Extinction debt

In ecology, extinction debt is the future extinction of species due to events in the past. The phrases dead clade walking and survival without recovery express the same idea.[1]

Extinction debt occurs because of time delays between impacts on a species, such as destruction of habitat, and the species' ultimate disappearance. For instance, long-lived trees may survive for many years even after reproduction of new trees has become impossible, and thus they may be committed to extinction. Technically, extinction debt generally refers to the number of species in an area likely to become extinct, rather than the prospects of any one species, but colloquially it refers to any occurrence of delayed extinction.

Extinction debt may be local or global, but most examples are local as these are easier to observe and model. It is most likely to be found in long-lived species and species with very specific habitat requirements (specialists).[2] Extinction debt has important implications for conservation, as it implies that species may become extinct due to past habitat destruction, even if continued impacts cease, and that current reserves may not be sufficient to maintain the species that occupy them. Interventions such as habitat restoration may reverse extinction debt.

Immigration credit is the corollary to extinction debt. It refers to the number of species likely to migrate to an area after an event such as the restoration of an ecosystem.[3]

Terminology

The term extinction debt was first used in 1994 in a paper by David Tilman, Robert May, Clarence Lehman and Martin Nowak,[4] although Jared Diamond used the term "relaxation time" to describe a similar phenomenon in 1972.[5]

Extinction debt is also known by the terms dead clade walking and survival without recovery[1] when referring to the species affected. The phrase "dead clade walking" was coined by David Jablonski as early as 2001[1] as a reference to Dead Man Walking,[6] a film whose title is based on American prison slang for a condemned prisoner's last walk to the execution chamber. "Dead clade walking" has since appeared in other scientists' writings about the aftermaths of mass extinctions.[7][8]

In discussions of threats to biodiversity, extinction debt is analogous to the "climate commitment" in climate change, which states that inertia will cause the earth to continue to warm for centuries even if no more greenhouse gasses are emitted. Similarly, the current extinction may continue long after human impacts on species halt.

Causes

Jablonski recognized at least four patterns in the fossil record following mass extinctions:[1]

(1) survival without recovery
also called “dead clade walking” – a group dwindling to extinction or relegation to precarious, minor ecological niches
(2) continuity with setbacks
patterns disturbed by the extinction event but soon continuing on the previous trajectory
(3) unbroken continuity
large-scale patterns continuing with little disruption
(4) unbridled diversification
an increase in diversity and species richness, as in the mammals following the end-Cretaceous extinction event

Extinction debt is caused by many of the same drivers as extinction. The most well-known drivers of extinction debt are habitat fragmentation and habitat destruction.[2] These cause extinction debt by reducing the ability of species to persist via immigration to new habitats. Under equilibrium conditions, species may become extinct in one habitat patch, yet continues to survive because it can disperse to other patches. However, as other patches have been destroyed or rendered inaccessible due to fragmentation, this "insurance" effect is reduced and the species may ultimately become extinct.

Pollution may also cause extinction debt by reducing a species' birth rate or increasing its death rate so that its population slowly declines.[9] Extinction debts may also be caused by invasive species[10] or by climate change.

Extinction debt may also occur due to the loss of mutualist species. In New Zealand, the local extinction of several species of pollinating birds in 1870 has caused a long-term reduction in the reproduction of the shrub species Rhabdothamnus solandri, which requires these birds to produce seeds. However, as the plant is slow-growing and long-lived, its populations persist.[11]

Jablonski found that the extinction rate of marine invertebrates was significantly higher in the stage (major subdivision of an epoch – typically 2–10 million years' duration) following a mass extinction than in the stages preceding the mass extinction. His analysis focused on marine molluscs since they constitute the most abundant group of fossils and are therefore the least likely to produce sampling errors. Jablonski suggested that two possible explanations deserved further study:

  • Post-extinction physical environments differed from pre-extinction environments in ways which were disadvantageous to the "dead clades walking".
  • Ecosystems that developed after recoveries from mass extinctions may have been less favorable for the "dead clades walking".[6]

Time scale

The time to "payoff" of extinction debt can be very long. Islands that lost habitat at the end of the last ice age 10,000 years ago still appear to be losing species as a result.[5] It has been shown that some bryozoans, a type of microscopic marine organism, became extinct due to the volcanic rise of the Isthmus of Panama. This event cut off the flow of nutrients from the Pacific Ocean to the Caribbean 3–4.5 million years ago. While bryozoan populations dropped severely at this time, extinction of these species took another 1–2 million years.[12]

