (non-endorheic). Continental divides are indicated by dark lines.
Endorheic basin 78
Antarctica
Endorheic lakes in Antarctica are located in the McMurdo Dry Valleys, Victoria Land, Antarctica, the largest
ice-free area in Antarctica.
• Don Juan Pond in Wright Valley is fed by groundwater from a rock glacier and remains unfrozen throughout
the
year.
• Lake Vanda in Wright Valley has a perennial ice cover, the edges of which melt in the summer allowing flow
from the longest river in Antarctica, the Onyx River. The lake is over 70 m deep and is hypersaline.
• Lake Bonney is in Taylor Valley and has a perennial ice over and two lobes separated by the Bonney Riegel.
The
lake is fed by glacial melt and discharge from Blood Falls. Its unique glacial history has resulted in a hypersaline
brine in the bottom waters and fresh water at the surface.
• Lake Hoare, in Taylor Valley, is the freshest of the Dry Valley lakes receiving its melt almost exclusively from
the Canada Glacier. The lake has an ice cover and forms a moat during the Austral summer.
• Lake Fryxell, in adjacent to the Ross Sea in Taylor Valley. The lake has an ice cover and receives its water
from
numerous glacial meltwater streams for approximately 6 weeks out of the year. Its salinity increases with depth.
Asia
Caspian Sea, a giant inland basin
Lake Van, Turkey
Much of western and Central Asia is a giant endorheic region made up
of a number of contiguous closed basins. The region contains several
basins and terminal lakes, including:
• The Caspian Sea, the largest lake on Earth. A large part of Eastern
Europe, drained by the Volga River, is part of the Caspian's basin.
• The Aral Sea, whose tributary rivers have been diverted, leading to
a dramatic shrinkage of the lake. The resulting ecological disaster
has brought the plight faced by internal drainage basins to public
attention.
• Lake Balkhash, in Kazakhstan.
• Issyk-Kul Lake, Son-Kul Lake, and Chatyr-Kul Lake in
Kyrgyzstan.
• Lop Lake, in the Tarim Basin of China's Xinjiang Uygur
Autonomous Region.
• The Central Asian Internal Drainage Basin, in southern and western
Mongolia, contains a series of closed drainage basins, such as the
Khyargas Nuur basin, the Uvs Nuur basin, and the Pu-Lun-To River
Basin.[6]
• Qaidam Basin, in Qinghai Province, China.
• Sistan Basin covering areas of Iran and Afghanistan
Other endorheic lakes and basins in Asia include:
• The Dead Sea, the lowest surface point on Earth and one of its
saltiest bodies of water, lies between Israel and Jordan.
• Sambhar Lake, in Rajasthan, north-western India, is also the
terminal point of an endorheic basin.
• Lake Van, in Turkey, is one of the world's largest endorheic lakes.
• Sabkhat al-Jabbul, extensive salt flats and a 100 square kilometres
(39 sq mi) lake in Syria.
• Solar Lake, Sinai, near the Israeli-Egypt border.
Endorheic basin 79
Australia
A false-colour satellite photo of Australia’s Lake
Eyre
Image credit: NASA’s Earth Observatory
Australia, being very dry and having exceedingly low runoff ratios due
to its ancient soils, has many endorheic drainages. The most important
are:
• Lake Eyre Basin, which drains into the highly variable Lake Eyre
and includes Lake Frome.
• Lake Torrens, to the west of the Flinders Ranges in South Australia.
• Lake Corangamite, a highly saline crater lake in western Victoria.
• Lake George, formerly connected to the Murray-Darling Basin
Africa
Large endorheic regions in Africa are located in the Sahara Desert, the
Kalahari Desert, and the Great Rift Valley:
• Lake Chad, (between Chad and Cameroon), fed by the Chari and
Logon rivers.
• Qattara Depression, in Egypt.
• Chott Melrhir, in Algeria.
• Chott el Djerid, in Tunisia.
• The Okavango River, in the Kalahari Desert, is part of an endorheic basin region which also includes the
Okavango Delta, Lake Ngami, the Nata River, and a number of salt pans such as Makgadikgadi Pan.
• Etosha pan in Namibia’s Etosha National Park.
• Lake Turkana, in Kenya, whose basin includes the Omo River of Ethiopia.
• Lake Chilwa, in Malawi.
• Afar Depression, in Eritrea, Ethiopia, and Djibouti.
• Some Rift Valley lakes, such as Lake Abijatta, Lake Chew Bahir, Lake Shala, Lake Chamo, and Lake Awasa.
• Lake Mweru Wantipa, in Zambia.
North and Central America
The dry lake at Badwater in Death Valley
National Park.
• The Valley of Mexico. In Pre-Columbian times, the Valley was
substantially covered with five lakes, including Lake Texcoco, Lake
Xochimilco, and Lake Chalco.
• Guzmán Basin, in northern Mexico and the southwestern United
States.
• Lago de Atitlán, in the highlands of Guatemala.
• Bolsón de Mapimí, in northern Mexico.
• The Great Basin[7] , which covers much of Nevada and Utah, is a
large region of contiguous closed basins, including:
• Great Salt Lake, in Utah, the largest terminal lake in the Western
Hemisphere.
• The Black Rock Desert in Nevada, location of the Thrust2 and ThrustSSC landspeed record runs, and the
annual home to the Burning Man festival.
• Death Valley in California and Nevada, the lowest land point in the United States. It had a system of tributary
basins:
Endorheic basin 80
Great Salt Lake, Satellite photo (2003) after five
years of drought
• Rogers Lake, at Edwards Air Force Base in California.
• Owens Lake and the Owens River basin.
• Mono Lake in California.
• Groom Dry Lake in Nevada, location of Area 51.
• Salton Sea in California, a lake accidentally recreated in 1905
when irrigation canals ruptured, filling a desert endorheic basin
and recreating an ancient saline sea.
• Sevier Lake, in Utah.
• Pyramid Lake in Nevada, whose drainage basin includes Lake
Tahoe.
• Humboldt Sink and the Humboldt River basin in Nevada.
• Tulare Lake at the southern end of the San Joaquin Valley fed by
the Kaweah, Kern and Tule Rivers plus southern distributaries of
the Kings, however agricultural development and irrigation diversions have left the lake dry.
• Crater Lake, in Oregon.
• The Great Divide Basin in Wyoming, a small endorheic basin that straddles the Continental Divide of the
Americas.
• Devils Lake, in North Dakota.
• Devil's Lake, in Wisconsin.
• Little Manitou Lake in Saskatchewan.
• Old Wives Lake, on the Laurentian Divide in Saskatchewan.
• Pakowki Lake, on the Laurentian Divide in Alberta.
• New Mexico has a number of desert endorheic basins including:
• The Tularosa Basin, a rift valley.
• Zuni Salt Lake, a maar.
• The Mimbres River Basin, in Grant County.
Many small lakes and ponds in North Dakota and Manitoba are endorheic; some of them have salt encrustations
along their shores.
Europe
• Lake Neusiedl, in Austria and Hungary.
• Lake Trasimeno, in Italy.
• Lake Velence, in Hungary.
• Lake Prespa, between Albania, Greece and the Republic of Macedonia.
• Rahasane turlough, the largest turlough in Ireland.
• Laacher See, in Germany.
All these lakes are drained, however, either through manmade canals or via karstic phenomena. Minor additional
endorheic lakes exist throughout the Mediterranean countries Spain (e.g. Laguna de Gallocanta), Italy, Cyprus
(Larnaca and Akrotiri salt lakes) and Greece.
Endorheic basin 81
South America
MODIS image from November 4, 2001 showing
Lake Titicaca, the Salar de Uyuni, and the Salar
de Coipasa. These are all parts of the Altiplano
• The Altiplano includes a number of closed basins such as Poopó
Lake.
• Lake Valencia, in Venezuela.
• Salar de Atacama, in the Atacama Desert, Chile.
Ancient
Some of the Earth’s ancient endorheic systems include:
• The Black Sea, until its merger with the Mediterranean.
• The Mediterranean Sea itself and all its tributary basins, during its
Messinian desiccation (5 m.y. BP approx.) as it became
disconnected from the Atlantic Ocean.
• Lake Lahontan in North America.
• Lake Bonneville in North America.
• Ebro and Duero basins, draining most of northern Spain during the
Neogene and perhaps Pliocene.
References
[1] This term is little-used among North American geoscientists.
[2] "Drainage systems" (http:/ / www. britannica. com/ eb/ topic-187043/ endorheic-system). Encyclopedia Britannica. . Retrieved 2008-02-
11.
[3] "Endorheic Lakes: Waterbodies That Don't Flow to the Sea" (http:/ / www. unep. or. jp/ ietc/ publications/ short_series/ lakereservoirs-2/
10.
asp). United Nations Environment Programme. . Retrieved 2008-02-11.
[4] "What is a watershed and why should I care?" (http:/ / www. wr. udel. edu/ cb/ whatwhycare. html). university of delaware. . Retrieved
2008-02-11.
[5] Saline Lake Ecosystems of the World (http:/ / books. google. com/ books?id=NOdvPFm6SyoC& pg=PA18& lpg=PA18&
dq=endorheic&
source=web& ots=EWyVvVeOhm& sig=tNrylNMQCGu6bUCBTyo1HHlGDRc). Springer. . Retrieved 2007-07-31.
[6] "Basins" (http:/ / www. mongolianriverresources. mn/ index. php?option=com_content& task=view& id=81& pid=366). Mongolian
River
Resources. . Retrieved 21 November 2010.
[7] Houghton, Samuel G. (1994). A trace of desert waters: the Great Basin story. Reno: University of Nevada Press.
External links
• Primer on endorheic lakes (http:/ / www. unep. or. jp/ ietc/ Publications/ Short_Series/ LakeReservoirs-2/ 10.
asp)
• The Silk Roads and Eurasian Geography (http:/ / depts. washington. edu/ silkroad/ geography/ geography.
html)
• The role of climate during high plateau formation. Insights from numerical experiments (http:/ / www.
sciencedirect. com/ science?_ob=ArticleURL& _udi=B6V61-4N5TNBS-2& _user=10& _rdoc=1& _fmt=&
_orig=search& _sort=d& view=c& _acct=C000050221& _version=1& _urlVersion=0& _userid=10&
md5=ba0480557981cd1ca96050abc5fdecb8)
Palearctic ecozone
The Palearctic ecozone.
The Palearctic or Palaearctic is one
of the eight ecozones dividing the
Earth's surface.
Physically, the Palearctic is the largest
ecozone. It includes the terrestrial
ecoregions of Europe, Asia north of the
Himalaya foothills, northern Africa,
and the northern and central parts of
the Arabian Peninsula.
Major ecological regions
The Palearctic ecozone includes mostly boreal and temperate climate ecoregions, which run across Eurasia from
western Europe to the Bering Sea.
Euro-Siberian region
The boreal and temperate Euro-Siberian region is the Palearctic's largest biogeographic region, which transitions
from tundra in the northern reaches of Russia and Scandinavia to the vast taiga, the boreal coniferous forests
which
run across the continent. South of the taiga are a belt of temperate broadleaf and mixed forests and temperate
coniferous forests. This vast Euro-Siberian region is characterized by many shared plant and animal species, and
has
many affinities with the temperate and boreal regions of the Nearctic ecoregion of North America. Eurasia and
North
America were often connected by the Bering land bridge, and have very similar mammal and bird fauna, with
many
Eurasian species having moved into North America, and fewer North American species having moved into
Eurasia.
Many zoologists consider the Palearctic and Nearctic to be a single Holarctic ecozone. The Palearctic and
Nearctic
also share many plant species, which botanists call the Arcto-Tertiary Geoflora.
Mediterranean Basin
The lands bordering the Mediterranean Sea in southern Europe, north Africa, and western Asia are home to the
Mediterranean basin ecoregions, which together constitute world's largest and most diverse mediterranean
climate
region of the world, with generally mild, rainy winters and hot, dry summers. The Mediterranean basin's mosaic
of
Mediterranean forests, woodlands, and scrub are home to 13,000 endemic species. The Mediterranean basin is
also
one of the world's most endangered biogeographic regions; only 4% of the region's original vegetation remains,
and
human activities, including overgrazing, deforestation, and conversion of lands for pasture, agriculture, or
urbanization, have degraded much of the region. Formerly the region was mostly covered with forests and
woodlands, but heavy human use has reduced much of the region to the sclerophyll shrublands known as
chaparral,
matorral, maquis, or garrigue. Conservation International has designated the Mediterranean basin as one of the
world's biodiversity hotspots.
Sahara and Arabian deserts
A great belt of deserts, including the Atlantic coastal desert, Sahara desert, and Arabian desert, separates the
Palearctic and Afrotropic ecoregions. This scheme includes these desert ecoregions in the palearctic ecozone;
other
biogeographers identify the ecozone boundary as the transition zone between the desert ecoregions and the
Mediterranean basin ecoregions to the north, which places the deserts in the Afrotropic, while others place the
boundary through the middle of the desert.
Western and Central Asia
The Caucasus mountains, which run between the Black Sea and the Caspian Sea, are a particularly rich mix of
coniferous, broadleaf, and mixed forests, and include the temperate rain forests of the Euxine-Colchic deciduous
forests ecoregion.
Central Asia and the Iranian plateau are home to dry steppe grasslands and desert basins, with montane forests,
woodlands, and grasslands in the region's high mountains and plateaux. In southern Asia the boundary of the
Palearctic is largely altitudinal. The middle altitude foothills of the Himalaya between about 2000–2500 m form
the
boundary between the Palearctic and Indomalaya ecoregions.
East Asia
China, Korea and Japan are more humid and temperate than adjacent Siberia and Central Asia, and are home to
rich
temperate coniferous, broadleaf, and mixed forests, which are now mostly limited to mountainous areas, as the
densely populated lowlands and river basins have been converted to intensive agricultural and urban use. East
Asia
was not much affected by glaciation in the ice ages, and retained 96 percent of Pliocene tree genera, while
Europe
retained only 27 percent. In the subtropical southern parts of China and, the Palearctic temperate forests
transition to
