Island ecology
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Island An island (or isle) is an isolated piece of habitat that is surrounded by a dramatically different habitat, such as water. Very small islands such as emergent land features on atolls can be called islets, skerries, cays or keys. An island ...
ecology Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overl ...
is the study of island organisms and their interactions with each other and the environment. Islands account for nearly 1/6 of earth’s total land area,Paulay, G. 1994. Biodiversity on Oceanic Islands: Its Origin and Extinction. ''American Zoology'' 34: 134-144. yet the ecology of island ecosystems is vastly different from that of mainland communities. Their isolation and high availability of empty
niche Niche may refer to: Science *Developmental niche, a concept for understanding the cultural context of child development *Ecological niche, a term describing the relational position of an organism's species *Niche differentiation, in ecology, the ...
s lead to increased speciation. As a result, island ecosystems comprise 30% of the world’s biodiversity hotspots, 50% of marine tropical diversity, and some of the most unusual and rare species.Gerlach, Justin. 2008
Island biodiversity - issues and opportunities
. World Conservation Congress
Many species still remain unknown. The diversity of species on islands is highly impacted by human activities such as
deforestation Deforestation or forest clearance is the removal of a forest or stand of trees from land that is then converted to non-forest use. Deforestation can involve conversion of forest land to farms, ranches, or urban use. The most concentrated ...
and introduction of the
exotic 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 response, ecologists and managers are directing attention towards
conservation Conservation is the preservation or efficient use of resources, or the conservation of various quantities under physical laws. Conservation may also refer to: Environment and natural resources * Nature conservation, the protection and managem ...
and restoration of island species. Because they are simple systems, islands provide an opportunity to study processes of
extinction Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
that can be extrapolated to larger ecosystems.


Ecological processes on islands

Islands are attractive sites for ecological research because they provide clear examples of evolution in action. They show interesting patterns of colonization, adaptation, and speciation.


Colonization and establishment

Islands are surrounded by water, and may or may not exist as part of a continental land mass. Oceanic islands arise due to volcanic activity or reef growth, and usually subside over time due to erosion and changing sea levels. When islands emerge, they undergo the process of ecological succession as species colonize the island (see theory of
island biogeography Insular biogeography or island biogeography is a field within biogeography that examines the factors that affect the species richness and diversification of isolated natural communities. The theory was originally developed to explain the pattern of ...
). New species cannot immigrate via land, and instead must arrive via air, water, or wind. As a result, organisms with high dispersal capabilities, such as plants and birds, are much more common on islands than are poorly dispersing taxa like mammals. However, some mammals are present on islands, presumably from swimming or riding on natural “rafts” that are washed away from the mainland. Of the species that arrive, only some will be able to survive and establish populations. As a result, islands have fewer species than mainland habitats. Island populations are small and exhibit low genetic variability (see founder effect), but are isolated from the predators and competitors that they initially evolved with. This can lead to a process called
ecological release Ecological release refers to a population increase or population explosion that occurs when a species is freed from limiting factors in its environment. Sometimes this may occur when a plant or animal species is introduced, for example, to an islan ...
, where a species is released from its ancestral community interactions and then colonizes new niches.