Extinction debts incurred due to human actions have shorter timescales. Local extinction of birds from rainforest fragmentation occurs over years or decades,[13] while plants in fragmented grasslands show debts lasting 50–100 years.[14] Tree species in fragmented temperate forests have debts lasting 200 years or more.[15]

Theoretical development

Origins in metapopulation models

Tilman et al. demonstrated that extinction debt could occur using a mathematical ecosystem model of species metapopulations. Metapopulations are multiple populations of a species that live in separate habitat patches or islands but interact via immigration between the patches. In this model, species persist via a balance between random local extinctions in patches and colonization of new patches. Tilman et al. used this model to predict that species would persist long after they no longer had sufficient habitat to support them. When used to estimate extinction debts of tropical tree species, the model predicted debts lasting 50–400 years.[4]

One of the assumptions underlying the original extinction debt model was a trade-off between species' competitive ability and colonization ability. That is, a species that competes well against other species, and is more likely to become dominant in an area, is less likely to colonize new habitats due to evolutionary trade-offs. One of the implications of this assumption is that better competitors, which may even be more common than other species, are more likely to become extinct than rarer, less competitive, better dispersing species. This has been one of the more controversial components of the model, as there is little evidence for this trade-off in many ecosystems, and in many empirical studies dominant competitors were least likely species to become extinct.[16] A later modification of the model showed that these trade-off assumptions may be relaxed, but need to exist partially, in order for the theory to work.[17]

Development in other models

Further theoretical work has shown that extinction debt can occur under many different circumstances, driven by different mechanisms and under different model assumptions. The original model predicted extinction debt as a result of habitat destruction in a system of small, isolated habitats such as islands. Later models showed that extinction debt could occur in systems where habitat destruction occurs in small areas within a large area of habitat, as in slash-and-burn agriculture in forests, and could also occur due to decreased growth of species from pollutants.[9] Predicted patterns of extinction debt differ between models, though. For instance, habitat destruction resembling slash-and-burn agriculture is thought to affect rare species rather than poor colonizers. Models that incorporate stochasticity, or random fluctuation in populations, show extinction debt occurring over different time scales than classic models.[18]

Most recently, extinction debts have been estimated through the use models derived from neutral theory. Neutral theory has very different assumptions than the metapopulation models described above. It predicts that the abundance and distribution of species can be predicted entirely through random processes, without considering the traits of individual species. As extinction debt arises in models under such different assumptions, it is robust to different kinds of models. Models derived from neutral theory have successfully predicted extinction times for a number of bird species, but perform poorly at both very small and very large spatial scales.[19]

Mathematical models have also shown that extinction debt will last longer if it occurs in response to large habitat impacts (as the system will move farther from equilibrium), and if species are long-lived. Also, species just below their extinction threshold, that is, just below the population level or habitat occupancy levels required sustain their population, will have long-term extinction debts. Finally, extinction debts are predicted to last longer in landscapes with a few large patches of habitat, rather than many small ones.[20]

Detection

Extinction debt is difficult to detect and measure. Processes that drive extinction debt are inherently slow and highly variable (noisy), and it is difficult to locate or count the very small populations of near-extinct species. Because of these issues, most measures of extinction debt have a great deal of uncertainty.[2]

Experimental evidence

Due to the logistical and ethical difficulties of inciting extinction debt, there are few studies of extinction debt in controlled experiments. However, experiments microcosms of insects living on moss habitats demonstrated that extinction debt occurs after habitat destruction. In these experiments, it took 6–12 months for species to die out following the destruction of habitat.[13]

Observational methods

Long-term observation

Extinction debts that reach equilibrium in relatively short time scales (years to decades) can be observed via measuring the change in species numbers in the time following an impact on habitat. For instance, in the Amazon rainforest, researchers have measured the rate at which bird species disappear after forest is cut down.[21] As even short-term extinction debts can take years to decades to reach equilibrium, though, such studies take many years and good data are rare.

