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An ecotone is a transition area between two biological communities, where two communities meet and integrate. It may be narrow or wide, and it may be local (the zone between a field and forest) or regional (the transition between forest and grassland
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syst ...
s). An ecotone may appear on the ground as a gradual blending of the two communities across a broad area, or it may manifest itself as a sharp boundary line.


Etymology

The word ''ecotone'' was coined by
Alfred Russel Wallace Alfred Russel Wallace (8 January 1823 – 7 November 1913) was a British natural history, naturalist, explorer, geographer, anthropologist, biologist and illustrator. He is best known for independently conceiving the theory of evolution thro ...
, who first observed the abrupt boundary between two biomes in 1859. It is formed as a combination of ''ecology'' plus ''-tone'', from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
''tonos'' or tension – in other words, a place where ecologies are in tension.


Features

There are several distinguishing features of an ecotone. First, an ecotone can have a sharp vegetation transition, with a distinct line between two communities. For example, a change in colors of grasses or plant life can indicate an ecotone. Second, a change in
physiognomy Physiognomy (from the Greek , , meaning "nature", and , meaning "judge" or "interpreter") is the practice of assessing a person's character or personality from their outer appearance—especially the face. The term can also refer to the genera ...
(physical appearance of a plant species) can be a key indicator. Water bodies, such as estuaries, can also have a region of transition, and the boundary is characterized by the differences in heights of the macrophytes or plant species present in the areas because this distinguishes the two areas' accessibility to light. Scientists look at color variations and changes in plant height. Third, a change of species can signal an ecotone. There will be specific organisms on one side of an ecotone or the other. Other factors can illustrate or obscure an ecotone, for example,
migration Migration, migratory, or migrate may refer to: Human migration * Human migration, physical movement by humans from one region to another ** International migration, when peoples cross state boundaries and stay in the host state for some minimum le ...
and the establishment of new plants. These are known as spatial mass effects, which are noticeable because some organisms will not be able to form self-sustaining populations if they cross the ecotone. If different species can survive in both communities of the two biomes, then the ecotone is considered to have
species richness Species richness is the number of different species represented in an ecological community, landscape or region. Species richness is simply a count of species, and it does not take into account the abundances of the species or their relative ab ...
; ecologists measure this when studying the
food chain A food chain is a linear network of links in a food web starting from producer organisms (such as grass or algae which produce their own food via photosynthesis) and ending at an apex predator species (like grizzly bears or killer whales), de ...
and success of organisms. Lastly, the abundance of
introduced species An introduced species, alien species, exotic species, adventive species, immigrant species, foreign species, non-indigenous species, or non-native species is a species living outside its native distributional range, but which has arrived there ...
in an ecotone can reveal the type of biome or efficiency of the two communities sharing space. Because an ecotone is the zone in which two communities integrate, many different forms of life have to live together and compete for space. Therefore, an ecotone can create a diverse ecosystem.


