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Species reintroduction is the deliberate release of a
species In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
into the wild, from captivity or other areas where the organism is capable of survival. The goal of species reintroduction is to establish a healthy, genetically diverse, self-sustaining population to an area where it has been extirpated, or to augment an existing
population Population typically refers to the number of people in a single area, whether it be a city or town, region, country, continent, or the world. Governments typically quantify the size of the resident population within their jurisdiction using ...
. Species that may be eligible for reintroduction are typically threatened or endangered in the wild. However, reintroduction of a species can also be for
pest control Pest control is the regulation or management of a species defined as a pest; any animal, plant or fungus that impacts adversely on human activities or environment. The human response depends on the importance of the damage done and will range ...
; for example, wolves being reintroduced to a wild area to curb an
overpopulation Overpopulation or overabundance is a phenomenon in which a species' population becomes larger than the carrying capacity of its environment. This may be caused by increased birth rates, lowered mortality rates, reduced predation or large scale m ...
of deer. Because reintroduction may involve returning native species to localities where they had been extirpated, some prefer the term "reestablishment". Humans have been reintroducing species for food and pest control for thousands of years. However, the practice of reintroducing for conservation is much younger, starting in the 20th century.


Methods for sourcing individuals

There are a variety of approaches to species reintroduction. The optimal strategy will depend on the biology of the organism. The first matter to address when beginning a species reintroduction is whether to source individuals ''in situ'', from wild populations, or ''ex situ'', from captivity in a zoo or botanic garden, for example.


''In situ'' sourcing

''
In situ ''In situ'' (; often not italicized in English) is a Latin phrase that translates literally to "on site" or "in position." It can mean "locally", "on site", "on the premises", or "in place" to describe where an event takes place and is used in ...
'' sourcing for restorations involves moving individuals from an existing wild population to a new site where the species was formerly extirpated. Ideally, populations should be sourced ''in situ'' when possible due to the numerous risks associated with reintroducing organisms from captive populations to the wild. To ensure that reintroduced populations have the best chance of surviving and reproducing, individuals should be sourced from populations that genetically and ecologically resemble the recipient population. Generally, sourcing from populations with similar environmental conditions to the reintroduction site will maximize the chance that reintroduced individuals are well adapted to the habitat of the reintroduction site . One consideration for ''in situ'' sourcing is at which life stage the organisms should be collected, transported, and reintroduced. For instance, with plants, it is often ideal to transport them as seeds as they have the best chance of surviving translocation at this stage. However, some plants are difficult to establish as seed and may need to be translocated as juveniles or adults.


''Ex situ'' sourcing

In situations where ''in situ'' collection of individuals is not feasible, such as for rare and endangered species with too few individuals existing in the wild, ''ex situ'' collection is possible. ''Ex situ'' collection methods allow storage of individuals that have high potential for reintroduction. Storage examples include
germplasm Germplasm are living genetic resources such as seeds or tissues that are maintained for the purpose of animal and plant breeding, preservation, and other research uses. These resources may take the form of seed collections stored in seed banks, t ...
stored in seed banks, sperm and egg banks,
cryopreservation Cryo-preservation or cryo-conservation is a process where organisms, organelles, cells, tissues, extracellular matrix, organs, or any other biological constructs susceptible to damage caused by unregulated chemical kinetics are preserved by coo ...
, and tissue culture. Methods that allow for storage of a high numbers of individuals also aim to maximize genetic diversity. Stored materials generally have long lifespans in storage, but some species do lose viability when stored as seed. Tissue culture and cryopreservation techniques have only been perfected for a few species. Organisms may also be kept in living collections in captivity. Living collections are more costly than storing
germplasm Germplasm are living genetic resources such as seeds or tissues that are maintained for the purpose of animal and plant breeding, preservation, and other research uses. These resources may take the form of seed collections stored in seed banks, t ...
and hence can support only a fraction of the individuals that ''ex situ'' sourcing can. Risk increases when sourcing individuals to add to living collections. Loss of genetic diversity is a concern because fewer individuals stored. Individuals may also become genetically adapted to captivity, which often adversely affects the reproductive fitness of individuals. Adaptation to captivity may make individuals less suitable for reintroduction to the wild. Thus, efforts should be made to replicate wild conditions and minimize time spent in captivity whenever possible.