Adaptation

In response to these changing ecological pressures, island species can become much more docile than their mainland counterparts, and may grow larger (see
island gigantism Island gigantism, or insular gigantism, is a biological phenomenon in which the size of an animal species isolated on an island increases dramatically in comparison to its mainland relatives. Island gigantism is one aspect of the more general Fos ...
) or smaller (see
island dwarfism Insular dwarfism, a form of phyletic dwarfism, is the process and condition of large animals evolving or having a reduced body size when their population's range is limited to a small environment, primarily islands. This natural process is disti ...
). Some of these unique adaptations are reflected in charismatic island species such as the Malagasy hippopotamus,
Komodo dragon The Komodo dragon (''Varanus komodoensis''), also known as the Komodo monitor, is a member of the monitor lizard family Varanidae that is endemic to the Indonesian islands of Komodo, Rinca, Flores, and Gili Motang. It is the largest extant ...
, or pygmy mammoths. Although, the giant tortoises of the
Galápagos Islands The Galápagos Islands (Spanish: , , ) are an archipelago of volcanic islands. They are distributed on each side of the equator in the Pacific Ocean, surrounding the centre of the Western Hemisphere, and are part of the Republic of Ecuador ...
and the
Seychelles Seychelles (, ; ), officially the Republic of Seychelles (french: link=no, République des Seychelles; Creole: ''La Repiblik Sesel''), is an archipelagic state consisting of 115 islands in the Indian Ocean. Its capital and largest city, ...
(the
Galápagos tortoise The Galápagos tortoise or Galápagos giant tortoise (''Chelonoidis niger'') is a species of very large tortoise in the genus ''Chelonoidis'' (which also contains three smaller species from mainland South America). It comprises 15 subspecies ( ...
and
Aldabrachelys ''Aldabrachelys'' is the recognised genus for the Seychelles and Madagascan radiations of giant tortoises, including the Aldabra giant tortoise (''Aldabrachelys gigantea''). Naming This name is problematic in that the type specimen is actually r ...
respectively) are sometimes given as examples of insular gigantism, they are now thought to represent the last remaining populations of historically widespread giant tortoises i.e. gigantism is an ancestral trait that occurred in the absence of insular selection pressures. The collection of differences in
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines * Morphology (archaeology), study of the shapes or forms of artifacts * Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies ...
,
ecology Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overl ...
,
physiology Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemical ...
and behaviour of insular species compared to their continental counterparts is termed
Island syndrome Island syndrome describes the differences in morphology, ecology, physiology and behaviour of insular species compared to their continental counterparts. These differences evolve due to the different ecological pressures affecting insular speci ...
. After immigration, birds, and some reptiles or mammals, tend to become larger and predatory, showing relaxed
intraspecific competition Intraspecific competition is an interaction in population ecology, whereby members of the same species compete for limited resources. This leads to a reduction in fitness for both individuals, but the more fit individual survives and is able to r ...
. For mammals, small species will increase in size and large species will decrease in size. This is referred to as the “island rule,” and is suggested to minimize energy expenditure.McNab, B.K. 2002. Minimizing energy expenditure facilitates vertebrate persistence on oceanic islands. Ecology Letters 5(5): 693-704. Other adaptations to life on islands include increased
poikilothermy A poikilotherm () is an animal whose internal temperature varies considerably. Poikilotherms have to survive and adapt to environmental stress. One of the most important stressors is temperature change, which can lead to alterations in membrane ...
, relaxed anti-predator behavior, and reduced
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex ( ...
in animals, and loss of herbivore defenses and reduced dispersal in plants.


Speciation

The formation of new islands and their isolation from the mainland provides many unoccupied niches for species to adapt to. Since immigration of predators and competitors is limited, many organisms are able to persist in these new niches. This results in a high occurrence of
endemism Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found else ...
, where species are unique to a localized area. For example, 50% of endemic bird areas are found on islands. Endemism is often the result of
adaptive radiation In evolutionary biology, adaptive radiation is a process in which organisms diversify rapidly from an ancestral species into a multitude of new forms, particularly when a change in the environment makes new resources available, alters biotic int ...
. Adaptive radiation is when a single species colonizes an area and rapidly diversifies to fill all of the available niches. A common example is the assemblage of finch species documented by
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended ...
in the Galapagos Islands. Darwin’s finches exhibited adaptive radiation by evolving different beak sizes to exploit the diversity of seeds present on the different islands. Because the distributions of these populations are limited by their island habitats, they tend to have fewer individuals than their mainland counterparts and lower genetic variation. This, along with the behavioral and ecological factors mentioned above, makes island species more vulnerable to extinction.


Survival

The continued survival of species on islands depends on factors such as natural
selection Selection may refer to: Science * Selection (biology), also called natural selection, selection in evolution ** Sex selection, in genetics ** Mate selection, in mating ** Sexual selection in humans, in human sexuality ** Human mating strateg ...
, genetic variation, natural disturbances (hurricanes, volcanic eruptions) and human-caused disturbances (introduced species,
habitat loss Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
). Human-caused disturbances tend to be the greatest cause of mortality, and understanding the causes of extinction facilitates conservation efforts.


Human impacts on island ecosystems

The movement of humans to islands has led to rapid extinction of native island species either from hunting,
habitat destruction Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
, or
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 ther ...
.


Hunting

Many large animals on islands have been hunted to extinction by humans. A well-known example is the
dodo The dodo (''Raphus cucullatus'') is an extinct flightless bird that was endemic to the island of Mauritius, which is east of Madagascar in the Indian Ocean. The dodo's closest genetic relative was the also-extinct Rodrigues solitaire. The ...
, once found on the island of
Mauritius Mauritius ( ; french: Maurice, link=no ; mfe, label= Mauritian Creole, Moris ), officially the Republic of Mauritius, is an island nation in the Indian Ocean about off the southeast coast of the African continent, east of Madagascar. It ...
. It evolved to become large, flightless and docile, and was subsequently driven to extinction by humans and introduced predators.