Formation

Changes in the physical environment may produce a sharp
boundary Boundary or Boundaries may refer to: * Border, in political geography Entertainment * ''Boundaries'' (2016 film), a 2016 Canadian film * ''Boundaries'' (2018 film), a 2018 American-Canadian road trip film *Boundary (cricket), the edge of the pla ...
, as in the example of the interface between areas of
forest A forest is an area of land dominated by trees. Hundreds of definitions of forest are used throughout the world, incorporating factors such as tree density, tree height, land use, legal standing, and ecological function. The United Nations' ...
and cleared land. Elsewhere, a more gradually blended interface area will be found, where species from each community will be found together as well as unique local species.
Mountain range A mountain range or hill range is a series of mountains or hills arranged in a line and connected by high ground. A mountain system or mountain belt is a group of mountain ranges with similarity in form, structure, and alignment that have ari ...
s often create such ecotones, due to the wide variety of climatic conditions experienced on their
slope In mathematics, the slope or gradient of a line is a number that describes both the ''direction'' and the ''steepness'' of the line. Slope is often denoted by the letter ''m''; there is no clear answer to the question why the letter ''m'' is use ...
s. They may also provide a boundary between species due to the obstructive nature of their
terrain Terrain or relief (also topographical relief) involves the vertical and horizontal dimensions of land surface. The term bathymetry is used to describe underwater relief, while hypsometry studies terrain relative to sea level. The Latin word ...
.
Mont Ventoux Mont Ventoux (; oc, Ventor, label= Provençal ) is a mountain in the Provence region of southern France, located some northeast of Carpentras, Vaucluse. On the north side, the mountain borders the department of Drôme. At , it is the highest ...
in
France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...
is a good example, marking the boundary between the
flora Flora is all the plant life present in a particular region or time, generally the naturally occurring ( indigenous) native plants. Sometimes bacteria and fungi are also referred to as flora, as in the terms '' gut flora'' or '' skin flora''. ...
and
fauna Fauna is all of the animal life present in a particular region or time. The corresponding term for plants is '' flora'', and for fungi, it is '' funga''. Flora, fauna, funga and other forms of life are collectively referred to as '' biota''. ...
of northern and
southern France Southern France, also known as the South of France or colloquially in French as , is a defined geographical area consisting of the regions of France that border the Atlantic Ocean south of the Marais Poitevin,Louis Papy, ''Le midi atlantique'', A ...
. Most
wetland A wetland is a distinct ecosystem that is flooded or saturated by water, either permanently (for years or decades) or seasonally (for weeks or months). Flooding results in oxygen-free (Anoxic waters, anoxic) processes prevailing, especially in t ...
s are ecotones. The spatial variation of ecotones often form due to disturbances, creating patches that separate patches of vegetation. Different intensity of disturbances can cause landslides, land shifts, or movement of sediment that can create these vegetation patches and ecotones. Plants in competition extend themselves on one side of the ecotone as far as their ability to maintain themselves allows. Beyond this competitors of the adjacent community take over. As a result, the ecotone represents a shift in dominance. Ecotones are particularly significant for mobile animals, as they can exploit more than one set of
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
s within a short distance. The ecotone contains not only species common to the communities on both sides; it may also include a number of highly adaptable species that tend to colonize such transitional areas. The phenomenon of increased variety of plants as well as animals at the community junction is called the
edge effect In ecology, edge effects are changes in population or community structures that occur at the boundary of two or more habitats. Areas with small habitat fragments exhibit especially pronounced edge effects that may extend throughout the range. ...
and is essentially due to a locally broader range of suitable environmental conditions or
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
s.


Ecotones and ecoclines

An ecotone is often associated with an ecocline: a "physical transition zone" between two systems. The ecotone and ecocline concepts are sometimes confused: an ecocline can signal an ecotone chemically (ex: pH or
salinity Salinity () is the saltiness or amount of salt (chemistry), salt dissolved in a body of water, called saline water (see also soil salinity). It is usually measured in g/L or g/kg (grams of salt per liter/kilogram of water; the latter is dimensio ...
gradient In vector calculus, the gradient of a scalar-valued differentiable function of several variables is the vector field (or vector-valued function) \nabla f whose value at a point p is the "direction and rate of fastest increase". If the gr ...
), or microclimatically (
hydrothermal Hydrothermal circulation in its most general sense is the circulation of hot water (Ancient Greek ὕδωρ, ''water'',Liddell, H.G. & Scott, R. (1940). ''A Greek-English Lexicon. revised and augmented throughout by Sir Henry Stuart Jones. with th ...
gradient) between two ecosystems. In contrast: * an ecocline is a variation of the
physicochemical Physical chemistry is the study of macroscopic and microscopic phenomena in chemical systems in terms of the principles, practices, and concepts of physics such as motion, energy, force, time, thermodynamics, quantum chemistry, statistical mecha ...
environment dependent of one or two physico-chemical factors of life, and thus presence/absence of certain species. An ecocline can be a
thermocline A thermocline (also known as the thermal layer or the metalimnion in lakes) is a thin but distinct layer in a large body of fluid (e.g. water, as in an ocean or lake; or air, e.g. an atmosphere) in which temperature changes more drastically with ...
,
chemocline A chemocline is a type of cline, a layer of fluid with different properties, characterized by a strong, vertical chemistry gradient within a body of water. In bodies of water where chemoclines occur, the cline separates the upper and lower layer ...
(chemical gradient),
halocline In oceanography, a halocline (from Greek ''hals'', ''halos'' 'salt' and ''klinein'' 'to slope') is a cline, a subtype of chemocline caused by a strong, vertical salinity gradient within a body of water. Because salinity (in concert with te ...
(salinity gradient) or
pycnocline A pycnocline is the cline or layer where the density gradient () is greatest within a body of water. An ocean current is generated by the forces such as breaking waves, temperature and salinity differences, wind, Coriolis effect, and tides cause ...
(variations in density of water induced by temperature or salinity). * ecocline transitions are less distinct (less clear-cut), have more stable conditions within, hence a higher plant species richness. * an ecotone describes a variation in species prevalence and is often not strictly dependent on a major physical factor separating one ecosystem from another, with resulting habitat variability. An ecotone is often unobtrusive and harder to measure. * an ecotone is the area where two communities interact. Ecotones can be easily identified by distinct change in soil gradient and soil composition between two communities. * ecotone transitions are more clear-cut (distinct), conditions are less stable, hence they have a low species richness.