the subtropical and tropical forests of Indomalaya, creating a rich and diverse mix of plant and animal species.
The
mountains of southwest China are also designated as a biodiversity hotspot. In Southeastern Asia, high mountain
ranges form tongues of Palearctic flora and fauna in northern Indochina and southern China. Isolated small
outposts
(sky islands) occur as far south as central Myanmar (on Nat Ma Taung, 3050 m), northernmost Vietnam (on Fan
Si
Pan, 3140 m) and the high mountains of Taiwan.
Freshwater
The ecozone contains several important freshwater ecoregions as well, including the heavily developed rivers of
Europe, the rivers of Russia, which flow into the Arctic, Baltic, Black, and Caspian seas, Siberia's Lake Baikal,
the
oldest and deepest lake on the planet, and Japan's ancient Lake Biwa.
Flora and fauna
One bird family, the accentors (Prunellidae) is endemic to the Palearctic region. The Holarctic has four other
endemic bird families: the divers or loons (Gaviidae), grouse (Tetraoninae), auks (Alcidae), and waxwings
(Bombycillidae).
There are no endemic mammal orders in the region, but several families are endemic: Calomyscidae (mouse-like
hamsters), Prolagidae, and Ailuridae (red pandas). Several mammal species originated in the Palearctic, and
spread
to the Nearctic during the ice ages, including the Brown Bear (Ursus arctos, known in North America as the
Grizzly), Red Deer (Cervus elaphus) in Europe and the closely related Elk (Cervus canadensis) in far
eastern
Siberia, American Bison (Bison bison), and Reindeer (Rangifer tarandus, known in North America as the
Caribou).
Palearctic terrestrial ecoregions
Palearctic Tropical and subtropical moist broadleaf forests
Guizhou Plateau broadleaf and mixed forests China
Yunnan Plateau subtropical evergreen forests China
Palearctic Temperate broadleaf and mixed forests
Appenine deciduous montane forests Italy
Atlantic mixed forests Belgium, Denmark, France, Germany. Netherlands
Azores temperate mixed forests Portugal
Balkan mixed forests Bulgaria, Greece, Macedonia, Romania, Serbia (Kosovo), Turkey
Baltic mixed forests Denmark, Germany, Poland, Sweden
Cantabrian mixed forests Portugal, Spain
Caspian Hyrcanian mixed forests Azerbaijan, Iran
Caucasus mixed forests Armenia, Azerbaijan, Georgia, Russia, Turkey
Celtic broadleaf forests Ireland, United Kingdom
Central Anatolian deciduous forests Turkey
Central China loess plateau mixed
forests
China
Central European mixed forests Austria, Belarus, Czech Republic, Germany, Lithuania, Moldova, Poland
Central Korean deciduous forests North Korea, South Korea
Changbai Mountains mixed forests China, North Korea
Changjiang Plain evergreen forests China
Crimean Submediterranean forest
complex
Russia, Ukraine
Daba Mountains evergreen forests China
Dinaric Mountains mixed forests Albania, Bosnia and Herzegovina, Croatia, Italy, Montenegro, Serbia, Slovenia
East European forest steppe Bulgaria, Moldova, Romania, Russia, Ukraine
Eastern Anatolian deciduous forests Turkey
English Lowlands beech forests United Kingdom
Euxine-Colchic deciduous forests Georgia, Turkey
Hokkaido deciduous forests Japan
Huang He Plain mixed forests China
Madeira evergreen forests Portugal
Manchurian mixed forests China, North Korea, Russia, South Korea
Nihonkai evergreen forests Japan
Nihonkai montane deciduous forests Japan
North Atlantic moist mixed forests Ireland, United Kingdom
Northeast China Plain deciduous forests China
Pannonian mixed forests Austria, Bosnia and Herzegovina, Croatia, Czech Republic, Hungary, Romania, Serbia, Slovakia,
Slovenia, Ukraine
Po Basin mixed forests Italy
Pyrenees conifer and mixed forests Andorra, France, Spain
Qin Ling Mountains deciduous forests China
Rodope montane mixed forests Bulgaria, Greece, Macedonia, Serbia
Sarmatic mixed forests Belarus, Estonia, Finland, Latvia, Lithuania, Norway, Russia, Sweden
Sichuan Basin evergreen broadleaf
forests
China
South Sakhalin-Kurile mixed forests Russia
Southern Korea evergreen forests South Korea
Taiheiyo evergreen forests Japan
Taiheiyo montane deciduous forests Japan
Tarim Basin deciduous forests and
steppe
Xinjiang, China
Ussuri broadleaf and mixed forests Russia
West Siberian broadleaf and mixed
forests
Russia
Western European broadleaf forests Austria, Czech Republic, France, Germany, Switzerland
Zagros Mountains forest steppe Iran
Palearctic Temperate coniferous forests
Alps conifer and mixed forests Austria, France, Germany, Italy, Slovenia, Switzerland
Altai montane forest and forest steppe China, Kazakhstan, Mongolia, Russia
Caledonian conifer forests United Kingdom
Carpathian montane conifer forests Czech Republic, Poland, Romania, Slovakia, Ukraine
Da Hinggan-Dzhagdy Mountains conifer forests China, Russia
East Afghan montane conifer forests Afghanistan, Pakistan
Elburz Range forest steppe Iran
Helanshan montane conifer forests China
Hengduan Mountains subalpine conifer forests China
Hokkaido montane conifer forests Japan
Honshu alpine conifer forests Japan
Khangai Mountains conifer forests Mongolia, Russia
Mediterranean conifer and mixed forests Algeria, Morocco, Tunisia
Northeastern Himalayan subalpine conifer forests China, India, Bhutan
Northern Anatolian conifer and deciduous forests Turkey
Nujiang Langcang Gorge alpine conifer and mixed forests China
Qilian Mountains conifer forests China
Qionglai-Minshan conifer forests China
Sayan montane conifer forests Mongolia, Russia
Scandinavian coastal conifer forests Norway, Finland, Sweden
Tian Shan montane conifer forests China, Kazakhstan, Kyrgyzstan, Tajikistan, Uzbekistan
Palearctic Boreal forests/taiga
East Siberian taiga Russia
Iceland boreal birch forests and alpine tundra Iceland
Kamchatka-Kurile meadows and sparse forests Russia
Kamchatka-Kurile taiga Russia
Northeast Siberian taiguhh Russia
Okhotsk-Manchurian taiga Russia
Sakhalin Island taiga Russia
Scandinavian and Russian taiga Finland, Norway, Russia, Sweden
Trans-Baikal conifer forests Mongolia, Russia
Urals montane tundra and taiga Russia
West Siberian taiga Russia
Puszcza Romnicka Poland, Russia
Palearctic Temperate grasslands, savannas, and shrublands
Alai-Western Tian Shan steppe Kazakhstan, Tajikistan, Uzbekistan
Altai steppe and semi-desert Kazakhstan
Central Anatolian steppe Turkey
Daurian forest steppe China, Mongolia, Russia
Eastern Anatolian montane steppe Armenia, Iran, Turkey
Emin Valley steppe China, Kazakhstan
Faroe Islands boreal grasslands Faroe Islands, part of Denmark
Gissaro-Alai open woodlands Kyrgyzstan, Tajikistan, Uzbekistan
Kazakh forest steppe Kazakhstan, Russia
Kazakh steppe Kazakhstan, Russia
Kazakh upland Kazakhstan
Middle East steppe Iraq, Syria
Mongolian-Manchurian grassland China, Mongolia, Russia
Pontic steppe Kazakhstan, Moldova, Romania, Russia, Ukraine
Sayan Intermontane steppe Russia
Selenge-Orkhon forest steppe Mongolia, Russia
South Siberian forest steppe Russia
Tian Shan foothill arid steppe China, Kazakhstan, Kyrgyzstan
Palearctic Flooded grasslands and savannas
Amur meadow steppe China, Russia
Bohai Sea saline meadow China
Nenjiang River grassland China
Nile Delta flooded savanna Egypt
Saharan halophytics Algeria, Egypt, Mauritania, Tunisia, Western Sahara
Tigris-Euphrates alluvial salt marsh Iraq, Iran
Ussuri-Wusuli meadow and forest meadow China, Russia
Yellow Sea saline meadow China
Palearctic Montane grasslands and shrublands
Altai alpine meadow and tundra People's Republic of China, Kazakhstan, Mongolia, Russia
Central Tibetan Plateau alpine steppe China
Eastern Himalayan alpine shrub and meadows Bhutan, Burma, People's Republic of China, India, Nepal
Ghorat-Hazarajat alpine meadow Afghanistan
Hindu Kush alpine meadow Afghanistan, Pakistan
Karakoram-West Tibetan Plateau alpine steppe Afghanistan, People's Republic of China, India, Pakistan
Khangai Mountains alpine meadow Mongolia
Kopet Dag woodlands and forest steppe Iran, Turkmenistan
Kuhrud-Kohbanan Mountains forest steppe Iran
Mediterranean High Atlas juniper steppe Morocco
North Tibetan Plateau-Kunlun Mountains alpine desert China
Northwestern Himalayan alpine shrub and meadows People's Republic of China, India, Pakistan
Ordos Plateau steppe China
Pamir alpine desert and tundra Afghanistan, China, Kyrgyzstan, Tajikistan
Qilian Mountains subalpine meadows China
Sayan Alpine meadows and tundra Mongolia, Russia
Southeast Tibet shrub and meadows China
Sulaiman Range alpine meadows Afghanistan, Pakistan
Tian Shan montane steppe and meadows China, Kazakhstan, Kyrgyzstan
Tibetan Plateau alpine shrub and meadows People's Republic of China
Western Himalayan alpine shrub and meadows India, Nepal
Yarlung Zambo arid steppe China
Palearctic Tundra
Arctic desert Russia, Svalbard (Norway)
Bering tundra Russia
Cherskii-Kolyma mountain tundra Russia
Chukchi Peninsula tundra Russia
Kamchatka Mountain tundra and forest tundra Russia
Kola Peninsula tundra Norway, Russia
Northeast Siberian coastal tundra Russia
Northwest Russian-Novaya Zemlya tundra Russia
Novosibirsk Islands arctic desert Russia
Scandinavian Montane Birch forest and grasslands Finland, Norway, Sweden
Taimyr-Central Siberian tundra Russia
Trans-Baikal Bald Mountain tundra Russia
Wrangel Island arctic desert Russia
Yamalagydanskaja tundra Russia
Palearctic Mediterranean forests, woodlands, and scrub
Aegean and Western Turkey sclerophyllous and mixed forests Greece, Macedonia, Turkey
Anatolian conifer and deciduous mixed forests Turkey
Canary Islands dry woodlands and forests Spain
Corsican montane broadleaf and mixed forests France
Crete Mediterranean forests Greece
Cyprus Mediterranean forests Cyprus
Eastern Mediterranean conifer-sclerophyllous-broadleaf forests Israel, Jordan, Lebanon, Syria, Turkey
Iberian conifer forests Spain
Iberian sclerophyllous and semi-deciduous forests Portugal, Spain
Illyrian deciduous forests Albania, Bosnia and Herzegovina, Croatia, Greece, Italy, Slovenia
Italian sclerophyllous and semi-deciduous forests France, Italy
Mediterranean acacia-argania dry woodlands and succulent thickets Morocco, Canary Islands (Spain)
Mediterranean dry woodlands and steppe Algeria, Egypt, Libya, Morocco, Tunisia
Mediterranean woodlands and forests Algeria, Morocco, Tunisia
Northeastern Spain and Southern France Mediterranean forests France, Monaco, Spain
Northwest Iberian montane forests Portugal, Spain
Pindus Mountains mixed forests Albania, Greece, Macedonia
South Appenine mixed montane forests Italy
Southeastern Iberian shrubs and woodlands Spain
Southern Anatolian montane conifer and deciduous forests Israel, Jordan, Lebanon, Syria, Turkey
Southwest Iberian Mediterranean sclerophyllous and mixed forests Spain
Tyrrhenian-Adriatic sclerophyllous and mixed forests Croatia, France, Italy, Malta
Palearctic Deserts and xeric shrublands
Afghan Mountains semi-desert Afghanistan
Alashan Plateau semi-desert China, Mongolia
Arabian Desert and East Sahero-Arabian xeric shrublands Egypt, Jordan, Oman, Saudi Arabia, Syria, United Arab Emirates, Yemen
Atlantic coastal desert Mauritania, Western Sahara
Azerbaijan shrub desert and steppe Azerbaijan, Georgia, Iran
Badkhiz-Karabil semi-desert Afghanistan, Iran, Tajikistan, Turkmenistan, Uzbekistan
Baluchistan xeric woodlands Afghanistan, Pakistan
Caspian lowland desert Iran, Kazakhstan, Russia, Turkmenistan
Central Afghan Mountains xeric woodlands Afghanistan
Central Asian northern desert Kazakhstan, Uzbekistan
Central Asian riparian woodlands Kazakhstan
Central Asian southern desert Kazakhstan, Turkmenistan, Uzbekistan
Central Persian desert basins Afghanistan, Iran
Eastern Gobi desert steppe China, Mongolia
Gobi Lakes Valley desert steppe Mongolia
Great Lakes Basin desert steppe Mongolia, Russia
Junggar Basin semi-desert China, Mongolia
Kazakh semi-desert Kazakhstan
Kopet Dag semi-desert Iran, Turkmenistan
Mesopotamian shrub desert Iraq, Jordan, Syria
North Saharan steppe and woodlands Algeria, Egypt, Libya, Morocco, Tunisia, Western Sahara
Paropamisus xeric woodlands Afghanistan
Persian Gulf desert and semi-desert Bahrain, Kuwait, Qatar, Saudi Arabia, United Arab Emirates
Qaidam Basin semi-desert China
Red Sea Nubo-Sindian tropical desert and semi-desert Iraq, Jordan, Oman, Saudi Arabia, Yemen
Rigestan-North Pakistan sandy desert Afghanistan, Iran, Pakistan
Sahara desert Algeria, Chad, Egypt, Libya, Mali, Mauritania, Niger, Sudan
South Iran Nubo-Sindian desert and semi-desert Iran, Iraq, Pakistan
South Saharan steppe and woodlands Algeria, Chad, Mali, Mauritania, Niger, Sudan
Taklimakan desert China
Tibesti-Jebel Uweinat montane xeric woodlands Chad, Egypt, Libya, Sudan
West Saharan montane xeric woodlands Algeria, Mali, Mauritania, Niger
References
• Amorosi, T. "Contributions to the zooarchaeology of Iceland: some preliminary notes." in
'The Anthropology of
Iceland' (eds. E.P. Durrenberger & G. Pálsson). Iowa City: University of Iowa Press, pages 203-227, 1989.
• Buckland, P.C., et al. "Holt in Eyjafjasveit, Iceland: a paleoecological study of the impact of
Landnám." in Acta
Archaeologica 61: pp. 252–271. 1991.
External links
• Map of the ecozones [1]
References
[1] http:/ / www. panda. org/ about_wwf/ where_we_work/ ecoregions/ maps/ index. cfm
Cold seep
A cold seep (sometimes called a cold vent) is an area of the ocean floor where hydrogen sulfide, methane
and other
hydrocarbon-rich fluid seepage occurs, often in the form of a brine pool. Cold seeps constitute a biome
supporting
several endemic species.
Cold seeps develop unique topography over time, where reactions between methane and seawater create
carbonate
rock formations and reefs. These reactions may also be dependent on bacterial activity. Ikaite, a hydrous calcium
carbonate, can be associated with oxidizing methane at cold seeps.
Types
These craters mark the formation of brine pools,
from which salt has seeped through the seafloor
and encrusted the nearby substrate.