Comparing the past and present

Most studies of extinction debt compare species numbers with habitat patterns from the past and habitat patterns in the present. If the present populations of species are more closely related to past habitat patterns than present, extinction debt is a likely explanation. The magnitude of extinction debt (i.e., number of species likely to become extinct) can not be estimated by this method.[2]

If one has information on species populations from the past in addition to the present, the magnitude of extinction debt can be estimated. One can use the relationship between species and habitat from the past to predict the number of species expected in the present. The difference between this estimate and the actual number of species is the extinction debt.[2]

This method requires the assumption that in the past species and their habitat were in equilibrium, which is often unknown. Also, a common relationship used to equate habitat and species number is the species-area curve, but as the species-area curve arises from very different mechanisms than those in metapopulation based models, extinction debts measured in this way may not conform with metapopulation models' predictions.[9] The relationship between habitat and species number can also be represented by much more complex models that simulate the behavior of many species independently.[15]

Comparing impacted and pristine habitats

If data on past species numbers or habitat are not available, species debt can also be estimated by comparing two different habitats: one which is mostly intact, and another which has had areas cleared and is smaller and more fragmented. One can then measure the relationship of species with the condition of habitat in the intact habitat, and, assuming this represents equilibrium, use it to predict the number of species in the cleared habitat. If this prediction is lower than the actual number of species in the cleared habitat, then the difference represents extinction debt.[2] This method requires many of the same assumptions as methods comparing the past and present.

Examples

Grasslands

Studies of European grasslands show evidence of extinction debt through both comparisons with the past and between present-day systems with different levels of human impacts. The species diversity of grasslands in Sweden appears to be a remnant of more connected landscapes present 50 to 100 years ago.[14] In alvar grasslands in Estonia that have lost area since the 1930s, 17–70% of species are estimated to be committed to extinction.[22] However, studies of similar grasslands in Belgium, where similar impacts have occurred, show no evidence of extinction debt.[23] This may be due to differences in the scale of measurement or the level of specialization of grass species.[24]

Forests

Forests in Vlaams-Brabant, Belgium, show evidence of extinction debt remaining from deforestation that occurred between 1775 and 1900. Detailed modeling of species behavior, based on similar forests in England that did not experience deforestation, showed that long-lived and slow-growing species were more common than equilibrium models would predict, indicating that their presence was due to lingering extinction debt.[15]

In Sweden, some species of lichens show an extinction debt in fragments of ancient forest. However, species of lichens that are habitat generalists, rather than specialists, do not.[25]

Insects

Extinction debt has been found among species of butterflies living in the grasslands on Saaremaa and Muhu – islands off the western coast of Estonia. Butterfly species distributions on these islands are better explained by the habitat in the past than current habitats.[26]

On the islands of the Azores Archipelago, more than 95% of native forests have been destroyed in the past 600 years. As a result, more than half of arthropods on these islands are believed to be committed to extinction, with many islands likely to lose more than 90% of species.[27]

Vertebrates

80–90% of extinction from past deforestation in the Amazon has yet to occur, based on modeling based on species-area relationships. Local extinctions of approximately 6 species are expected in each 2500 km2 region by 2050 due to past deforestation.[28] Birds in the Amazon rain forest continued to become extinct locally for 12 years following logging that broke up contiguous forest into smaller fragments. The extinction rate slowed, however, as forest regrew in the spaces in between habitat fragments.[21]

Countries in Africa are estimated to have, on average, a local extinction debt of 30% for forest-dwelling primates. That is, they are expected to have 30% of their forest primate species to become extinct in the future due to loss of forest habitat. The time scale for these extinctions has not been estimated.[29]

Based on historical species-area relationships, Hungary currently has approximately nine more species of raptors than are thought to be able to be supported by current nature reserves.[30]

Applications to conservation

The existence of extinction debt in many different ecosystems has important implications for conservation. It implies that in the absence of further habitat destruction or other environmental impacts, many species are still likely to become extinct. Protection of existing habitats may not be sufficient to protect species from extinction.[30] However, the long time scales of extinction debt may allow for habitat restoration in order to prevent extinction,[2] as occurred in the slowing of extinction in Amazon forest birds above.[21] In another example, it has been found that grizzly bears in very small reserves in the Rocky Mountains are likely to become extinct, but this finding allows the modification of reserve networks to better support their populations.[31]

The extinction debt concept may require revision of the value of land for species conservation, as the number of species currently present in a habitat may not be a good measure of the habitat's ability to support species (see carrying capacity) in the future.[25] As extinction debt may last longest near extinction thresholds, it may be hardest to detect the threat of extinction for species that conservation could benefit the most.[20]

Economic analyses have shown that including extinction in management decision-making process changes decision outcomes, as the decision to destroy habitat changes conservation value in the future as well as the present. It is estimated that in Costa Rica, ongoing extinction debt may cost between $88 million and $467 million.[32]

In popular culture

References

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Abiotic component

In biology and ecology, abiotic components or abiotic factors are non-living chemical and physical parts of the environment that affect living organisms and the functioning of ecosystems. Abiotic factors and the phenomena associated with them underpin all biology.