Examples

* The Kra ecotone between 11°N and 13°N latitude just north of the
Kra Isthmus The Kra Isthmus ( th, คอคอดกระ, ) in Thailand is the narrowest part of the Malay Peninsula. The western part of the isthmus belongs to Ranong Province and the eastern part to Chumphon Province, both in Southern Thailand. The ...
that connects the
Thai-Malay Peninsula The Malay Peninsula ( Malay: ''Semenanjung Tanah Melayu'') is a peninsula in Mainland Southeast Asia. The landmass runs approximately north–south, and at its terminus, it is the southernmost point of the Asian continental mainland. The ar ...
with mainland
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...
is an example of a regional scale ecotone. It marks the transition zone between the moist
deciduous In the fields of horticulture and Botany, the term ''deciduous'' () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed leaves, usually in the autumn; to the shedding of petals, ...
forest in the mainland
Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical south-eastern region of Asia, consisting of the regions that are situated south of mainland ...
biogeographical Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, ...
region in the north and the wet seasonal dipterocarp forest in the
Sundaland Sundaland (also called Sundaica or the Sundaic region) is a biogeographical region of South-eastern Asia corresponding to a larger landmass that was exposed throughout the last 2.6 million years during periods when sea levels were lower. It ...
region in the south. It has been shown to be the biogeographical transition between Indochinese and Sundaic faunas. Approximately 152 species of bird were found to have northern or southern range limits between these latitudes.
Population genetics Population genetics is a subfield of genetics that deals with genetic differences within and between populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as adaptation, speciation, and po ...
studies have also found that the Kra ecotone is the major physical barrier that limits gene flow in the honeybees ''
Apis cerana ''Apis cerana'', the eastern honey bee, Asiatic honey bee or Asian honey bee, is a species of honey bee native to South, Southeast and East Asia. This species is the sister species of ''Apis koschevnikovi'' and both are in the same subgenus as ...
'' and ''
Apis dorsata ''Apis dorsata'', the giant honey bee, सिङ्गुस in Nepali, is a honey bee of South and Southeast Asia, found mainly in forested areas such as the Terai of Nepal. They are typically around long. Nests are mainly built in exposed pla ...
'' and the stingless bees ''
Trigona collina ''Trigona collina'' is one of the stingless bee Stingless bees, sometimes called stingless honey bees or simply meliponines, are a large group of bees (about 550 described species), comprising the tribe Meliponini (or subtribe Meliponina accor ...
'' and '' Trigona pagdeni''. * The
Wallace Line The Wallace Line or Wallace's Line is a faunal boundary line drawn in 1859 by the British naturalist Alfred Russel Wallace and named by English biologist Thomas Henry Huxley that separates the biogeographical realms of Asia and Wallacea, a trans ...
running through the Lombok Strait between the
Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Gui ...
n islands of
Bali Bali () is a province of Indonesia and the westernmost of the Lesser Sunda Islands. East of Java and west of Lombok, the province includes the island of Bali and a few smaller neighbouring islands, notably Nusa Penida, Nusa Lembongan, and ...
and
Lombok Lombok is an island in West Nusa Tenggara province, Indonesia. It forms part of the chain of the Lesser Sunda Islands, with the Lombok Strait separating it from Bali to the west and the Alas Strait between it and Sumbawa to the east. It is ...
is a faunal boundary that separates the
Indomalayan realm The Indomalayan realm is one of the eight biogeographic realms. It extends across most of South and Southeast Asia and into the southern parts of East Asia. Also called the Oriental realm by biogeographers, Indomalaya spreads all over the Ind ...
from
Wallacea Wallacea is a biogeographical designation for a group of mainly Indonesian islands separated by deep-water straits from the Asian and Australian continental shelves. Wallacea includes Sulawesi, the largest island in the group, as well as ...
. It is named for