Type of cold seeps can be distinguished as according to the depth to
shallows cold seeps and deep cold seeps.[1] Cold seeps can also be
distinguished in detail as follows:
• oil/gas seeps[1]
• gas seeps:[1] methane seeps
• gas hydrate seeps[1]
• brine seeps[1] are forming brine pools
• pockmarks[1]
• mud volcanos[1]
Cold seep 91
Formation and ecological succession
Cold seeps occur over fissures on the seafloor caused by tectonic activity. Oil and methane "seep" out of those
fissures, get diffused by sediment, and emerge over an area several hundred meters wide.[2]
Methane (CH4) is the main component of what we commonly refer to as natural gas.[2] But in addition to being
an
important energy source for humans, methane also forms the basis of a cold seep ecosystem.[2]
Chemosynthetic communities
Bacterial mat consisting of sulfide-oxidizing
bacteria Beggiatoa spp. at a seep on Blake Ridge,
off South Carolina. The red dots are
range-finding laser beams.
Organisms living in cold seeps are known as extremophiles. Biological
research in cold seeps and hydrothermal vents has been mostly focused
on the microbiology and the prominent chemosynthetic
macro-invertebrates.[1] Much less research has been done on the
smaller benthic fraction at the size of the meiofauna (<1 mm).[1]
Community composition's orderly shift from one set of species to
another is called ecological succession:[2]
The first type of organism to take advantage of this deep-sea energy
source is bacteria.[2] Aggregating into bacterial mats at cold seeps,
these bacteria metabolize methane and hydrogen sulfide (another gas
that emerges from seeps) for energy.[2] This process of obtaining
energy from chemicals is known as chemosynthesis.[2]
A mussel bed at the edge of the brine pool.
During this initial stage, when methane is relatively abundant, dense
mussel beds also form near the cold seep.[2] Mostly composed of
species in the genus Bathymodiolus, these mussels do not directly
consume food.[2] Instead, they are nourished by symbiotic bacteria that
also produce energy from methane, similar to their relatives that form
mats.[2]
This microbial activity produces calcium carbonate (CaCO3), which is
deposited on the seafloor and forms a layer of rock.[2] During a period
lasting up to several decades, these rock formations attract
vestimentiferan tubeworms (family Siboglinidae), which settle and grow along with the mussels.[2] Like the
mussels,
tubeworms rely on chemosynthetic bacteria (in this case, a type that needs hydrogen sulfide instead of methane)
for
survival.[2] True to any symbiotic relationship, a tubeworm also provides for their bacteria by appropriating
hydrogen sulfide from the environment.[2] The sulfide not only comes from the water, but is also mined from the
sediment through an extensive "root" system a tubeworm "bush" establishes in the hard, carbonate substrate.[2] A
tubeworm bush can contain hundreds of individual worms, which can grow a meter or more above the sediment.
[2]
Cold seeps do not last indefinitely. As the rate of gas seepage slowly decrease, the shorter-lived, methane-hungry
mussels (or more precisely, their methane-hungry bacterial symbionts) start to die off.[2] At this stage,
tubeworms
become the dominant organism in a seep community.[2] As long as there is some sulfide in the sediment, the
sulfide-mining tubeworms can persist.[2] Individuals of one tubeworm species Lamellibrachia luymesi have
been
estimated to live for over 250 years in such conditions.[2]
Cold seep 92
"Roots" of tubeworms also provide a supply of
hydrogen sulfide from the sediment to the
bacteria inside these tubeworms.
Symbiotic vestimentiferan tubeworm
Lamellibrachia luymesi from a cold seep at 550
m depth in the Gulf of Mexico. In the sediments
around the base are orange bacterial mats of the
sulfide-oxidizing bacteria Beggiatoa spp. and
empty shells of various clams and snails, which
are also common inhabitants of the seeps.[3]
Tubeworms, soft corals and chemosynthetic
mussels at a seep located 3000 m (9843 ft) down
on the Florida Escarpment. Eelpouts, a Galatheid
crab and an alvinocarid shrimp feed on mussels
damaged during a sampling exercise.
Comparison with other communities
Cold seeps and hydrothermal vents are communities that do not rely on photosynthesis for food and energy
production.[1] These systems are largely driven by chemosynthetic derived energy.[1] Both systems share
common
characteristics such as the presence of reduced chemical compounds (H2S and hydrocarbonates), local hypoxia
or
even anoxia, a high abundance and metabolic activity of bacterial populations, and the production of
autochthonous,
organic material by chemoautotrophic bacteria.[1] Both hydrothermal vents and cold seeps show regularly,
highly
increased levels of metazoan biomass in association with a low local diversity.[1] This is explained through the
presence of dense aggregations of foundation species and epizooic animals, living within these aggregations.[1]
However, hydrothermal vents and cold seeps differ also in many ways. Compared to the more stable cold seeps,
vents are characterized by locally high temperatures, strongly fluctuating temperatures, pH, sulphide and oxygen
concentrations, often the absence of sediments, a relatively young age, and often unpredictable conditions, such
as
waxing and waning of vent fluids or volcanic eruptions.[1] Unlike hydrothermal vents, which are volatile and
ephemeral environments, cold seeps emit at a slow and dependable rate. Likely owing to the cooler temperatures
and
stability, many cold seep organisms are much longer-lived than those inhabiting hydrothermal vents.
End of cold seep community
Finally, as cold seeps become inactive, tubeworms also start to disappear, clearing the way for corals to settle on
the
now exposed carbonate substrate.[2] The corals do not rely on hydrocarbons seeping out of the seafloor.[2]
Studies on
Lophelia pertusa suggest they derive their nutrition primarily from the ocean surface.[2] Chemosynthesis
plays only a
very small role, if any, in their settlement and growth.[2] While deepwater corals do not seem to be
chemosynthesis-based organisms, the chemosynthetic organisms that come before them enable the corals'
existence.[2] This hypothesis about establishment of deep water coral reefs is called hydraulic theory.[4] [5]
Cold seep 93
Distribution
Cold seeps were discovered in 1983 by Dr. Charles Paull in the Gulf of Mexico at a depth of 3200 meters (10499
ft).
Since then, seeps have been discovered in other parts of the world's oceans. They have also been grouped into
several biogeographic provinces: Gulf of Mexico, Atlantic, Mediterranean, East Pacific, West Pacific[6] and
under an
ice shelf in Antarctica.[7] Those localities include the Monterey Canyon just off Monterey Bay, California, the
Sea of
Japan, off the Pacific coast of Costa Rica, in the Atlantic off of Africa and in waters off the coast of Alaska. The
deepest seep community known is found in the Japan trench at a depth of 7326 m (24035 ft).
• Map of some cold seeps (marked with a star) [8]
In the Gulf of Mexico
The manned submersible DSV Alvin, which made
possible the discovery of chemosynthetic
communities in the Gulf of Mexico in 1983.
Discoveries in the Gulf of Mexico
The chemosynthetic communities of the Gulf of Mexico have been
studied extensively over the past 20 years, and communities first
discovered on the upper slope are likely the best understood seep
communities in the world.[9] The history of the discovery of these
remarkable animals has all occurred within the last 30 years.[9]
Interestingly, each major discovery was unexpected―from the first
hydrothermal vent communities anywhere in the world to the first cold
seep communities in the Gulf of Mexico.[9]
Communities were discovered in the Eastern Gulf of Mexico in 1983
using the manned submersible DSV Alvin, during a cruise investigating
the bottom of the Florida Escarpment in areas of “cold” brine seepage,
where they unexpectedly discovered tubeworms and mussels (Paull et
al., 1984).[9]
Two groups fortuitously discovered chemosynthetic communities in
the Central Gulf of Mexico concurrently in November 1984.[9] During
investigations by Texas A&M University to determine the effects of oil
seepage on benthic ecology (until this investigation, all effects of oil seepage were assumed to be detrimental),
bottom trawls unexpectedly recovered extensive collections of chemosynthetic organisms, including tube worms
and
clams (Kennicutt et al., 1985).[9] At the same time, LGL Ecological Research Associates was conducting a
research
cruise as part of the multiyear MMS Northern Gulf of Mexico Continental Slope Study (Gallaway et al., 1988).
[9]
Bottom photography (processed on board the vessel) resulted in clear images of vesicomyid clam
chemosynthetic
communities coincidentally in the same manner as the first discovery by camera sled in the Pacific in 1977.[9]
Photography during the same LGL/MMS cruise also documented tube-worm communities in situ in the Central
Gulf
of Mexico for the first time (not processed until after the cruise; Boland, 1986) prior to the initial submersible
investigations and firsthand descriptions of Bush Hill in 1986 (Rosman et al., 1987a; MacDonald et al., 1989b).
[9]
The site was targeted by acoustic “wipeout” zones or lack of substrate structure caused by seeping hydrocarbons.
[9]
This was determined using an acoustic pinger system during the same cruise on the R/V Edwin Link (the old
one,
only 113 ft (34 m)), which used one of the Johnson Sea Link submersibles.[9] The site is characterized by
dense
tubeworm and mussel accumulations, as well as exposed carbonate outcrops with numerous gorgonian and
Lophelia
coral colonies. Bush Hill has become one of the most thoroughly studied chemosynthetic sites in the world.[9]
Cold seep 94
Distribution in the Gulf of Mexico
Chemosynthetic communities in the northern part
of Gulf Of Mexico around cold seeps known in
2000.
There is a clear relationship between known hydrocarbon discoveries
at great depth in the Gulf slope and chemosynthetic communities,
hydrocarbon seepage, and authigenic minerals including carbonates at
the seafloor (Sassen et al., 1993a and b).[9] While the hydrocarbon
reservoirs are broad areas several kilometers beneath the Gulf,
chemosynthetic communities occur in isolated areas with thin veneers
of sediment only a few meters thick.[9]
The northern Gulf of Mexico slope includes a stratigraphic section
more than 10 km (6 mi) thick and has been profoundly influenced by
salt movement.[9] Mesozoic source rocks from Upper Jurassic to Upper
Cretaceous generate oil in most of the Gulf slope fields (Sassen et al.,
1993a and b).[9] Migration conduits supply fresh hydrocarbon materials
through a vertical scale of 6–8 km (4-5 mi) toward the surface.[9] The surface expressions of hydrocarbon
migration
are referred to as seeps.[9] Geological evidence demonstrates that hydrocarbon and brine seepage persists in
spatially
discrete areas for thousands of years.[9]
The time scale for oil and gas migration (combination of buoyancy and pressure) from source systems is on the
scale
of millions of years (Sassen, 1997).[9] Seepage from hydrocarbon sources through faults towards the surface
tends to
be diffused through the overlying sediment, carbonate outcroppings, and hydrate deposits so the corresponding
hydrocarbon seep communities tend to be larger (a few hundred meters wide) than chemosynthetic communities
found around the hydrothermal vents of the Eastern Pacific (MacDonald, 1992).[9] There are large differences in
the
concentrations of hydrocarbons at seep sites.[9] Roberts (2001) presented a spectrum of responses to be expected
under a variety of flux rate conditions varying from very slow seepage to rapid venting.[9] Very slow seepage
sites do
not support complex chemosynthetic communities; rather, they usually only support simple microbial mats
(Beggiatoa sp.).[9]
In the upper slope environment, the hard substrates resulting from carbonate precipitation can have associated
communities of nonchemosynthetic animals, including a variety of sessile cnidarians such as corals and
anemones.[9]
At the rapid flux end of the spectrum fluidized sediment generally accompanies hydrocarbons and formation
fluids
arriving at the seafloor.[9] Mud volcanoes and mud flows result.[9] Somewhere between these two end members
exists the conditions that support densely populated and diverse communities of chemosynthetic organisms
(microbial mats, siboglinid tube worms, bathymodioline mussels, lucinid and vesycomyid clams, and associated
organisms).[9] These areas are frequently associated with surface or near-surface gas hydrate deposits.[9] They
also
have localized areas of lithified seafloor, generally authigenic carbonates but sometimes more exotic minerals
such
as barite are present.[9]
Chemosynthetic communities in the northern part
of Gulf Of Mexico around cold seeps known in
2006 include more than 50 communities.
The widespread nature of Gulf of Mexico chemosynthetic communities
was first documented during contracted investigations by the
Geological and Environmental Research Group (GERG) of Texas
A&M University for the Offshore Operators Committee (Brooks et al.,
1986).[9] This survey remains the most widespread and comprehensive,
although numerous additional communities have been documented
since that time.[9] Industry exploring for energy reserves in the Gulf of
Mexico has also documented numerous new communities through a
wide range of depths, including the deepest known occurrence in the
Central Gulf of Mexico in Alaminos Canyon Block 818 at a depth of
Cold seep 95
2,750 m (9,022 ft).[9] The occurrence of chemosynthetic organisms dependent on hydrocarbon seepage has been
documented in water depths as shallow as 290 m (951 ft) (Roberts et al., 1990) and as deep as 2,744 m (9,003 ft)