Successes and failures

Reintroduction biology is a relatively young discipline and continues to be a work in progress. No strict and accepted definition of reintroduction success exists, but it has been proposed that the criteria widely used to assess the conservation status of endangered taxa, such as the
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biol ...
criteria, should be used to assess reintroduction success. Successful reintroduction programs should yield viable and self-sustainable populations in the long-term. The IUCN/SSC Re-introduction Specialist Group & Environment Agency, in their 2011 Global Re-introduction Perspectives, compiled reintroduction case studies from around the world.Soorae, P. S. (ed.) (2011). Global Re-introduction Perspectives: 2011.More case studies from around the globe. Gland, Switzerland: IUCN/SSC Re-introduction Specialist Group and Abu Dhabi, UAE: Environment Agency-Abu Dhabi. xiv + 250 pp.SBN: 978-2-8317-1432-5 https://portals.iucn.org/library/sites/library/files/documents/2011-073.pdf 184 case studies were reported on a range of species which included
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s,
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% ...
, amphibians, reptiles,
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s, mammals, and
plant Plants are predominantly Photosynthesis, photosynthetic eukaryotes of the Kingdom (biology), kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all curr ...
s. Assessments from all of the studies included goals, success indicators, project summary, major difficulties faced, major lessons learned, and success of project with reasons for success or failure. A similar assessment focused solely on plants found high rates of success for rare species reintroductions. An analysis of data from the Center for Plant Conservation International Reintroduction Registry found that, for the 49 cases where data were available, 92% of the reintroduced plant populations survived two years. The
Siberian tiger The Siberian tiger or Amur tiger is a population of the tiger subspecies ''Panthera tigris tigris'' native to the Russian Far East, Northeast China and possibly North Korea. It once ranged throughout the Korean Peninsula, but currently inhabi ...
population has rebounded from 40 individuals in the 1940s to around 500 in 2007. The Siberian tiger population is now the largest un-fragmented tiger population in the world. Yet, a high proportion of translocations and reintroductions have not been successful in establishing viable populations. For instance, in China reintroduction of captive Giant Pandas have had mixed effects. The initial pandas released from captivity all died quickly after reintroduction. Even now that they have improved their ability to reintroduce pandas, concern remains over how well the captive-bred pandas will fare with their wild relatives. Many factors can attribute to the success or failure of a reintroduction. Predators, food, pathogens, competitors, and weather can all affect a reintroduced population's ability to grow, survive, and reproduce. The number of animals reintroduced in an attempt should also vary with factors such as social behavior, expected rates of predation, and density in the wild. Animals raised in captivity may experience stress during captivity or translocation, which can weaken their immune systems. The IUCN reintroduction guidelines emphasize the need for an assessment of the availability of suitable habitat as a key component of reintroduction planning.Prepared by the SSC Re-introduction Specialist Group (May 1995) IUCN/SSC Guidelines for Re-Introductions http://www.iucnsscrsg.org Poor assessment of the release site can increase the chances that the species will reject the site and perhaps move to a less suitable environment. This can decrease the species fitness and thus decrease chances for survival. They state that restoration of the original habitat and amelioration of causes of extinction must be explored and considered as essential conditions for these projects. Unfortunately, the monitoring period that should follow reintroductions often remains neglected.