Alfred Russel Wallace Alfred Russel Wallace (8 January 1823 – 7 November 1913) was a British natural history, naturalist, explorer, geographer, anthropologist, biologist and illustrator. He is best known for independently conceiving the theory of evolution thro ...
, who first observed the abrupt boundary between the two
biomes A biome () is a biogeographical unit consisting of a biological community that has formed in response to the physical environment in which they are found and a shared regional climate. Biomes may span more than one continent. Biome is a broader ...
in 1859. Biologists believe it was the depth of the Lombok Strait itself that kept the animals on either side isolated from one another. However, it has been found that some flightless animals such as certain weevil species have, in the past, been involved in several transgression events in which species from land east of the Wallace Line relocated to Bali. When sea levels dropped during the
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
ice age An ice age is a long period of reduction in the temperature of Earth's surface and atmosphere, resulting in the presence or expansion of continental and polar ice sheets and alpine glaciers. Earth's climate alternates between ice ages and gre ...
, the islands of Bali,
Java Java (; id, Jawa, ; jv, ꦗꦮ; su, ) is one of the Greater Sunda Islands in Indonesia. It is bordered by the Indian Ocean to the south and the Java Sea to the north. With a population of 151.6 million people, Java is the world's mo ...
and
Sumatra Sumatra is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the sixth-largest island in the world at 473,481 km2 (182,812 mi.2), not including adjacent i ...
were all connected to one another and to the mainland of
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...
. They shared the Asian fauna. The Lombok Strait's deep water kept Lombok and the Lesser Sunda archipelago isolated from the Asian mainland. These islands were, instead, colonized by Australasian fauna. *
Mbam Djerem National Park The Mbam Djerem National Park is found in Cameroon. It was established in 2000 and covers . Overview A protected area was created in January 2000 in this ecosystem in the centre of Cameroon, and named the Mbam Djerem National Park. It covers 420 ...
's ecotone in
Cameroon Cameroon (; french: Cameroun, ff, Kamerun), officially the Republic of Cameroon (french: République du Cameroun, links=no), is a country in west-central Africa. It is bordered by Nigeria to the west and north; Chad to the northeast; the ...
is up to 1,000 km wide in places and differences within species are believed to be precursors to
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution withi ...
. * General examples of ecotones include
salt marsh A salt marsh or saltmarsh, also known as a coastal salt marsh or a tidal marsh, is a coastal ecosystem in the upper coastal intertidal zone between land and open saltwater or brackish water that is regularly flooded by the tides. It is domin ...
es and
riparian zone A riparian zone or riparian area is the interface between land and a river or stream. Riparian is also the proper nomenclature for one of the terrestrial biomes of the Earth. Plant habitats and communities along the river margins and banks a ...
s.


See also

*
Cline (biology) In biology, a cline (from the Greek κλίνειν ''klinein'', meaning "to lean") is a measurable gradient in a single character (or biological trait) of a species across its geographical range. First coined by Julian Huxley in 1938, the "cha ...
- ''Ecocline'' *
Ecotype In evolutionary ecology, an ecotype,Greek: ''οίκος'' = home and ''τύπος'' = type, coined by Göte Turesson in 1922 sometimes called ecospecies, describes a genetically distinct geographic variety, population, or race within a species ...
*
Phylogenetic niche conservatism The term phylogenetic niche conservatism has seen increasing use in recent years in the scientific literature, though the exact definition has been a matter of some contention. Fundamentally, phylogenetic niche conservatism refers to the tendency ...
* Plant functional type * Satoyama *
Biogeographic realm A biogeographic realm or ecozone is the broadest biogeographic division of Earth's land surface, based on distributional patterns of terrestrial organisms. They are subdivided into bioregions, which are further subdivided into ecoregions. ...


References


External links

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