(Allen, personal communication, 2005).[9] This depth range specifically places chemosynthetic communities in
the
deepwater region of the Gulf of Mexico, which is defined as water depths greater than 305 m (1,000 ft).[9]
Chemosynthetic communities are not found on the continental shelf although they do appear in the fossil record
in
water shallower than 200 m (656 ft).[9] One theory explaining this is that predation pressure has varied
substantially
over the time period involved (Callender and Powell 1999).[9] More than 50 communities are now known to
exist in
43 OCS blocks.[9] Although a systematic survey has not been done to identify all chemosynthetic communities
in the
Gulf of Mexico, there is evidence indicating that many more such communities may exist.[9] The depth limits of
discoveries probably reflect the limits of exploration (lack of submersibles capable of depths over 1,000 m
(3,281 ft)).[9]
MacDonald et al. (1993 and 1996) have analyzed remote-sensing images from space that reveal the presence of
oil
slicks across the north-central Gulf of Mexico.[9] Results confirmed extensive natural oil seepage in the Gulf of
Mexico, especially in water depths greater than 1,000 m (3,281 ft).[9] A total of 58 additional potential locations
were
documented where seafloor sources were capable of producing perennial oil slicks (MacDonald et al., 1996).[9]
Estimated seepage rates ranged from 4 bbl/d (0.64 m3/d) to 70 bbl/d (11 m3/d) compared to less than 0.1 bbl/d
(0.016 m3/d) for ship discharges (both normalized for 1,000 mi2 (640,000 ac)).[9] This evidence considerably
increases the area where chemosynthetic communities dependent on hydrocarbon seepage may be expected.[9]
The densest aggregations of chemosynthetic organisms have been found at water depths of around 500 m (1,640
ft)
and deeper.[9] The best known of these communities was named Bush Hill by the investigators who first
described it
(MacDonald et al., 1989b).[9] It is a surprisingly large and dense community of chemosynthetic tube worms and
mussels at a site of natural petroleum and gas seepage over a salt diapir in Green Canyon Block 185. The seep
site is
a small knoll that rises about 40 m (131 ft) above the surrounding seafloor in about 580-m (1,903-ft) water
depth.[9]
Stability
According to Sassen (1997) the role of hydrates at chemosynthetic communities has been greatly
underestimated.[9]
The biological alteration of frozen gas hydrates was first discovered during the MMS study " Stability and
Change in
Gulf of Mexico Chemosynthetic Communities [10]".[9] It is hypothesized (MacDonald, 1998b) that the
dynamics of
hydrate alteration could play a major role as a mechanism for regulation of the release of hydrocarbon gases to
fuel
biogeochemical processes and could also play a substantial role in community stability.[9] Recorded bottom-
water
temperature excursions of several degrees in some areas such as the Bush Hill site (4-5 °C at 500-m (1,640-ft)
depth)
are believed to result in dissociation of hydrates, resulting in an increase in gas fluxes (MacDonald et al., 1994).
[9]
Although not as destructive as the volcanism at vent sites of the mid-ocean ridges, the dynamics of shallow
hydrate
formation and movement will clearly affect sessile animals that form part of the seepage barrier.[9] There is
potential
of a catastrophic event where an entire layer of shallow hydrate could break free of the bottom and result in
considerable impact to local communities of chemosynthetic fauna.[9] At deeper depths (>1,000 m, >3,281 ft),
the
bottom-water temperature is colder (by approximately 3 °C) and undergoes less fluctuation.[9] The formation of
more stable and probably deeper hydrates influences the flux of light hydrocarbon gases to the sediment surface,
thus
influencing the surface morphology and characteristics of chemosynthetic communities.[9] Within complex
communities such as Bush Hill, petroleum seems less important than previously thought (MacDonald, 1998b).[9]
Through taphonomic studies (death assemblages of shells) and interpretation of seep assemblage composition
from
cores, Powell et al. (1998) reported that, overall, seep communities were persistent over periods of 500-1,000
years
and probably throughout the entire Pleistocene.[9] Some sites retained optimal habitat over geological time
scales.[9]
Powell reported evidence of mussel and clam communities persisting in the same sites for 500-4,000 years.[9]
Powell
also found that both the composition of species and trophic tiering of hydrocarbon seep communities tend to be
fairly
constant across time, with temporal variations only in numerical abundance.[9] He found few cases in which the
Cold seep 96
community type changed (from mussel to clam communities, for example) or had disappeared completely.[9]
Faunal
succession was not observed.[9] Surprisingly, when recovery occurred after a past destructive event, the same
chemosynthetic species reoccupied a site.[9] There was little evidence of catastrophic burial events, but two
instances
were found in mussel communities in Green Canyon Block 234.[9] The most notable observation reported by
Powell
(1995) was the uniqueness of each chemosynthetic community site.[9]
Precipitation of authigenic carbonates and other geologic events will undoubtedly alter surface seepage patterns
over
periods of many years, although through direct observation, no changes in chemosynthetic fauna distribution or
composition were observed at seven separate study sites (MacDonald et al., 1995).[9] A slightly longer period
(19
years) can be referenced in the case of Bush Hill, the first Central Gulf of Mexico community described in situ
in
1986.[9] No mass die-offs or large-scale shifts in faunal composition have been observed (with the exception of
collections for scientific purposes) over the19-year history of research at this site.[9]
All chemosynthetic communities are located in water depths beyond the impact of severe storms, including
hurricanes, and there would have been no alteration of these communities caused from surface storms, including
hurricanes.[9]
Biology
The mussel species Bathymodiolus childressi is
the dominant species in the mytilid type of cold
seep communities in the Gulf of Mexico.
MacDonald et al. (1990) has described four general community
types.[9] These are communities dominated by Vestimentiferan tube
worms (Lamellibrachia c.f. barhami and Escarpia spp.), mytilid
mussels (Seep Mytilid Ia, Ib, and III, and others), vesicomyid clams
(Vesicomya cordata and Calyptogena ponderosa), and infaunal lucinid
or thyasirid clams (Lucinoma sp. or Thyasira sp.).[9] Bacterial mats are
present at all sites visited to date.[9] These faunal groups tend to display
distinctive characteristics in terms of how they aggregate, the size of
aggregations, the geological and chemical properties of the habitats in
which they occur and, to some degree, the heterotrophic fauna that
occur with them.[9] Many of the species found at these cold seep
communities in the Gulf of Mexico are new to science and remain
undescribed.[9]
Individual lamellibranchid tube worms, the longer of two taxa found at seeps can reach lengths of 3 m (10 ft)
and
live hundreds of years (Fisher et al., 1997; Bergquist et al., 2000).[9] Growth rates determined from recovered
marked tube worms have been variable, ranging from no growth of 13 individuals measured one year to a
maximum
growth of 9.6 cm/yr (3.8 in/yr) in a Lamellibrachia individual (MacDonald, 2002).[9] Average growth rate
was
2.19 cm/yr (0.86 in/yr) for the Escarpia-like species and 2.92 cm/yr (1.15 in/yr) for lamellibrachids.[9] These
are
slower growth rates than those of their hydrothermal vent relatives, but Lamellibrachia individuals can reach
lengths
2-3 times that of the largest known hydrothermal vent species.[9] Individuals of Lamellibrachia sp. in excess
of 3 m
(10 ft) have been collected on several occasions, representing probable ages in excess of 400 years (Fisher,
1995).[9]
Vestimentiferan tube worm spawning is not seasonal and recruitment is episodic.[9]
Tubeworms are either male or female. One recent discovery indicates that the spawning of female
Lamellibrachia
appears to have produced a unique association with the large bivalve Acesta bullisi, which lives permanently
attached to the anterior tube opening of the tubeworm, and feeds on the periodic egg release (Järnegren et al.,
2005).[9] This close association between the bivalves and tubeworms was discovered in 1984 (Boland, 1986) but
not
fully explained.[9] Virtually all mature Acesta individuals are found on female rather than male tubeworms.[9]
This
evidence and other experiments by Järnegren et al. (2005) seem to have solved this mystery.[9]
Cold seep 97
Growth rates for methanotrophic mussels at cold seep sites have been reported (Fisher, 1995).[9] General growth
rates were found to be relatively high.[9] Adult mussel growth rates were similar to mussels from a littoral
environment at similar temperatures.[9] Fisher also found that juvenile mussels at hydrocarbon seeps initially
grow
rapidly, but the growth rate drops markedly in adults; they grow to reproductive size very quickly.[9] Both
individuals and communities appear to be very long lived.[9] These methane-dependent mussels have strict
chemical
requirements that tie them to areas of the most active seepage in the Gulf of Mexico.[9] As a result of their rapid
growth rates, mussel recolonization of a disturbed seep site could occur relatively rapidly.[9] There is some
evidence
that mussels also have some requirement of a hard substrate and could increase in numbers if suitable substrate is
increased on the seafloor (Fisher, 1995).[9] Two associated species are always found associated with mussel beds
–
the gastropod Bathynerita naticoidea and a small Alvinocarid shrimp – suggesting these endemic species
have
excellent dispersal abilities and can tolerate a wide range of conditions (MacDonald, 2002).[9]
Unlike mussel beds, chemosynthetic clam beds may persist as a visual surface phenomenon for an extended
period
without input of new living individuals because of low dissolution rates and low sedimentation rates.[9] Most
clam
beds investigated by Powell (1995) were inactive.[9] Living individuals were rarely encountered.[9] Powell
reported
that over a 50-year timespan, local extinctions and recolonization should be gradual and exceedingly rare.[9]
Contrasting these inactive beds, the first community discovered in the Central Gulf of Mexico consisted of
numerous
actively plowing clams.[9] The images obtained of this community were used to develop length/frequency and
live/dead ratios as well as spatial patters (Rosman et al., 1987a).[9]
Extensive bacterial mats of free-living bacteria are also evident at all hydrocarbon seep sites.[9] These bacteria
may
compete with the major fauna for sulfide and methane energy sources and may also contribute substantially to
overall production (MacDonald, 1998b).[9] The white, nonpigmented mats were found to be an autotrophic
sulfur
bacteria Beggiatoa species, and the orange mats possessed an unidentified nonchemosynthetic metabolism
(MacDonald, 1998b).[9]
Heterotrophic species at seep sites are a mixture of species unique to seeps (particularly molluscs and crustacean
invertebrates) and those that are a normal component from the surrounding environment.[9] Carney (1993) first
reported a potential imbalance that could occur as a result of chronic disruption.[9] Because of sporadic
recruitment
patterns, predators could gain an advantage, resulting in exterminations in local populations of mussel beds.[9] It
is
clear that seep systems do interact with the background fauna but conflicting evidence remains as to what degree
outright predation on some specific community components such as tubeworms occurs (MacDonald, 2002).[9]
The
more surprising results from this recent work is why background species do not utilize seep production more
than
seems to be evident.[9] In fact, seep-associated consumers such as galatheid crabs and nerite gastropods had
isotopic
signatures, indicating that their diets were a mixture of seep and background production.[9] At some sites,
endemic
seep invertebrates that would have been expected to obtain much if not all their diet from seep production
actually
consumed as much as 50 percent of their diets from the background.[9]
Cold seep 98
In the Atlantic Ocean
Map of cold seeps in in the Atlantic Equatorial
Belt.
BR - Blake Ridge diapir
BT - Barbados trench
OR - Orenoque sectors
EP - El Pilar sector
NIG - Nigerian slope
GUI - Guiness area
REG - Regab pockmark.
Cold-seep communities in the western Atlantic Ocean have also been
described from a few dives on mud volcanoes and diapirs between
1000 and 5000 m depth in the Barbados accretionary prism area and
from the Blake Ridge diapir off North Carolina.[6] More recently seep
communities have been discovered in the eastern Atlantic, on a giant
pockmark cluster in the Gulf of Guinea near the Congo deep channel,
also on other pockmarks of the Congo margin, Gabon margin and
Nigeria margin and in the Gulf of Cadiz.[6]
Cold seeps are also available in Northern Atlantic Ocean.[1] (map
shows three localities there [11])
Extensive faunal sampling has been conducted from 400 to 3300 m in
the Atlantic Equatorial Belt from the Gulf of Mexico to the Gulf of
Guinea including Barbados accretionary prism, the Blake Ridge diapir,
and in the Eastern Atlantic from the Congo and Gabon margins and the recently explored Nigeria margin during
Census of Marine Life ChEss project.[6] Of the 72 taxa identified at the species level, a total of 9 species or
species
complexes are identified as amphi-Atlantic.[6]
The Atlantic Equatorial Belt seep megafauna community structure is influenced primarily by depth rather than