Genetic considerations

When a species has been extirpated from a site where it previously existed, individuals that will comprise the reintroduced population must be sourced from wild or captive populations. When sourcing individuals for reintroduction, it is important to consider
local adaptation Local adaptation is a mechanism in evolutionary biology whereby a population of organisms evolves to be more well-suited to its local environment than other members of the same species that live elsewhere. Local adaptation requires that different po ...
, adaptation to captivity (for ''ex situ'' conservation), the possibility of
inbreeding depression Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. ...
and
outbreeding depression In biology, outbreeding depression happens when crosses between two genetically distant groups or populations result in a reduction of fitness. This is particularly likely if the subspecies have different habitats or if no genetic exchange has occ ...
, and
taxonomy Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
,
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 ...
, and
genetic diversity Genetic diversity is the total number of Genetics, genetic characteristics in the genetic makeup of a species, it ranges widely from the number of species to differences within species and can be attributed to the span of survival for a species. ...
of the source population. Reintroduced populations experience increased vulnerability to influences of drift,
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 strat ...
, and
gene flow In population genetics, gene flow (also known as gene migration or geneflow and allele flow) is the transfer of genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalen ...
evolutionary processes due to their small sizes, climatic and ecological differences between source and native habitats, and presence of other mating-compatible populations. If the species slated for reintroduction is rare in the wild, it is likely to have unusually low population numbers, and care should be taken to avoid
inbreeding Inbreeding is the production of offspring from the mating or breeding of individuals or organisms that are closely related genetically. By analogy, the term is used in human reproduction, but more commonly refers to the genetic disorders an ...
and
inbreeding depression Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. ...
. Inbreeding can change the frequency of allele distribution in a population, and potentially result in a change to crucial genetic diversity. Additionally,
outbreeding depression In biology, outbreeding depression happens when crosses between two genetically distant groups or populations result in a reduction of fitness. This is particularly likely if the subspecies have different habitats or if no genetic exchange has occ ...
can occur if a reintroduced population can hybridize with existing populations in the wild, which can result in offspring with reduced fitness, and less adaptation to local conditions. To minimize both, practitioners should source for individuals in a way that captures as much genetic diversity as possible, and attempt to match source site conditions to local site conditions as much as possible. Capturing as much
genetic diversity Genetic diversity is the total number of Genetics, genetic characteristics in the genetic makeup of a species, it ranges widely from the number of species to differences within species and can be attributed to the span of survival for a species. ...
as possible, measured as
heterozygosity Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism. Mo ...
, is suggested in species reintroductions. Some protocols suggest sourcing approximately 30 individuals from a population will capture 95% of the genetic diversity. Maintaining genetic diversity in the recipient population is crucial to avoiding the loss of essential local adaptations, minimizing inbreeding depression, and maximizing fitness of the reintroduced population.


Ecological similarity

Plants or animals that undergo reintroduction may exhibit reduced fitness if they are not sufficiently adapted to local environmental conditions. Therefore, researchers should consider ecological and environmental similarity of source and recipient sites when selecting populations for reintroduction. Environmental factors to consider include climate and soil traits (pH, percent clay, silt and sand, percent combustion carbon, percent combustion nitrogen, concentration of Ca, Na, Mg, P, K). Historically, sourcing plant material for reintroductions has followed the rule "local is best," as the best way to preserve local adaptations, with individuals for reintroductions selected from the most geographically proximate population. However, geographic distance was shown in a common garden experiment to be an insufficient predictor of fitness. Additionally, projected climatic shifts induced by
climate change In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to ...
have led to the development of new seed sourcing protocols that aim to source seeds that are best adapted to project climate conditions. Conservation agencies have developed seed transfer zones that serve as guidelines for how far plant material can be transported before it will perform poorly. Seed transfer zones take into account proximity, ecological conditions, and climatic conditions in order to predict how plant performance will vary from one zone to the next. A study of the reintroduction of ''
Castilleja levisecta ''Castilleja levisecta'' is a rare species of flowering plant in the family Orobanchaceae known by the common name golden paintbrush, or golden Indian paintbrush, listed under the Endangered Species Act in 1997. It is native to British Columbia a ...
'' found that the source populations most physically near the reintroduction site performed the poorest in a field experiment, while those from the source population whose ecological conditions most closely matched the reintroduction site performed best, demonstrating the importance of matching the evolved adaptations of a population to the conditions at the reintroduction site.


Adaptation to captivity

Some reintroduction programs use plants or animals from captive populations to form a reintroduced population. When reintroducing individuals from a captive population to the wild, there is a risk that they have adapted to captivity due to differential selection of genotypes in captivity versus the wild. The genetic basis of this adaptation is selection of rare,
recessive In genetics, dominance is the phenomenon of one variant (allele) of a gene on a chromosome masking or overriding the effect of a different variant of the same gene on the other copy of the chromosome. The first variant is termed dominant and ...
allele An allele (, ; ; modern formation from Greek ἄλλος ''állos'', "other") is a variation of the same sequence of nucleotides at the same place on a long DNA molecule, as described in leading textbooks on genetics and evolution. ::"The chro ...
s that are deleterious in the wild but preferred in captivity. Consequently, animals adapted to captivity show reduced stress tolerance, increased tameness, and loss of local adaptations. Plants also can show adaptations to captivity through changes in drought tolerance, nutrient requirements, and seed dormancy requirements. Extent of adaptation is directly related to intensity of selection, genetic diversity,
effective population size The effective population size (''N'e'') is a number that, in some simplified scenarios, corresponds to the number of breeding individuals in the population. More generally, ''N'e'' is the number of individuals that an idealised population w ...
and number of generations in captivity. Characteristics selected for in captivity are overwhelmingly disadvantageous in the wild, so such adaptations can lead to reduced fitness following reintroduction. Reintroduction projects that introduce wild animals generally experience higher success rates than those that use captive-bred animals. Genetic adaptation to captivity can be minimized through management methods: by maximizing generation length and number of new individuals added to the captive population; minimizing effective population size, number of generations spent in captivity, and
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 strat ...
pressure; and reducing genetic diversity by fragmenting the population. For plants, minimizing adaptation to captivity is usually achieved by sourcing plant material from a
seed bank A seed bank (also seed banks or seeds bank) stores seeds to preserve genetic diversity; hence it is a type of gene bank. There are many reasons to store seeds. One is to preserve the genes that plant breeders need to increase yield, disease res ...
, where individuals are preserved as wild-collected seeds, and have not had the chance to adapt to conditions in captivity. However, this method is only plausible for plants with
seed dormancy Seed dormancy is an evolutionary adaptation that prevents seeds from germinating during unsuitable ecological conditions that would typically lead to a low probability of seedling survival. Dormant seeds do not germinate in a specified period of ...
.