Abiotic components include physical conditions and non-living resources that affect living organisms in terms of growth, maintenance, and reproduction. Resources are distinguished as substances or objects in the environment required by one organism and consumed or otherwise made unavailable for use by other organisms.

Component degradation of a substance occurs by chemical or physical processes, e.g. hydrolysis. All non-living components of an ecosystem, such as atmospheric conditions and water resources, are called abiotic components.

Bacterivore

Bacterivores are free-living, generally heterotrophic organisms, exclusively microscopic, which obtain energy and nutrients primarily or entirely from the consumption of bacteria. Many species of amoeba are bacterivores, as well as other types of protozoans. Commonly, all species of bacteria will be prey, but spores of some species, such as Clostridium perfringens, will never be prey, because of their cellular attributes.

Copiotroph

A copiotroph is an organism found in environments rich in nutrients, particularly carbon. They are the opposite to oligotrophs, which survive in much lower carbon concentrations.

Copiotrophic organisms tend to grow in high organic substrate conditions. For example, copiotrophic organisms grow in Sewage lagoons. They grow in organic substrate conditions up to 100x higher than oligotrophs.

Decomposer

Decomposers are organisms that break down dead or decaying organisms, and in doing so, they carry out the natural process of decomposition. Like herbivores and predators, decomposers are heterotrophic, meaning that they use organic substrates to get their energy, carbon and nutrients for growth and development. While the terms decomposer and detritivore are often interchangeably used, detritivores must ingest and digest dead matter via internal processes while decomposers can directly absorb nutrients through chemical and biological processes hence breaking down matter without ingesting it. Thus, invertebrates such as earthworms, woodlice, and sea cucumbers are technically detritivores, not decomposers, since they must ingest nutrients and are unable to absorb them externally.

Depensation

In population dynamics, depensation is the effect on a population (such as a fish stock) whereby, due to certain causes, a decrease in the breeding population (mature individuals) leads to reduced production and survival of eggs or offspring. The causes may include predation levels rising per offspring (given the same level of overall predator pressure) and the allee effect, particularly the reduced likelihood of finding a mate.

Dominance (ecology)

Ecological dominance is the degree to which a taxon is more numerous than its competitors in an ecological community, or makes up more of the biomass.

Most ecological communities are defined by their dominant species.

In many examples of wet woodland in western Europe, the dominant tree is alder (Alnus glutinosa).

In temperate bogs, the dominant vegetation is usually species of Sphagnum moss.

Tidal swamps in the tropics are usually dominated by species of mangrove (Rhizophoraceae)

Some sea floor communities are dominated by brittle stars.

Exposed rocky shorelines are dominated by sessile organisms such as barnacles and limpets.

Energy Systems Language

The Energy Systems Language, also referred to as Energese, Energy Circuit Language, or Generic Systems Symbols, was developed by the ecologist Howard T. Odum and colleagues in the 1950s during studies of the tropical forests funded by the United States Atomic Energy Commission. They are used to compose energy flow diagrams in the field of systems ecology.

Extinction threshold

Extinction threshold is a term used in conservation biology to explain the point at which a species, population or metapopulation, experiences an abrupt change in density or number because of an important parameter, such as habitat loss. It is at this critical value below which a species, population, or metapopulation, will go extinct, though this may take a long time for species just below the critical value, a phenomenon known as extinction debt.Extinction thresholds are important to conservation biologists when studying a species in a population or metapopulation context because the colonization rate must be larger than the extinction rate, otherwise the entire entity will go extinct once it reaches the threshold.Extinction thresholds are realized under a number of circumstances and the point in modeling them is to define the conditions that lead a population to extinction. Modeling extinction thresholds can explain the relationship between extinction threshold and habitat loss and habitat fragmentation.

Feeding frenzy

In ecology, a feeding frenzy occurs when predators are overwhelmed by the amount of prey available. For example, a large school of fish can cause nearby sharks, such as the lemon shark, to enter into a feeding frenzy. This can cause the sharks to go wild, biting anything that moves, including each other or anything else within biting range. Another functional explanation for feeding frenzy is competition amongst predators. This term is most often used when referring to sharks or piranhas. It has also been used as a term within journalism.