by
geographic distance.[6] The bivalves Bathymodiolinae (within Mytilidae) species or complexes of species are the
most widespread in the Atlantic.[6] The Bathymodiolus boomerang complex is found at the Florida
escarpment site,
the Blake Ridge diapir, the Barbados prism and the Regab site of Congo.[6] The Bathymodiolus childressi
complex is
also widely distributed along the Atlantic Equatorial Belt from the Gulf of Mexico across to the Nigerian
Margin,
although not on the Regab or Blake ridge sites.[6] The commensal polynoid, Branchipolynoe seepensis is
known
from the Gulf of Mexico, Gulf of Guinea and Barbados.[6] Other species with distributions extending from the
eastern to western Atlantic are: gastropod Cordesia provannoides, the shrimp Alvinocaris muricola, the
galatheids
Munidopsis geyeri and Munidopsis livida and probably the holothurid Chiridota heheva.[6]
There have been found cold seeps also in Amazon deepsea fan. High-resolution seismic profiles near the shelf
edge
show evidence of near-surface slumps and faulting 20–50 m in the subsurface and concentrations (about 500 m2)
of
methane gas. Several studies (e.g., Amazon Shelf Study—AMASEDS, LEPLAC, REMAC, GLORIA, Ocean
Drilling Program) indicate that there is evidence for gas seepage on the slope off the Amazon fan based on the
incidence of bottom-simulating reflections (BSRs), mud volcanoes, pock marks, gas in sediments, and deeper
hydrocarbon occurrences. The existence of methane at relatively shallow depths and extensive areas of gas
hydrates
have been mapped in this region. Also, gas chimneys have been reported, and exploratory wells have discovered
subcommercial gas accumulations and pock marks along fault planes. A sound geological and geophysical
understanding of the Foz do Amazonas Basin is already available and used by the energy companies.[12]
Exploration of new areas, such as potential seep sites off of the east coast of the U.S. and the Laurentian fan
where
chemosynthetic communities are known deeper than 3500 m, and shallower sites in the Gulf of Guinea are need
to
study in the future.[6]
Cold seep 99
In the Mediterranean
The first biological evidence for reduced environments in the Mediterranean Sea was the presence of Lucinidae
and
Vesicomyidae bivalve shells cored on the top of the Napoli mud volcano, located at 1,900 m depth on the
Mediterranean Ridge in the subduction zone of the African plate. This was followed by the description of a new
Lucinidae bivalve species, Lucinoma kazani, associated with bacterial endosymbionts. In the southeastern
Mediterranean, communities of polychaetes and bivalves were also found associated with cold seeps and
carbonates
near Egypt and the Gaza Strip at depths of 500–800 m, but no living fauna was collected. The first in situ
observations of extensive living chemosynthetic communities in the Eastern Mediterranean Sea prompted
cooperation between biologists, geochemists, and geologists. During submersible dives, communities comprising
large fields of small bivalves (dead and alive), large siboglinid tube worms, isolated or forming dense
aggregations,
large sponges, and associated endemic fauna were observed in various cold seep habitats associated with
carbonate
crusts at 1,700–2,000 m depth. Two mud volcano fields were first explored, one along the Mediterranean Ridge,
where most of them were partially (Napoli, Milano mud volcanoes) or totally (Urania, Maidstone mud
volcanoes)
affected by brines, and the other on the Anaximander mounds south of Turkey. The latter area includes the large
Amsterdam mud volcano, which is affected by recent mudflows, and the smaller Kazan or Kula mud volcanoes.
Gas
hydrates have been sampled at the Amsterdam and Kazan mud volcanoes, and high methane levels have been
recorded above the seafloor. Several provinces of the Nile deep-sea fan have been explored recently. These
include
the very active brine seepage named the Menes Caldera in the eastern province between 2,500 m and 3,000 m,
the
pockmarks in the central area along mid- and lower slopes, and the mud volcanoes of the eastern province, as
well as
one in the central upper slope (North Alex area) at 500 m depth.[13]
During these first exploratory dives, symbiont-bearing taxa that are similar to those observed on the Olimpi and
Anaximander mud fields were sampled and identified. This similarity is not surprising, as most of these taxa
were
originally described from dredging in the Nile fan.[13] Up to five species of bivalves harboring bacterial
symbionts
colonized these methane- and sulfide-rich environments. A new species of Siboglinidae polychaete,
Lamellibrachia
anaximandri, the tubeworm colonizing cold seeps from the Mediterranean ridge to the Nile deep-sea fan, has
just
been described in 2010.[13] [14] Moreover, the study of symbioses revealed associations with chemoautotrophic
Bacteria, sulfur oxidizers in Vesicomyidae and Lucinidae bivalves and Siboglinidae tubeworms, and highlighted
the
exceptional diversity of Bacteria living in symbiosis with small Mytilidae. The Mediterranean seeps appear to
represent a rich habitat characterized by megafauna species richness (e.g., gastropods) or the exceptional size of
some species such as sponges (Rhizaxinella pyrifera) and crabs (Chaceon mediterraneus), compared
with their
background counterparts. This contrasts with the low macro- and mega-faunal abundance and diversity of the
deep
Eastern Mediterranean. Seep communities in the Mediterranean that include endemic chemosynthetic species
and
associated fauna differ from the other known seep communities in the world at the species level but also by the
absence of the large size bivalve genera Calyptogena or Bathymodiolus. The isolation of the
Mediterranean seeps
from the Atlantic Ocean after the Messinian crisis led to the development of unique communities, which are
likely to
differ in composition and structure from those in the Atlantic Ocean. Further expeditions involved quantitative
sampling of habitats in different areas, from the Mediterranean Ridge to the eastern Nile deep-sea fan.[13] Cold
seeps
discovered in the Sea of Marmara in 2008[15] have also revealed chemosynthesis-based communities that
showed a
considerable similarity to the symbiont-bearing fauna of eastern Mediterranean cold seeps.[13]
Cold seep 100
In the West Pacific
Native aluminium has been reported also in cold seeps in the northeastern continental slope of the South China
Sea
and Chen et al. (2011)[16] have proposed a theory of its origin as resulting by reduction from
tetrahydroxoaluminate
Al(OH)4
- to metallic aluminium by bacteria.[16]
Japan
Methane seep communities in Japan are distributed along plate convergence areas because of the accompanying
tectonic activity.[17] Many seeps have been found in the Japan Trench, Nankai Trough, Ryukyu Trench, Sagami
Bay
and Suruga Bay, and the Sea of Japan.[17] Many of species in cold seeps of Japan are endemic.[17]
New Zealand
Off the mainland coast of the New Zealand, shelf-edge instability is enhanced in some locations by cold seeps of
methane-rich fluids that likewise support chemosynthetic faunas and carbonate concretions.[18] [19] [20] [21]
Dominant
animals are tube worms of the family Siboglinidae and bivalves of families Vesicomyidae and Mytilidae
(Bathymodiolus).[21] Numerous of its species appear to be endemic.[21] The deep bottom trawling has highly
negative impact on cold seep communities and those ecosystems are threatened.[21] Depths down to 2,000 m
including cold seeps belongs among as-yet-unmapped topographic and chemical complexity of habitats.[18] The
scale of new-species discovery in these poorly studied or unexplored ecosystems is likely to be high.[18]
In the East Pacific
Monterey Bay Aquarium Research Institute has
used remotely operated underwater vehicle
Ventana in the research of Monterey Bay cold
seeps.
In the deep sea the COMARGE project has studied the biodiversity
patterns along and across the Chilean margin through a complexity of
ecosystems such as methane seeps and oxygen minimum zones
reporting that such habitat heterogeneity may influence the biodiversity
patterns of the local fauna.[12] [22] [23] [24] Seep fauna include bivalves
of families Lucinidae, Thyasiridae, Solemyidae (Acharax sp.), and
Vesicomyidae (Calyptogena gallardoi) and polychates Lamellibrachia
sp. and two other polychate species.[23] Furthermore, in these soft
reduced sediments below the oxygen minimum zone off the Chilean
margin, a diverse microbial community composed by a variety of large
prokaryotes (mainly large multi-cellular filamentous “mega bacteria” of
the genera Thioploca and Beggiatoa, and of “macrobacteria” including
a diversity of phenotypes), protists (ciliates, flagellates, and
foraminifers), as well as small metazoans (mostly nematodes and
polychaetes) has been found.[12] [25] Gallardo et al. (2007)[25] argue
that the likely chemolithotrophic metabolism of most of these megaand
macrobacteria offer an alternative explanation to fossil findings, in
particular to those from obvious non-littoral origins, suggesting that
traditional hypotheses on the cyanobacterial origin of some fossils may
have to be revised.[12]
Cold seeps (pockmark) are also known from depths of 130 m in the Hecate Strait, British Columbia, Canada.[26]
Unobvious fauna (also unobvious for cold seeps) has been found there with these dominating species: sea snail
Fusitriton oregonensis, anemone Metridium giganteum, encrusting sponges and bivalve Solemya
reidi.[26]
Cold seeps with chemosynthetic communities along the USA Pacific coast occur in Monterey Canyon, just off
Monterey Bay, California on a mud volcano.[27] There have been found, for example Calyptogena clams
Cold seep 101
Calyptogena kilmeri and Calyptogena pacifica[28] and foraminiferan Spiroplectammina biformis.
[29]
• map of cold seeps in the Monterey Bay [30]
In the Antarctic
The relatively few investigations to the Antarctic deep sea have shown the presence of deep-water habitats,
including
hydrothermal vents, cold seeps, and mud volcanoes.[31] Other than the Antarctic Benthic Deep-Sea Biodiversity
Project (ANDEEP) cruises, little work has been done in the deep sea.[31] There are more species waiting to be
described.[31]
Detection
With continuing experience, particularly on the upper continental slope in the Gulf of Mexico, the successful
prediction of the presence of tubeworm communities continues to improve, however chemosynthetic
communities
cannot be reliably detected directly using geophysical techniques.[9] Hydrocarbon seeps that allow
chemosynthetic
communities to exist do modify the geological characteristics in ways that can be remotely detected, but the time
scales of co-occurring active seepage and the presence of living communities is always uncertain.[9] These
known
sediment modifications include (1) precipitation of authigenic carbonate in the form of micronodules, nodules, or
rock masses; (2) formation of gas hydrates; (3) modification of sediment composition through concentration of
hard
chemosynthetic organism remains (such as shell fragments and layers); (4) formation of interstitial gas bubbles
or
hydrocarbons; and (5) formation of depressions or pockmarks by gas expulsion.[9] These features give rise to
acoustic effects such as wipeout zones (no echoes), hard bottoms (strongly reflective echoes), bright spots
(reflection
enhanced layers), or reverberant layers (Behrens, 1988; Roberts and Neurauter, 1990).[9] "Potential" locations for
most types of communities can be determined by careful interpretation of these various geophysical
modifications,
but to date, the process remains imperfect and confirmation of living communities requires direct visual
techniques.[9]
Fossilized records
Late Cretaceous cold seep deposit in the Pierre
Shale, southwest South Dakota.
Cold seep deposits are found throughout the Phanerozoic rock record,
especially in the Late Mesozoic and Cenozoic (see for example Kaim
et al., 2008).[32] These fossil cold seeps are characterized by
mound-like topography (where preserved), coarsely crystalline
carbonates, and abundant mollusks and brachiopods.
References
This article incorporates a public domain work of the United States
Government from references[2] [9] and CC-BY-2.5 from references[1]
[3] [6] [12] [13] [17] [18] [31]
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Biogeography of Free-Living Nematodes Associated with Chemosynthetic Environments in the Deep Sea: A Review". PLoS ONE 5(8):
e12449. doi:10.1371/journal.pone.0012449.
[2] Hsing P.-Y. (October 19, 2010). "Gas-powered Circle of Life – Succession in a Deep-sea Ecosystem" (http:/ / oceanexplorer. noaa. gov/
explorations/ 10lophelia/ logs/ oct18/ oct18. html). NOAA Ocean Explorer | Lophelia II 2010: Oil Seeps and Deep Reefs | October 18 Log.
Retrieved 25 January 2011.
[3] Boetius A. (2005). "Microfauna–Macrofauna Interaction in the Seafloor: Lessons from the Tubeworm". PLoS Biology 3(3): e102.
doi:10.1371/journal.pbio.0030102
[4] Hovland M. & Thomsen E. (1997) "Cold-water corals—are they hydrocarbon seep related?" Marine Geology 137(1-2): 159-164.
doi:10.1016/S0025-3227(96)00086-2.
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[5] Hovland M. (2008). Deep-water coral reefs: unique biodiversity hot-spots. 8.10 Summary and re-iteration of the hydraulic
theory. Springer,
278 pp. ISBN 978-1-4020-8461-4. Pages 204 (http:/ / books. google. com/ books?id=CIc7w6SuhY8C& lpg=PA204& ots=Y4gU_7bLi4&
dq=hydraulic theory coral& hl=cs& pg=PA204#v=onepage& q=hydraulic theory coral& f=false)-205.
[6] Olu K., Cordes E. E., Fisher C. R., Brooks J. M., Sibuet M. & Desbruyčres D. (2010). "Biogeography and Potential Exchanges Among
the
Atlantic Equatorial Belt Cold-Seep Faunas". PLoS ONE 5(8): e11967. doi:10.1371/journal.pone.0011967.