Genetic trade-offs

In reintroductions from captivity, translocation of animals from captivity to the wild has implications for both captive and wild populations. Reintroduction of genetically valuable animals from captivity improves genetic diversity of reintroduced populations while depleting captive populations; conversely, genetically valuable captive-bred animals may be closely related to individuals in the wild and thus increase risk of inbreeding depression if reintroduced. Increasing genetic diversity is favored with removal of genetically overrepresented individuals from captive populations and addition of animals with low genetic relatedness to the wild. However, in practice, initial reintroduction of individuals with low genetic value to the captive population is recommended to allow for genetic assessment before translocation of valuable individuals.


Improving research techniques

A cooperative approach to reintroduction by ecologists and biologists could improve research techniques. For both preparation and monitoring of reintroductions, increasing contacts between academic population biologists and wildlife managers is encouraged within the Survival Species Commission and the IUCN. The IUCN states that a re-introduction requires a multidisciplinary approach involving a team of persons drawn from a variety of backgrounds. A survey by Wolf et al. in 1998 indicated that 64% of reintroduction projects have used subjective opinion to assess habitat quality. This means that most reintroduction evaluation has been based on human anecdotal evidence and not enough has been based on statistical findings. Seddon et al. (2007) suggest that researchers contemplating future reintroductions should specify goals, overall ecological purpose, and inherent technical and biological limitations of a given reintroduction, and planning and evaluation processes should incorporate both experimental and modeling approaches. Monitoring the health of individuals, as well as the survival, is important; both before and after the reintroduction. Intervention may be necessary if the situation proves unfavorable. Population dynamics models that integrate demographic parameters and behavioral data recorded in the field can lead to simulations and tests of a priori hypotheses. Using previous results to design further decisions and experiments is a central concept of
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 ove ...
. In other words, learning by doing can help in future projects. Population ecologists should therefore collaborate with biologists, ecologists, and wildlife management to improve reintroduction programs.BGCI: The global network. Retrieved April 30, 2012 from http://www.bgci.org/ourwork/Ecosystems/


Genetic monitoring

For reintroduced populations to successfully establish and maximize reproductive fitness, practitioners should perform genetic tests to select which individuals will be the founders of reintroduced populations and to continue monitoring populations post-reintroduction. A number of methods are available to measure the genetic relatedness between and variation among individuals within populations. Common genetic diversity assessment tools include
microsatellite A microsatellite is a tract of repetitive DNA in which certain DNA motifs (ranging in length from one to six or more base pairs) are repeated, typically 5–50 times. Microsatellites occur at thousands of locations within an organism's genome. ...
markers, mitochondrial DNA analyses,
alloenzyme Alloenzymes (or also called allozymes) are variant forms of an enzyme which differ structurally but not functionally from other allozymes coded for by different alleles at the same locus. These are opposed to isozymes, which are enzymes that perfo ...
s, and
amplified fragment length polymorphism AFLP-PCR or just AFLP is a PCR-based tool used in genetics research, DNA fingerprinting, and in the practice of genetic engineering. Developed in the early 1990s by KeyGene, AFLP uses restriction enzymes to digest genomic DNA, followed by lig ...
markers. Post-reintroduction, genetic monitoring tools can be used to obtain data such as population abundance,
effective population size The effective population size (''N'e'') is a number that, in some simplified scenarios, corresponds to the number of breeding individuals in the population. More generally, ''N'e'' is the number of individuals that an idealised population w ...
, and population structure, and can also be used to identify instances of
inbreeding Inbreeding is the production of offspring from the mating or breeding of individuals or organisms that are closely related genetically. By analogy, the term is used in human reproduction, but more commonly refers to the genetic disorders an ...
within reintroduced populations or hybridization with existing populations that are genetically compatible. Long-term genetic monitoring is recommended post-reintroduction to track changes in genetic diversity of the reintroduced population and determine success of a reintroduction program. Adverse genetic changes such as loss of
heterozygosity Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism. Mo ...
may indicate management intervention, such as population supplementation, is necessary for survival of the reintroduced population.