Habitat destruction

The depletion of natural resources can have dramatic effects on island ecology. On
Easter Island Easter Island ( rap, Rapa Nui; es, Isla de Pascua) is an island and special territory of Chile in the southeastern Pacific Ocean, at the southeasternmost point of the Polynesian Triangle in Oceania. The island is most famous for its ne ...
, the depletion of the forest by humans not only resulted in widespread loss of species, but also the collapse of the island civilization. Today there are over 500 million people on islands, all dependent on local resources either directly (traditional use) or indirectly (ecotourism revenue). Population growth and development result in heavy
deforestation Deforestation or forest clearance is the removal of a forest or stand of trees from land that is then converted to non-forest use. Deforestation can involve conversion of forest land to farms, ranches, or urban use. The most concentrated ...
,
pollution Pollution is the introduction of contaminants into the natural environment that cause adverse change. Pollution can take the form of any substance (solid, liquid, or gas) or energy (such as radioactivity, heat, sound, or light). Pollutants, the ...
, and
over-exploitation Overexploitation, also called overharvesting, refers to harvesting a renewable resource to the point of diminishing returns. Continued overexploitation can lead to the destruction of the resource, as it will be unable to replenish. The term ap ...
. Overharvesting of ocean fauna is particularly troubling as yields of
coral reef fish Coral reef fish are fish which live amongst or in close relation to coral reefs. Coral reefs form complex ecosystems with tremendous biodiversity. Among the myriad inhabitants, the fish stand out as colourful and interesting to watch. Hundreds ...
species are an important food source for island populations.


Introduced species

Humans have contributed to globalization and decreased effective isolation of island communities, allowing for invasion of exotic species. This can have a profound effect on the native species. In
Guam Guam (; ch, Guåhan ) is an organized, unincorporated territory of the United States in the Micronesia subregion of the western Pacific Ocean. It is the westernmost point and territory of the United States (reckoned from the geographic cent ...
, the introduced
brown tree snake The brown tree snake (''Boiga irregularis''), also known as the brown catsnake, is an arboreal rear-fanged colubrid snake native to eastern and northern coastal Australia, eastern Indonesia (Sulawesi to Papua), Papua New Guinea, and many islands ...
ate nearly all of the native vertebrate species to extinction. Feral cats and dogs have also greatly diminished native vertebrate populations on islands, through both predation and disease. Introduced
ungulate Ungulates ( ) are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraffes, ...
s are another major threat, as they graze on native vegetation and can destroy entire forests. Exotic grasses can out-compete native
understory In forestry and ecology, understory (American English), or understorey (Commonwealth English), also known as underbrush or undergrowth, includes plant life growing beneath the forest canopy without penetrating it to any great extent, but abo ...
species and increase the risk of fire. Lastly, social insects such as ants also cause major problems. Nonnative species introduced onto islands can have profound effects on an island's ecosystem, more so than nonnative species introduced to continental land (Platenberg). The higher impact of introduced and nonnative species on islands is largely a result of lower biodiversity levels (Platenburg et al). Biodiversity on islands is especially threatened by logging, hunting, fishing and plant gathering. However, islands’ vulnerability to introduced species is also due to agricultural, economic, and health differences as compared to continental land (Russel et al). The smaller land area and population sizes of islands compared to continents create greater vulnerability to the impacts of introduced species (Russel et al). Introduced species negatively impact ecosystems through altered predator-prey interactions that can cause harm or even local extinction to native species populations (Towns et al). There are many examples of animal species such as birds, reptiles, and aquatic insects being harmed by the introduction of predators such as rats, cats, and ants. For example, seabirds on islands in Hawaii are impacted by non-native predators like barn owls and ants. At a colony in Hawaii, at least 20% of wedge-tailed shearwater eggs were taken by mynas, dark-plumaged birds of the starling family (Towns et al). Also in Hawaii, more than forty four introduced ant species, especially tropical fire ants, caused detriments to the growth of shearwater chicks, such as loss of tissue in their feet (Towns et al). Other examples of high-impact species to island ecosystems are the Cuban treefrog and cane toad, which were introduced to the US Virgin Islands under various circumstances (Platenberg). The Cuban treefrog has been present in the USVI since arriving unintentionally on a cargo ship in the 1970s (Platenberg). The Cuban tree frog is known for its ability to survive under harsh conditions, and it is highly adaptable, as it will eat a wide variety of organisms (Platenberg). Native frog and anole lizard populations have declined, likely due to Cuban tree frog interference (Platenberg). In contrast to the accidental introduction of the Cuban treefrog, the cane toad was deliberately introduced to control agricultural pests (Platenberg). Similar to the Cuban treefrog, the cane toad is also omnivorous. Cane toad tadpoles compete for limited freshwater resources with the white-lipped frog, a native species, thus limiting their resources (Platenberg). In general, when the introduction of a nonnative species results in extinction, the ecosystem experiences losses in some trophic levels (Platenberg). For example, vertebrate herbivores that are prone to extinction change the ecosystem function in plant communities. This phenomenon is seen in New Zealand, where the loss of bird species may have changed dynamics in avian-induced vegetation communities and impacted abundances of forest plants (Platenberg). Also in New Zealand, small mammalian predators such as rodents have little direct effect on vegetation but have greater effect on island faunas through extinction and displacement (Wood et al). Pacific rats, for example, were thought to be causing the local extinction of large, nocturnal ground-dwelling lizards, and they were assumed to have minimal effect on diurnal species, those that sleep at night, such as shore skinks (Wood et al). In places where Pacific rats were removed, however, shore skink populations rapidly changed, indicating that the rats’ impact has been underestimated (Wood et al). The loss of a keystone species, a species that all other species in the ecosystem depend on such as seabirds, can also have significant effects (Towns). Seabirds are essential to the structure of the ecosystem because they transport large amounts of nutrients into ecosystems and burrow into soils, helping vegetation (Towns). However, since human settlement, seabird communities have been severely impacted on islands across the globe (Towns). While some exotic species may perform ecological roles similar to those of extinct species, there are many roles that cannot be fulfilled by other species. Therefore, similar exotic species do not offer complete replacement for extinct species in the community (Towns). Even if the number of species introduced to an island is roughly the same as the number of extinct ones, the ecological traits are not comparable enough to make up for losses (Towns).