[7] "Demise of Antarctic Ice Shelf Reveals New Life" (http:/ / www. nsf. gov/ discoveries/ disc_summ. jsp?cntn_id=109683). National
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Foundation. 2007. . Retrieved 2008-02-14.
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journal.
pone. 0010363. g001
[9] Minerals Management Service Gulf of Mexico OCS Region (November 2006). "Gulf of Mexico OCS Oil and Gas Lease Sales: 2007-
2012.
Western Planning Area Sales 204, 207, 210, 215, and 218. Central Planning Area Sales 205, 206, 208, 213, 216, and 222. Draft
Environmental
Impact Statement. Volume I: Chapters 1-8 and Appendices". U.S. Department of the Interior, Minerals Management Service, Gulf of Mexico
OCS Region, New Orleans. page 3-27 - 3-31 PDF (http:/ / www. gomr. mms. gov/ PDFs/ 2006/ 2006-062-Vol1. pdf).
[10] http:/ / www. gomr. mms. gov/ PI/ PDFImages/ ESPIS/ 2/ 3071. pdf
[11] http:/ / www. plosone. org/ article/ slideshow. action?uri=info:doi/ 10. 1371/ journal. pone. 0012449& imageURI=info:doi/ 10. 1371/
journal. pone. 0012449. g001
[12] Miloslavich P., Klein E., Díaz J. M., Hernández C. E., Bigatti G. et al. (2011). "Marine Biodiversity in the Atlantic and Pacific Coasts of
South America: Knowledge and Gaps". PLoS ONE 6(1): e14631. doi:10.1371/journal.pone.0014631.
[13] Danovaro R., Company J. B., Corinaldesi C., D'Onghia G., Galil B. et al. (2010). "Deep-Sea Biodiversity in the Mediterranean Sea: The
Known, the Unknown, and the Unknowable". PLoS ONE 5(8): e11832. doi:10.1371/journal.pone.0011832.
[14] Southward E., Andersen A., Hourdez S. (submitted 2010). "Lamellibrachia anaximandri n.sp., a new vestimentiferan tubeworm
from the
Mediterranean (Annelida)". Zoosystema.
[15] Zitter T. A. C, Henry P., Aloisi G., Delaygue G., Çagatay M. N. et al. (2008). "Cold seeps along the main Marmara Fault in the Sea of
Marmara (Turkey)". Deep Sea Research Part I: Oceanographic Research Papers 55(4): 552-570.
doi:10.1016/j.dsr.2008.01.002.
[16] Chen Z., Huang C.-Y., Zhao M., Yan W., Chien C.-W., Chen M., Yang H., Machiyama H. & Lin S. (2011). "Characteristics and
possible
origin of native aluminum in cold seep sediments from the northeastern South China Sea". Journal of Asian Earth Sciences 40(1):
363-370.
doi:doi:10.1016/j.jseaes.2010.06.006.
[17] Fujikura K., Lindsay D., Kitazato H., Nishida S. & Shirayama Y. (2010). "Marine Biodiversity in Japanese Waters". PLoS ONE 5(8):
e11836. doi:10.1371/journal.pone.0011836.
[18] Gordon D. P., Beaumont J., MacDiarmid A., Robertson D. A. & Ahyong S. T (2010). "Marine Biodiversity of Aotearoa New Zealand".
PLoS ONE 5(8): e10905. doi:10.1371/journal.pone.0010905.
[19] Lewis K. B. & Marshall B. A. (1996). "Seep faunas and other indicators of methane-rich dewatering on New Zealand convergent
margins".
New Zealand Journal of Geology and Geophysics 39(2): 181-200. doi:10.1080/00288306.1996.9514704. PDF (http:/ / www.
informaworld.
com/ smpp/ ftinterface~content=a920227278~fulltext=713240930~frm=content).
[20] Orpin A. R. (1997). "Dolomite chimneys as possible evidence of coastal fluid expulsion, uppermost Otago continental slope, southern
New
Zealand". Marine Geology 138(1-2): 51–67. doi:10.1016/S0025-3227(96)00101-6.
[21] Baco A. R., Rowden A. A., Levin L. A., Smith C. R., Bowden D. et al. (2009). "Initial characterization of cold seep faunal communities
on
the New Zealand margin". Marine Geology 272(1-4): 251-259. doi:10.1016/j.margeo.2009.06.015.
[22] Sellanes J., Neira C., Quiroga E. & Teixido N. (2010). "Diversity patterns along and across the Chilean margin: a continental slope
encompassing oxygen gradients and methane seep benthic habitats". Marine Ecology 31(1): 111-124. doi:10.1111/j.1439-
0485.2009.00332.x.
[23] Sellanes J., Quiroga E. & Neira C. (2008). "Megafauna community structure and trophic relationships at the recently discovered
Concepción
Methane Seep Area, Chile, ~36°S". ICES Journal of Marine Science 65(7): 1102-1111. doi:10.1093/icesjms/fsn099.
[24] Sellanes J., Quiroga E., Gallardo V. A. (2004). "First direct evidence of methane seepage and associated chemosynthetic communities in
the
bathyal zone off Chile". Journal of the Marine Biological Association of the UK 84(5): 1065-1066.
doi:10.1017/S0025315404010422h.
[25] Gallardo V. A. & Espinoza C. (2007). "Large multicellular filamentous bacteria under the oxygen minimum zone of the eastern South
Pacific: a forgotten biosphere". In: Hoover R. B., Levin G. V., Rozanov A. Y. & Davies P. C. W. (eds). San Diego, CA, USA: Proc. SPIE
6694: 66941H–11. doi:10.1117/12.782209.
[26] Barrie J. V., Cook S. & Conway K. W. (available online 4 March 2010). "Cold seeps and benthic habitat on the Pacific margin of
Canada".
Continental Shelf Research 31(2) Supplement 1: S85-S92. doi:10.1016/j.csr.2010.02.013.
[27] Lorenson T. D., Kvenvolden K. A., Hostettler F. D., Rosenbauer R. J., Martin J. B. & Orange D. L. (1999). "Hydrocarbons Associated
with
Fluid Venting Process in Monterey Bay, California" (http:/ / walrus. wr. usgs. gov/ monterey/ hydrocarbons/ sitemap. html). USGS Pacific
Coastal & Marine Science Center.
[28] Goffredi S. K. & Barry J. P. (2000). "Factors regulating productivity in chemoautotrophic symbioses; with emphasis on Calyptogena
kilmeri
and Calyptogena pacifica". Poster, Monterey Bay Aquarium Research Institute. accessed 3 February 2011. PDF (http:/ / www. mbari.
org/
benthic/ sposter. pdf).
[29] Bernhard J. M., Buck K. R. & Barry J. P. (2001). "Monterey Bay cold-seep biota: Assemblages, abundance, and ultrastructure of living
foraminifera". Deep Sea Research Part I: Oceanographic Research Papers 48(10): 2233-2249. doi:10.1016/S0967-
0637(01)00017-6.
[30] http:/ / www. mbari. org/ benthic/ coldseeploc. htm
Cold seep 103
[31] Griffiths H. J. (2010). "Antarctic Marine Biodiversity – What Do We Know About the Distribution of Life in the Southern Ocean?".
PLoS
ONE 5(8): e11683. doi:10.1371/journal.pone.0011683.
[32] Kaim A., Jenkins R. & Warén A. (2008). "Provannid and provannid-like gastropods from the Late Cretaceous cold seeps of Hokkaido
(Japan) and the fossil record of the Provannidae (Gastropoda: Abyssochrysoidea)". Zoological Journal of the Linnean Society
154(3):
421-436. doi:10.1111/j.1096-3642.2008.00431.x.
Further reading
• Bright M., Plum C., Riavitz L. A., Nikolov N., Martínez Arbizu P. & Cordes E. E. & Gollner S. (2010).
"Epizooic
metazoan meiobenthos associated with tubeworm and mussel aggregations from cold seeps of the Northern Gulf
of Mexico". Deep Sea Research Part II: Topical Studies in Oceanography 57(21-23): 1982-1989.
doi:10.1016/j.dsr2.2010.05.003.
• Lloyd K. G., Albert D. B., Biddle J. F., Chanton J. P., Pizarro O. & Teske A. (2010). "Spatial Structure and
Activity of Sedimentary Microbial Communities Underlying a Beggiatoa spp. Mat in a Gulf of Mexico
Hydrocarbon Seep". PLoS ONE 5(1): e8738. doi:10.1371/journal.pone.0008738.
• Metaxas A. & Kelly N. E. (2010). "Do Larval Supply and Recruitment Vary among Chemosynthetic
Environments of the Deep Sea?". PLoS ONE 5(7): e11646. doi:10.1371/journal.pone.0011646.
• Rodríguez E. & Daly M. (2010). "Phylogenetic Relationships among Deep-Sea and Chemosynthetic Sea
Anemones: Actinoscyphiidae and Actinostolidae (Actiniaria: Mesomyaria)". PLoS ONE 5(6): e10958.
doi:10.1371/journal.pone.0010958.
• Sibuet M. & Olu K. (1998). "Biogeography, biodiversity and fluid dependence of deep-sea cold-seep
communities at active and passive margins". Deep Sea Research Part II: Topical Studies in
Oceanography
45(1-3): 517-567. doi:10.1016/S0967-0645(97)00074-X.
External links
• Paul Yancy's vents and seeps page (http:/ / people. whitman. edu/ ~yancey/ califseeps. html)
• Monterey Bay Aquarium Research Institute's seeps page (http:/ / www. mbari. org/ benthic/ coldseeps. htm)
• ScienceDaily News: Tubeworms in deep sea discovered to have record long life spans (http:/ / www.
sciencedaily. com/ releases/ 2000/ 02/ 000203075002. htm)
Benthic zone 104
Benthic zone
The benthic zone is the ecological region at the lowest level of a body of water such as an ocean or a lake,
including
the sediment surface and some sub-surface layers. Organisms living in this zone are called benthos. They
generally
live in close relationship with the substrate bottom; many such organisms are permanently attached to the
bottom.
The superficial layer of the soil lining the given body of water, the benthic boundary layer, is an integral part of
the
benthic zone, as it influences greatly the biological activity which takes place there. Examples of contact soil
layers
include sand bottoms, rocky outcrops, coral, and bay mud.
Description
The benthic region begins at the shore line (intertidal or eulittoral zone) and extends downward along the surface
of
the continental shelf out to sea. The continental shelf is a gently sloping benthic region that extends away from
the
land mass. At the continental shelf edge, usually about 200 meters deep, the gradient greatly increases and is
known
as the continental slope. The continental slope drops down to the deep sea floor. The deep-sea floor is called the
abyssal plain and is usually about 4000 meters deep. The ocean floor is not all flat but has submarine ridges and
deep
ocean trenches known as the hadal zone.
For comparison, the pelagic zone is the descriptive term for the ecological region above the benthos, including
the
water-column up to the surface. Depending on the water-body, the benthic zone may include areas which are
only a
few inches below water, such as a stream or shallow pond; at the other end of the spectrum, benthos of the deep
ocean includes the bottom levels of the oceanic abyssal zone.
For information on animals that live in the deeper areas of the oceans see aphotic zone. Generally, these include
life
forms that tolerate cool temperatures and low oxygen levels, but this depends on the depth of the water.
Organisms
Benthos are the organisms which live in the benthic zone, and are different from those elsewhere in the water
column. Many are adapted to live on the substrate (bottom). In their habitats they can be considered as dominant
creatures. Many organisms adapted to deep-water pressure cannot survive in the upper parts of the water column.
The pressure difference can be very significant (approximately one atmosphere for each 10 meters of water
depth).
Because light does not penetrate very deep ocean-water, the energy source for the benthic ecosystem is often
organic
matter from higher up in the water column which drifts down to the depths. This dead and decaying matter
sustains
the benthic food chain; most organisms in the benthic zone are scavengers or detritivores. Some microorganisms
use
chemosynthesis to produce biomass.
Benthic organisms can be divided into two categories based on whether they make their home on the ocean floor
or
an inch or two into the ocean floor. Those living on the surface of the ocean floor are known as epifauna.[1]
Those
who live burrowed into the ocean floor are known as infauna.[2]
Benthic zone 105
Habitats
In oceanic environments, benthic habitats can be further zoned by depth. From the shallowest to the deepest are:
the
epipelagic (less than 200 meters), the mesopelagic (200–1000 metres), the bathyal (1000–4000 meters), the
abyssal
(4000–6000 meters) and the deepest, the hadal (below 6000 meters).
The lower zones are in deep, pressurized areas of the ocean. Because of the high pressures and seclusion neither
tidal
changes nor human impacts have had much of an effect on these areas, and the habitats have not changed much
over
the years. Many benthic organisms have retained their historic evolutionary characteristics. Some organisms are
significantly larger than their relatives living in shallower zones, largely because of higher oxygen concentration
in
deep water.[3]
It is not easy to map or observe these organisms and their habitats, and most observation has been done through
remote controlled submarines.
References
[1] http:/ / www. merriam-webster. com/ dictionary/ epifaunal
[2] http:/ / www. merriam-webster. com/ dictionary/ epifaunal
[3] Royal Belgian Institute of Natural Sciences, news item March 2005 (http:/ / www. naturalsciences. be/ active/ sciencenews/ archive2005/
polarregions/ page3/ document_view)
External links
• Data Archive for Seabed Species and Habitats (http:/ / www. dassh. ac. uk) from the UK Marine Data Archive
Centre
Kelp forest
Kelp forests are underwater areas with a high density of kelp. They are recognized as one of the most
productive
and dynamic ecosystems on Earth.[1] Smaller areas of anchored kelp are called kelp beds.
Kelp forests occur worldwide throughout temperate and polar coastal oceans.[1] In 2007, kelp forests were also
discovered in tropical waters near Ecuador.[2]
Physically formed by brown macroalgae of the order Laminariales, kelp forests provide a unique three-
dimensional
habitat for marine organisms and are a source for understanding many ecological processes. Over the last
century,
they have been the focus of extensive research, particularly in trophic ecology, and continue to provoke
important
ideas that are relevant beyond this unique ecosystem. For example, kelp forests can influence coastal
oceanographic
patterns[3] and provide many ecosystem services.[4]
However, the influence of humans has often contributed to kelp forest degradation. Of particular concern are the
effects of overfishing nearshore ecosystems, which can release herbivores from their normal population
regulation
and result in the over-grazing of kelp and other algae.[5] This can rapidly result in transitions to barren
landscapes
where relatively few species persist.[6] The implementation of marine protected areas (MPAs) is one
management
strategy useful for addressing such issues since it may limit the impacts of fishing and buffer the ecosystem from
additive effects of other environmental stressors.
Kelp forest 106
Kelp