Re-introduction Specialist Group (RSG)

The RSG is a network of specialists whose aim is to combat the ongoing and massive
loss of biodiversity Biodiversity loss includes the worldwide extinction of different species, as well as the local reduction or loss of species in a certain habitat, resulting in a loss of biological diversity. The latter phenomenon can be temporary or permanent, de ...
by using re-introductions as a responsible tool for the management and restoration of biodiversity. It does this by actively developing and promoting sound inter-disciplinary scientific information, policy, and practice to establish viable wild populations in their natural habitats. The role of the RSG is to promote the re-establishment of viable populations in the wild of animals and plants. The need for this role was felt due to the increased demand from re-introduction practitioners, the global conservation community and increase in re-introduction projects worldwide. Increasing numbers of animal and plant species are becoming rare, or even extinct in the wild. In an attempt to re-establish populations, species can – in some instances – be re-introduced into an area, either through translocation from existing wild populations, or by re-introducing captive-bred animals or artificially propagated plants.


Reintroduction programs


Africa

*
Addax The addax (''Addax nasomaculatus''), also known as the white antelope and the screwhorn antelope, is an antelope native to the Sahara Desert. The only member of the genus ''Addax'', it was first described scientifically by Henri de Blainvill ...
in
Morocco Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria ...
and
Tunisia ) , image_map = Tunisia location (orthographic projection).svg , map_caption = Location of Tunisia in northern Africa , image_map2 = , capital = Tunis , largest_city = capital , ...
,
Chad Chad (; ar, تشاد , ; french: Tchad, ), officially the Republic of Chad, '; ) is a landlocked country at the crossroads of North and Central Africa. It is bordered by Libya to the north, Sudan to the east, the Central African Repub ...
* African bush elephant into
Samara Private Game Reserve Samara Private Game Reserve is a 28,300 hectare private game reserve, located near Graaff-Reinet in the Eastern Cape, South Africa. Previously a network of 11 livestock farms, Samara was created in 1997 with the vision to restore the area to its ...
in Eastern Cape,
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring count ...
*
African wild dog The African wild dog (''Lycaon pictus''), also called the painted dog or Cape hunting dog, is a wild canine which is a native species to sub-Saharan Africa. It is the largest wild canine in Africa, and the only extant member of the genus '' Ly ...
to
Gorongosa National Park Gorongosa National Park is at the southern end of the Great African Rift Valley in the heart of central Mozambique, Southeast Africa. The more than park comprises the valley floor and parts of surrounding plateaus. Rivers originating on nearby ...
,
Mozambique Mozambique (), officially the Republic of Mozambique ( pt, Moçambique or , ; ny, Mozambiki; sw, Msumbiji; ts, Muzambhiki), is a country located in southeastern Africa bordered by the Indian Ocean to the east, Tanzania to the north, Mala ...
and Lekedi Park,
Gabon Gabon (; ; snq, Ngabu), officially the Gabonese Republic (french: République gabonaise), is a country on the west coast of Central Africa. Located on the equator, it is bordered by Equatorial Guinea to the northwest, Cameroon to the north ...
(successful) *
Black rhinoceros The black rhinoceros, black rhino or hook-lipped rhinoceros (''Diceros bicornis'') is a species of rhinoceros, native to eastern and southern Africa including Angola, Botswana, Kenya, Malawi, Mozambique, Namibia, South Africa, Eswatini, Tanzania ...
in
Malawi Malawi (; or aláwi Tumbuka: ''Malaŵi''), officially the Republic of Malawi, is a landlocked country in Southeastern Africa that was formerly known as Nyasaland. It is bordered by Zambia to the west, Tanzania to the north and northe ...
,
Zambia Zambia (), officially the Republic of Zambia, is a landlocked country at the