Global warming

Global warming is emerging as a strong cause of species loss on islands. This can be due to
sea level rise Globally, sea levels are rising due to human-caused climate change. Between 1901 and 2018, the globally averaged sea level rose by , or 1–2 mm per year on average.IPCC, 2019Summary for Policymakers InIPCC Special Report on the Ocean and Cry ...
, the intrusion of salt water into freshwater habitats, or species inability to adapt to increasing temperatures and extreme weather events. Plant species are particularly susceptible. In more isolated areas, such as the Southern Ocean Islands, indirect effects such as invasive species and global warming can play a greater role in influencing populations than overexploitation, pollution and
habitat loss Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
.


Trophic cascades

Human activities and the introduction of non-native species often cause
trophic cascade Trophic cascades are powerful indirect interactions that can control entire ecosystems, occurring when a trophic level in a food web is suppressed. For example, a top-down cascade will occur if predators are effective enough in predation to reduce t ...
s, where direct effects on one species result in indirect effects on other species in the food web. An example is on
Santa Cruz Island Santa Cruz Island ( Spanish: ''Isla Santa Cruz'', Chumash: ''Limuw'') is located off the southwestern coast of Ventura, California, United States. It is the largest island in California and largest of the eight islands in the Channel Islands ...
of the
California Channel Islands The Channel Islands () are an eight-island archipelago located within the Southern California Bight in the Pacific Ocean, off the coast of California. The four Northern Channel Islands are part of the Transverse Ranges geologic province, and ...
, where
DDT Dichlorodiphenyltrichloroethane, commonly known as DDT, is a colorless, tasteless, and almost odorless crystalline chemical compound, an organochloride. Originally developed as an insecticide, it became infamous for its environmental impacts. ...
poisoning reduced bald eagle populations. This, along with an abundance of introduced feral pigs for prey, allowed
golden eagle The golden eagle (''Aquila chrysaetos'') is a bird of prey living in the Northern Hemisphere. It is the most widely distributed species of eagle. Like all eagles, it belongs to the family Accipitridae. They are one of the best-known birds of ...
s to colonize the island and replace bald eagles. However, the golden eagles also ate native
island fox The island fox (''Urocyon littoralis'') is a small fox that is endemic to six of the eight Channel Islands of California. There are six subspecies, each unique to the island it lives on, reflecting its evolutionary history. They are generally d ...
es. Fox population levels decreased to near extinction, while skunk populations increased due to relaxed competition with foxes.