Rockfish swimming around giant kelp
A diver in a kelp forest off the coast of California
Giant kelp uses gas filled floats to keep the plant
suspended, allowing the kelp blades near the
ocean surface to capture light for photosynthesis.
The term 'kelp' refers to marine algae belonging to the taxonomic order
Laminariales (Phylum: Heterokontophyta). Though not considered a
taxonomically diverse order, kelps are highly diverse structurally and
functionally.[4] The most widely recognized species are the giant kelps
(Macrocystis spp.), although there are numerous other genera such as
Laminaria, Ecklonia, Lessonia, Alaria and Eisenia.
Frequently considered an ecosystem engineer, kelp provides a physical
substrate and habitat for kelp forest communities.[7] In algae
(Kingdom: Protista), the body of an individual organism is known as a
thallus rather than as a plant (Kingdom: Plantae). The morphological
structure of a kelp thallus is defined by three basic structural units:[6]
• The holdfast is a root-like mass that anchors the thallus to the sea
floor, though unlike true roots it is not responsible for absorbing
and delivering nutrients to the rest of the thallus;
• The stipe is analogous to a plant stalk, extending vertically from
the holdfast and providing a support framework for other
morphological features;
• The fronds are leaf- or blade-like attachments extending from the
stipe, sometimes along its full length, and are the sites of nutrient
uptake and photosynthetic activity.
In addition, many kelp species have pneumatocysts, or gas-filled
bladders, usually located at the base of fronds near the stipe. These
structures provide the necessary buoyancy for kelp to maintain an
upright position in the water column.
The environmental factors necessary for kelp to survive include hard
substrate (usually rock), high nutrients (e.g., nitrogen, phosphorus),
and light (minimum annual irradiance dose > 50 E m−2[8] ). Especially
productive kelp forests tend to be associated with areas of significant
oceanographic upwelling, a process that delivers cool nutrient-rich
water from depth to the ocean’s mixed surface layer.[8] Water flow and
turbulence facilitate nutrient assimilation across kelp fronds throughout
the water column.[9] Water clarity affects the depth to which sufficient
light can be transmitted. In ideal conditions, giant kelp (Macrocystis
spp.) can grow as much as 30-60 centimeters vertically per day. Some
species such as Nereocystis are annual while others like Eisenia are
perennial, living for more than 20 years.[10] In perennial kelp forests,
maximum growth rates occur during upwelling months (typically
spring and summer) and die-backs correspond to reduced nutrient
availability, shorter photoperiods and increased storm frequency.[6]
Kelps are primarily associated with temperate and arctic waters worldwide. Of the more dominant species,
Laminaria is mainly associated with both sides of the Atlantic Ocean and the coasts of
Kelp forest 107
Distribution of Kelp
China and Japan; Ecklonia is found in Australia, New Zealand, and
South Africa; and Macrocystis occurs throughout the northeastern and
southeastern Pacific Ocean, Southern Ocean archipelagos, and in
patches around Australia, New Zealand and South Africa.[6] The region
with the greatest diversity of kelps (>20 species) is the northeastern
Pacific, from north of San Francisco, California, to the Aleutian
Islands, Alaska.
Although kelp forests are unknown in tropical surface waters, a few
species of Laminaria have been known to occur exclusively in tropical deep waters.[11] [12] This general
absence of
kelp from the tropics is believed to be mostly due to insufficient nutrient levels associated with warm,
oligotrophic
waters.[6] One recent study spatially overlaid the requisite physical parameters for kelp with mean oceanographic
conditions has produced a model predicting the existence of subsurface kelps throughout the tropics worldwide
to
depths of 200 m. For a hotspot in the Galapagos Islands, the local model was improved with fine-scale data and
tested; the research team found thriving kelp forests in all 8 of their sampled sites, all of which had been
predicted by
the model and thus validated their approach. This suggests that their global model might actually be fairly
accurate
and if so, kelp forests would be prolific in tropical subsurface waters worldwide.[2] The importance of this
contribution has been rapidly acknowledged within the scientific community and prompts an entirely new
trajectory
of kelp forest research, particularly emphasizing the potential for a spatial refuge from climate change and
explanations to evolutionary patterns of kelps worldwide.[13]
Ecosystem architecture
Kelp forest exhibit at the Monterey Bay
Aquarium
The architecture of a kelp forest ecosystem is based on its physical
structure, which influences the associated species that define its
community structure. Structurally, the ecosystem includes three guilds
of kelp and two guilds occupied by other algae:[6]
• Canopy kelps include the largest species and often constitute
floating canopies that extend to the ocean surface (e.g.,
Macrocystis and Alaria);
• Stipitate kelps generally extend a few meters above the sea floor
and can grow in dense aggregations (e.g., Eisenia and Ecklonia);
• Prostrate kelps lie near and along the sea floor (e.g., Laminaria);
• The benthic assemblage is composed of other algal species (e.g.,
filamentous and foliose functional groups, articulated corallines)
and sessile organisms along the ocean bottom;
• Encrusting coralline algae directly and often extensively cover
geologic substrate.
Multiple kelp species often co-exist within a forest; the term
understory canopy refers to the stipitate and prostrate kelps. For
example, a Macrocystis canopy may extend many meters above the
seafloor towards the ocean surface, while an understory of the kelps
Eisenia and Pterygophora reaches upward only a few meters. Beneath
these kelps there may be a benthic assemblage of foliose red algae. The dense vertical infrastructure with
overlying
canopy forms a system of microenvironments similar to those observed in a terrestrial forest, with a sunny
canopy
region, a partially shaded middle, and darkened seafloor.[6] Each guild has associated organisms, which vary in
their
levels of dependence on the habitat, and the assemblage of these organisms can vary with kelp morphologies.[14]
[15]
Kelp forest 108
[16] For example, in California Macrocystis pyrifera forests, the nudibranch Melibe leonina and skeleton
shrimp
Caprella californica are closely associated with surface canopies; the kelp perch Brachyistius frenatus,
rockfish
Sebastes spp. and many other fishes are found within the stipitate understory; brittle stars and turban snails
Tegula
spp. are closely associated with the kelp holdfast, while various herbivores such as sea urchins and abalone live
under the prostrate canopy; many seastars, hydroids and benthic fishes live among the benthic assemblages;
solitary
corals, various gastropods and echinoderms live over the encrusting coralline algae.[14] In addition, pelagic
fishes
and marine mammals are loosely associated with kelp forests, usually interacting near the edges as they visit to
feed
on resident organisms.
Trophic ecology
Sea urchins like this purple sea urchin can
damage kelp forests by chewing through kelp
holdfasts
The sea otter is an important predator of sea
urchins.
The jeweled top snail Calliostoma annulatum
grazing on a blade of giant kelp
Classic studies in kelp forest ecology have largely focused on trophic
interactions (the relationships between organisms and their food webs),
particularly the understanding and top-down trophic processes.
Bottom-up processes are generally driven by the abiotic conditions
required for primary producers to grow, such as availability of light
and nutrients, and the subsequent transfer of energy to consumers at
higher trophic levels. For example, the occurrence of kelp is frequently
correlated with oceanographic upwelling zones, which provide
unusually high concentrations of nutrients to the local environment.[17]
[18] This allows kelp to grow and subsequently support herbivores,
which in turn support consumers at higher trophic levels.[19] By
contrast, in top-down processes, predators limit the biomass of species
at lower trophic levels through consumption. In the absence of
predation, these lower level species flourish because resources that
support their energetic requirements are non-limiting. In a well-studied
example from Alaskan kelp forests,[20] sea otters (Enhydra lutris)
control populations of herbivorous sea urchins through predation.
When sea otters are removed from the ecosystem (for example, by
human exploitation), urchin populations are released from predatory
control and grow dramatically. This leads to increased herbivore
pressure on local kelp stands. Deterioration of the kelp itself results in
the loss of physical ecosystem structure and subsequently, the loss of
other species associated with this habitat. In Alaskan kelp forest
ecosystems, sea otters are the keystone species that mediates this
trophic cascade. In Southern California, kelp forests persist without sea
otters and the control of herbivorous urchins is instead mediated by a
suite of predators including lobsters and large fishes. The effect of
removing one predatory species in this system differs from Alaska
because there is redundancy in the trophic levels and other predatory
species can continue to regulate urchins.[15] However, the removal of
multiple predators can effectively release urchins from predator
pressure and allow the system to follow trajectories towards kelp forest
degradation.[21] Similar examples exist in Nova Scotia,[22] South
Africa,[23] Australia[24] and Chile.[25] The relative importance of
top-down versus bottom-up control in kelp forest ecosystems and the
strengths of trophic interactions continue to be the subject of considerable scientific investigation.[26] [27] [28]
Kelp forest 109
The transition from macroalgal (i.e. kelp forest) to denuded landscapes dominated by sea urchins (or ‘urchin
barrens’)
is a widespread phenomenon,[29] [30] [4] [31] often resulting from trophic cascades like those described above; the
two
phases are regarded as alternative stable states of the ecosystem.[32] [33] The recovery of kelp forests from barren
states has been documented following dramatic perturbations, such as urchin disease or large shifts in thermal
conditions.[21] [34] [35] Recovery from intermediate states of deterioration is less predictable and depends on a
combination of abiotic factors and biotic interactions in each case.
Though urchins are usually the dominant herbivore, others with significant interaction strengths include seastars,
isopods, kelp crabs, and herbivorous fishes.[6] [26] In many cases, these organisms feed on kelp that has been
dislodged from substrate and drifts near the ocean floor rather than expend energy searching for intact thalli to
feed
on. When there is sufficient drift kelp, herbivorous grazers do not exert pressure on attached plants; when drift
subsidies are unavailable, grazers directly impact the physical structure of the ecosystem.[36] [37] Many studies in
Southern California have demonstrated that the availability of drift kelp specifically influences the foraging
behavior
of sea urchins.[38] [39] Drift kelp and kelp-derived particulate matter have also been important in subsidizing
adjacent
habitats, such as sandy beaches and the rocky intertidal.[40] [41] [42]
Patch dynamics
Another major area of kelp forest research has been directed at understanding the spatial-temporal patterns of
kelp
patches. Not only do such dynamics affect the physical landscape, but they also affect species that associate with
kelp for refuge or foraging activities.[14] [19] Large-scale environmental disturbances have offered important
insights
concerning mechanisms and ecosystem resilience. Examples of environmental disturbances include the
following:
• Acute and chronic pollution events have been shown to impact southern California kelp forests, though the
intensity of the impact seems to depend on both the nature of the contaminants and duration of exposure.[43]
[44] [45] [46] [47] Pollution can include sediment deposition and eutrophication from sewage, industrial
byproducts and contaminants like PCBs and heavy metals (for example, copper, zinc), runoff of
organophosphates from agricultural areas, anti-fouling chemicals used in harbors and marinas (for example,
TBT and creosote) and land-based pathogens like fecal coliform bacteria.
• Catastrophic storms can remove surface kelp canopies through wave activity but usually leave understory
kelps intact; they can also remove urchins when little spatial refuge is available.[32] [37] Interspersed canopy
clearings create a seascape mosaic where sunlight penetrates deeper into the kelp forest and species that are
normally light-limited in the understory can flourish. Similarly, substrate cleared of kelp holdfasts can provide
space for other sessile species to establish themselves and occupy the seafloor, sometimes directly competing
with juvenile kelp plants and even inhibiting their settlement.[48]
• El Nińo-Southern Oscillation (ENSO) events involve the depression of oceanographic thermoclines, severe
reductions of nutrient input, and changes in storm patterns.[32] [49] Stress due to warm water and nutrient
depletion can increase the susceptibility of kelp to storm damage and herbivorous grazing, sometimes even
prompting phase shifts to urchin-dominated landscapes.[35] [38] [50] In general, oceanographic conditions (that
is, water temperature, currents) influence the recruitment success of kelp and its competitors, which clearly