Lithoautotroph

A lithoautotroph or chemolithoautotroph is a microbe which derives energy from reduced compounds of mineral origin. Lithoautotrophs are a type of lithotrophs with autotrophic metabolic pathways. Lithoautotrophs are exclusively microbes; macrofauna do not possess the capability to use mineral sources of energy. Most lithoautotrophs belong to the domain Bacteria, while some belong to the domain Archaea. For lithoautotrophic bacteria, only inorganic molecules can be used as energy sources. The term "Lithotroph" is from Greek lithos (λίθος) meaning "rock" and trōphos (τροφοσ) meaning "consumer"; literally, it may be read "eaters of rock". Many lithoautotrophs are extremophiles, but this is not universally so.

Lithoautotrophs are extremely specific in using their energy source. Thus, despite the diversity in using inorganic molecules in order to obtain energy that lithoautotrophs exhibit as a group, one particular lithoautotroph would use only one type of inorganic molecule to get its energy.

Mesotrophic soil

Mesotrophic soils are soils with a moderate inherent fertility. An indicator of soil fertility is its base status, which is expressed as a ratio relating the major nutrient cations (calcium, magnesium, potassium and sodium) found there to the soil's clay percentage. This is commonly expressed in hundredths of a mole of cations per kilogram of clay, i.e. cmol (+) kg−1 clay.

Microecosystem

Microecosystems can exist in locations which are precisely defined by critical environmental factors within small or tiny spaces.

Such factors may include temperature, pH, chemical milieu, nutrient supply, presence of symbionts or solid substrates, gaseous atmosphere (aerobic or anaerobic) etc.

Mycotroph

A mycotroph is a plant that gets all or part of its carbon, water, or nutrient supply through symbiotic association with fungi. The term can refer to plants that engage in either of two distinct symbioses with fungi:

Many mycotrophs have a mutualistic association with fungi in any of several forms of mycorrhiza. The majority of plant species are mycotrophic in this sense. Examples include Burmanniaceae.

Some mycotrophs are parasitic upon fungi in an association known as myco-heterotrophy.

Organotroph

An organotroph is an organism that obtains hydrogen or electrons from organic substrates. This term is used in microbiology to classify and describe organisms based on how they obtain electrons for their respiration processes. Some organotrophs such as animals and many bacteria, are also heterotrophs. Organotrophs can be either anaerobic or aerobic.

Antonym: Lithotroph, Adjective: Organotrophic.

Overpopulation

Overpopulation occurs when a species' population exceeds the carrying capacity of its ecological niche. It can result from an increase in births (fertility rate), a decline in the mortality rate, an increase in immigration, or an unsustainable biome and depletion of resources. When overpopulation occurs, individuals limit available resources to survive.

The change in number of individuals per unit area in a given locality is an important variable that has a significant impact on the entire ecosystem.

Planktivore

A planktivore is an aquatic organism that feeds on planktonic food, including zooplankton and phytoplankton.

Recruitment (biology)

In biology, especially marine biology, recruitment occurs when a juvenile organism joins a population, whether by birth or immigration, usually at a stage whereby the organisms are settled and able to be detected by an observer.There are two types of recruitment: closed and open.In the study of fisheries, recruitment is "the number of fish surviving to enter the fishery or to some life history stage such as settlement or maturity".

Relative abundance distribution

In the field of ecology, the relative abundance distribution (RAD) or species abundance distribution describes the relationship between the number of species observed in a field study as a function of their observed abundance. The graphs obtained in this manner are typically fitted to a Zipf–Mandelbrot law, the exponent of which serves as an index of biodiversity in the ecosystem under study.

Rhabdothamnus

Rhabdothamnus solandri, is the only member of the genus Rhabdothamnus, and the only plant from the Gesneriaceae family native to New Zealand. The common names for the plant are New Zealand gloxinia, and in Māori language: taurepo, matata and waiuatua.

Rhabdothamnus solandri is a small shrub growing to 2 metres. It is only found in the North Island in a variety of locations such as forests, near streams, or on banks. R. solandri has a distinctive red and yellow trumpet shaped flower.Due to extinction debt, the future extinction of this shrub is nearly guaranteed. The local extinction of several species of pollinating birds in 1870 has caused a long-term reduction in the reproduction of the shrub species, which requires these birds to produce seeds. However, as the plant is slow-growing and long-lived, its populations persist.

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