Island conservation


Conservation on islands

Since island ecosystems are self-contained, it should be possible to mitigate many of the threats to species. Ecologists and managers are working together to prioritize areas for conservation and to quickly design and implement action plans. Not everything can be put into a reserve, so it is important to first compile pertinent information and prioritize areas of concern.Kepler, C.B. and J.M. Scott. 1985. Conservation of Island Ecosystems. ICBP Technical Publication Number 3: 255-271. Once an area has been chosen, managers must then acquire ownership and gain support. Local experts and indigenous populations should also be involved in this process. Having clearly defined goals will facilitate the many necessary interactions between people and agencies. Once a reserve is in place, managers can then practice
adaptive management Adaptive management, also known as adaptive resource management or adaptive environmental assessment and management, is a structured, iterative process of robust decision making in the face of uncertainty, with an aim to reducing uncertainty over ...
and do continued community education. On land, island conservation focuses on the protection of species and their habitat. In some cases, conservation can be integrated with agricultural production. For example, the ''
Acacia koa ''Acacia koa'' or commonly known as koa is a species of flowering tree in the family Fabaceae. It is endemic to the Hawaiian Islands, where it is the second most common tree. The highest populations are on Hawaii, Maui and Oahu. Name The name ...
'' plantations and wooded pastures in
Hawaii Hawaii ( ; haw, Hawaii or ) is a state in the Western United States, located in the Pacific Ocean about from the U.S. mainland. It is the only U.S. state outside North America, the only state that is an archipelago, and the only state ...
are anthropogenically altered ecosystems, yet allow connectivity between forest fragments and thus maintain higher diversity than would open pasture. Other directions include habitat restoration, and eradication of introduced predators, ungulates, and exotic plants (via hunting, removal or
biological control Biological control or biocontrol is a method of controlling pests, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically also i ...
). In marine ecosystems, there has been an increasing establishment of “no-take” reserves. This allows for the reestablishment of native species, and also the augmentation of commercially harvested species. However, in both terrestrial and marine systems, these actions are expensive and do not always result in the desired outcomes. For example, some non-natives become keystone species and their removal can cause more harm than good to the ecosystem. To be more effective, managers of island ecosystems should share information and learn from each other’s mistakes.Cronk, Q.C.B. 1997. Islands: stability, diversity, conservation. Biodiversity and Conservation 6: 477-493.


Island restoration

Island conservation tends to focus on preservation of individual species and their habitats. However, many ecologists caution that ecological and evolutionary processes should be conserved as well. The conservation of island communities as a whole is closely linked to restoration. Active restoration on islands can be done for both animal species (translocations, induced breeding) and plant species (reforestation). Creating goals for restoration can be challenging because it is often impossible to restore the ecosystem to its “historic” or “normal” state, if that state can even be clearly defined. Restoration is never complete, as ecological communities are always in a state of change.


Sustainable use

As
resource depletion Resource depletion is the consumption of a resource faster than it can be replenished. Natural resources are commonly divided between renewable resources and non-renewable resources (see also mineral resource classification). Use of either ...
is a major issue on islands, the needs of human populations must also be taken into account. On many islands, scientists and managers are studying traditional practices of indigenous populations as potential conservation solutions. In some cases, limited-take systems that serve the community may provide a better alternative to fully closed protected areas, if there are not enough resources for proper enforcement. Public education plays an important role.


See also

*
Disturbance (ecology) In ecology, a disturbance is a temporary change in environmental conditions that causes a pronounced change in an ecosystem. Disturbances often act quickly and with great effect, to alter the physical structure or arrangement of biotic and abioti ...
*
Island biogeography Insular biogeography or island biogeography is a field within biogeography that examines the factors that affect the species richness and diversification of isolated natural communities. The theory was originally developed to explain the pattern of ...
*
Island syndrome Island syndrome describes the differences in morphology, ecology, physiology and behaviour of insular species compared to their continental counterparts. These differences evolve due to the different ecological pressures affecting insular speci ...
* Hawaiian honeycreeper conservation *
Mammals of the Caribbean A unique and diverse albeit phylogenetically restricted mammal fauna is known from the Caribbean region. The region—specifically, all islands in the Caribbean Sea (except for small islets close to the continental mainland) and the Bahamas, Turks ...
*
Patch dynamics Patch dynamics is an ecological perspective that the structure, function, and dynamics of ecological systems can be understood through studying their interactive patches. Patch dynamics, as a term, may also refer to the spatiotemporal changes wi ...
*
Distance decay Distance decay is a geographical term which describes the effect of distance on cultural or spatial interactions. The distance decay effect states that the interaction between two locales declines as the distance between them increases. Once the d ...
*
Sky island Sky islands are isolated mountains surrounded by radically different lowland environments. The term originally referred to those found on the Mexican Plateau, and has extended to similarly isolated high-elevation forests. The isolation has s ...


References


External links


Island Conservation
{{DEFAULTSORT:Island Ecology Ecology Islands