affect subsequent species interactions and kelp forest dynamics.[32] [51]
• Overfishing higher trophic levels that naturally regulate herbivore populations is also recognized as an
important stressor in kelp forests.[5] [28] [52] As described in the previous section, the drivers and outcomes of
trophic cascades are important for understanding spatial-temporal patterns of kelp forests.[20] [21] [26]
In addition to ecological monitoring of kelp forests before, during, and after such disturbances, scientists try to
tease
apart the intricacies of kelp forest dynamics using experimental manipulations. By working on smaller
spatial-temporal scales, they can control for the presence or absence of specific biotic and abiotic factors to
discover
the operative mechanisms. For example, in southern Australia, manipulations of kelp canopy types demonstrated
that
the relative amount of Ecklonia radiata in a canopy could be used to predict understory species assemblages;
Kelp forest 110
consequently, the proportion of E. radiata can be used as an indicator of other species occurring in the
environment.[53]
Human use
Kelp forests have been important to human existence for thousands of years.[54] Indeed, many now theorise that
the
first colonisation of the Americas was due to fishing communities following the Pacific Kelp Forests during the
last
Ice Age. One theory contends that the kelp forests that would have stretched from northeast Asia to the
American
Pacific coast would have provided many benefits to ancient boaters. The kelp forests would have provided many
sustenance opportunities as well as acting as a type of buffer from rough water. Besides these benefits
researchers
believe that the kelp forests might have helped early boaters navigate, acting as a type of "kelp highway".
Theorists
also suggest that the kelp forests would have helped these ancient colonists by providing a stable way of life and
preventing them from having to adapt to new ecosystems and develop new survival methods even as they
traveled
thousands of miles.[55] Modern economies are based on fisheries of kelp-associated species like lobster and
rockfish.
Humans also harvest kelp directly to feed aquaculture species like abalone and to extract the compound alginic
acid,
which is used in products like toothpaste and antacids.[56] [57] Kelp forests are valued for recreational activities
such
as SCUBA diving and kayaking; the industries that support these sports represent one benefit related to the
ecosystem and the enjoyment derived from these activities represents another. All of these are examples of
ecosystem services provided specifically by kelp forests.
Threats and management
Marine protected areas are one way to guard kelp
forests as an ecosystem. The nudibranch Melibe
leonina on a Macrocystis frond (California).
Given the complexity of kelp forests – their variable structure,
geography and interactions – they pose a considerable challenge to
environmental managers. It is difficult to extrapolate even well-studied
trends to the future because interactions within the ecosystem will
change under variable conditions, not all relationships in the ecosystem
are understood, and there can be non-linear thresholds to transitions
that are not yet recognized.[58] With respect to kelp forests, major
issues of concern include marine pollution and water quality, kelp
harvesting and fisheries, invasive species and climate change.[4] It has
been argued that the most pressing threat to kelp forest preservation is
the overfishing of coastal ecosystems, which by removing higher
trophic levels facilitates their shift to depauperate urchin barrens.[5]
The maintenance of biodiversity is recognized as a way of generally
stabilizing ecosystems and their services through mechanisms such as
functional compensation and reduced susceptibility to foreign species
invasions.[59] [60] [61] [62]
In many places, managers have opted to regulate the harvest of kelp[18] [63] and/or the taking of kelp forest
species by
fisheries.[4] [52] While these may be effective in one sense, they do not necessarily protect the entirety of the
ecosystem. Marine protected areas (MPAs) offer a unique solution that encompasses not only target species for
harvesting but also the interactions surrounding them and the local environment as a whole.[64] [65] Direct
benefits of
MPAs to fisheries (for example, spillover effects) have been well documented around the world.[5] [66] [67] [68]
Indirect benefits have also been shown for several cases among species such as abalone and fishes in Central
California.[69] [70] Most importantly, studies have demonstrated that MPAs can be effective at protecting existing
kelp forest ecosystems and may also allow for the regeneration of those that have been impacted.[32] [71] [72]
Kelp forest 111
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External links
• Kelp Forest & Rocky Subtidal Habitats (http:/ / bonita. mbnms. nos. noaa. gov/ sitechar/ kelp. html)
• Monterey Bay Aquarium Kelp Cam (http:/ / www. mbayaq. org/ efc/ efc_hp/ hp_kelp_cam. asp) - watch a live
feed from the Monterey Bay Aquarium's kelp forest exhibit
• Kelp Watch (http:/ / www. kelpwatch. tas. gov. au) - a project by the Department of Primary Industries, Water
&
Environment of the government of Tasmania, Australia, with excellent general information on kelp forests as
well
as specific information on Tasmanian kelp forests.
Intertidal zone
The intertidal zone (also known as the foreshore and seashore and sometimes referred to as the littoral
zone) is the
area that is exposed to the air at low tide and underwater at high tide (for example, the area between tide marks).
This area can include many different types of habitats, with many types of animals like starfish, sea urchins, and
some species of coral. The well known area also includes steep rocky cliffs, sandy beaches, or wetlands (e.g.,
vast
mudflats). The area can be a narrow strip, as in Pacific islands that have only a narrow tidal range, or can include
many meters of shoreline where shallow beach slopes interact with high tidal excursion.
Organisms in the intertidal zone are adapted to an environment of harsh extremes. Water is available regularly
with
the tides but varies from fresh with rain to highly saline and dry salt with drying between tidal inundations. The
action of waves can dislodge residents in the littoral zone. With the intertidal zone's high exposure to the sun the
temperature range can be anything from very hot with full sun to near freezing in colder climates. Some
microclimates in the littoral zone are ameliorated by local features and larger plants such as mangroves.
Adaptation
in the littoral zone allows the use of nutrients supplied in high volume on a regular basis from the sea which is
actively moved to the zone by tides. Edges of habitats, in this case land and sea, are themselves often significant
ecologies, and the littoral zone is a prime example.
A typical rocky shore can be divided into a spray zone or splash zone (also known as the supratidal zone), which
is
above the spring high-tide line and is covered by water only during storms, and an intertidal zone, which lies
between the high and low tidal extremes. Along most shores, the intertidal zone can be clearly separated into the
following subzones: high tide zone, middle tide zone, and low tide zone. The intertidal zone is one of a number
of
marine biomes or habitats, including estuaries, neritic, surface and deep zones.
Intertidal zone 114
Zonation
Tide pools at Pillar Point showing zonation on the
edge of the rock ledge
A rock, seen at low tide, exhibiting typical
intertidal zonation, Kalaloch, Washington,
western USA.
Marine biologists and others divide the intertidal region into three
zones (low, middle, and high), based on the overall average exposure
of the zone. The low intertidal zone, which borders on the shallow
subtidal zone, is only exposed to air at the lowest of low tides and is
primarily marine in character. The mid intertidal zone is regularly
exposed and submerged by average tides. The high intertidal zone is
only covered by the highest of the high tides, and spends much of its
time as terrestrial habitat. The high intertidal zone borders on the
splash zone (the region above the highest still-tide level, but which
receives wave splash). On shores exposed to heavy wave action, the
intertidal zone will be influenced by waves, as the spray from breaking
waves will extend the intertidal zone.
Depending on the substratum and topography of the shore, additional
features may be noticed. On rocky shores, tide pools form in
depressions that fill with water as the tide rises. Under certain
conditions, such as those at Morecambe Bay, quicksand may form.
Low tide zone (lower littoral)
This subregion is mostly submerged - it is only exposed at the point of
low tide and for a longer period of time during extremely low tides.
This area is teeming with life; the most notable difference with this
subregion to the other three is that there is much more marine
vegetation, especially seaweeds. There is also a great biodiversity.
Organisms in this zone generally are not well adapted to periods of
dryness and temperature extremes. Some of the organisms in this area
are abalone, anemones, brown seaweed, chitons, crabs, green algae,
hydroids, isopods, limpets, mussels, nudibranchs, sculpin, sea
cucumber, sea lettuce, sea palms, sea stars, sea urchins, shrimp, snails,
sponges, surf grass, tube worms, and whelks. Creatures in this area can
grow to larger sizes because there is more available energy in the
localized ecosystem. Also, marine vegetation can grow to much greater
sizes than in the other three intertidal subregions due to the better water
coverage. The water is shallow enough to allow plenty of light to reach
the vegetation to allow substantial photosynthetic activity, and the
salinity is at almost normal levels. This area is also protected from
large predators such as fish because of the wave action and the
relatively shallow water.
Intertidal zone 115
Ecology
A California tide pool in the low tide zone
The intertidal region is an important model system for the study of
ecology, especially on wave-swept rocky shores. The region contains a
high diversity of species, and the zonation created by the tides causes
species ranges to be compressed into very narrow bands. This makes it
relatively simple to study species across their entire cross-shore range,
something that can be extremely difficult in, for instance, terrestrial
habitats that can stretch thousands of kilometers. Communities on
wave-swept shores also have high turnover due to disturbance, so it is
possible to watch ecological succession over years rather than decades.
Since the foreshore is alternately covered by the sea and exposed to the
air, organisms living in this environment must have adaptions for both wet and dry conditions. Hazards include
being
smashed or carried away by rough waves, exposure to dangerously high temperatures, and desiccation. Typical
inhabitants of the intertidal rocky shore include urchins, sea anemones, barnacles, chitons, crabs, isopods,
mussels,
sea stars, and many marine gastropod mollusks such as limpets, whelks, and even octopuses.
Legal issues
As with the dry sand part of a beach, legal and political disputes can arise over the ownership and use of the
foreshore. One recent example is the New Zealand foreshore and seabed controversy. In legal discussions the
foreshore is often referred to as the wet-sand area.
For privately-owned beaches in the United States, some states such as Massachusetts use the low water mark as
the
dividing line between the property of the State and that of the beach owner. Other states such as California use
the
high water mark.
In the UK, the foreshore is generally deemed to be owned by the Crown although there are notable exceptions,
especially what are termed several fisheries which can be historic deeds to title, dating back to King John's
time or
earlier, and the Udal Law, which applies generally in Orkney and Shetland.
Gallery
Mussels in the intertidal zone in
Cornwall, England.
Barnacles and limpets in the
intertidal zone near Newquay,
Cornwall.
A tidal pool in the intertidal zone
during low tide, Sunrise-on-Sea,
South Africa.
Unexplained crumbs of sand
which appear to have been
deposited around stone by
escaping air.
Intertidal zone 116
References
External links
• Enchanted Learning (http:/ / www. enchantedlearning. com/ biomes/ intertidal/ intertidal. shtml)
• Enyclopćdia Britannica (http:/ / www. britannica. com/ eb/ article-9048555)
• Watch the online documentary The Intertidal Zone (http:/ / www. nfb. ca/ film/ intertidal_zone/ )
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Image:Earth Western Hemisphere.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Earth_Western_Hemisphere.jpg License: Public Domain Contributors:
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Image:Pichavaram mangrove forest panorama.jpg Source: http://en.wikipedia.org/w/index.php?title=File:Pichavaram_mangrove_forest_panorama.jpg License:
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Secosky modified nasa
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modified nasa image.
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Bioms

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What are the different types of biomes?

How are biomes classified?

Bioms

Tropical and subtropical moist

Tropical and subtropical dry

Tropical and subtropical coniferous

Temperate broadleaf and mixed

Temperate coniferous forest

Tropical and subtropical grasslands,

Temperate grasslands, savannas,

Flooded grasslands and savannas

Montane grasslands and shrublands

Deserts and xeric shrublands

Upland and lowland (freshwater

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