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In
biogeography Biogeography is the study of the species distribution, distribution of species and ecosystems in geography, geographic space and through evolutionary history of life, geological time. Organisms and biological community (ecology), communities o ...
and
paleontology Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure ge ...
, a relict is a
population Population is a set of humans or other organisms in a given region or area. Governments conduct a census to quantify the resident population size within a given jurisdiction. The term is also applied to non-human animals, microorganisms, and pl ...
or
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
of organisms that was more widespread or more diverse in the past. A relictual population is a population currently inhabiting a restricted area whose range was far wider during a previous geologic epoch. Similarly, a relictual taxon is a taxon (e.g. species or other lineage) which is the sole surviving representative of a formerly diverse group.


Definition

A relict (or relic) plant or animal is a
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
that persists as a remnant of what was once a diverse and widespread population. Relictualism occurs when a widespread habitat or range changes and a small area becomes cut off from the whole. A subset of the population is then confined to the available hospitable area, and survives there while the broader population either shrinks or evolves divergently. This phenomenon differs from
endemism Endemism is the state of a species being found only 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 foun ...
in that the range of the population was not always restricted to the local region. In other words, the species or group did not necessarily arise in that small area, but rather was stranded, or insularized, by changes over time. The agent of change could be anything from
competition Competition is a rivalry where two or more parties strive for a common goal which cannot be shared: where one's gain is the other's loss (an example of which is a zero-sum game). Competition can arise between entities such as organisms, indi ...
from other organisms,
continental drift Continental drift is a highly supported scientific theory, originating in the early 20th century, that Earth's continents move or drift relative to each other over geologic time. The theory of continental drift has since been validated and inc ...
, or
climate change Present-day climate change includes both global warming—the ongoing increase in Global surface temperature, global average temperature—and its wider effects on Earth's climate system. Climate variability and change, Climate change in ...
such as an
ice age An ice age is a long period of reduction in the temperature of Earth's surface and atmosphere, resulting in the presence or expansion of continental and polar ice sheets and alpine glaciers. Earth's climate alternates between ice ages, and g ...
. When a relict is representative of taxa found in the
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
record, and yet is still living, such an organism is sometimes referred to as a
living fossil A living fossil is a Deprecation, deprecated term for an extant taxon that phenotypically resembles related species known only from the fossil record. To be considered a living fossil, the fossil species must be old relative to the time of or ...
. However, a relict need not be currently living. An evolutionary relict is any organism that was characteristic of the flora or fauna of one age and that persisted into a later age, with the later age being characterized by newly evolved flora or fauna significantly different from those that came before.


Examples

A notable example is the
thylacine The thylacine (; binomial name ''Thylacinus cynocephalus''), also commonly known as the Tasmanian tiger or Tasmanian wolf, was a carnivorous marsupial that was native to the Mainland Australia, Australian mainland and the islands of Tasmani ...
of Tasmania, a relict
marsupial Marsupials are a diverse group of mammals belonging to the infraclass Marsupialia. They are natively found in Australasia, Wallacea, and the Americas. One of marsupials' unique features is their reproductive strategy: the young are born in a r ...
carnivore that survived into modern times on an island, whereas the rest of its species on mainland Australia had gone extinct between 3000 and 2000 years ago. Another example is '' Omma,'' a genus of beetle with a fossil record extending back over 200 million years to the Late Triassic and found worldwide during the Jurassic and Cretaceous, now confined to a single living species in Australia. Another relict from the Triassic is '' Pholadomya'', a common clam genus during the Mesozoic, now confined to a single rare species in the Caribbean. The
tuatara The tuatara (''Sphenodon punctatus'') is a species of reptile endemic to New Zealand. Despite its close resemblance to lizards, it is actually the only extant member of a distinct lineage, the previously highly diverse order Rhynchocephal ...
endemic to New Zealand is the only living member of the once-diverse reptile order Rhynchocephalia, which has a fossil record stretching back 240 million years and during the Mesozoic era was globally distributed and ecologically diverse. An example from the fossil record would be a specimen of
Nimravidae Nimravidae is an extinct family (biology), family of carnivorans, sometimes known as false saber-toothed cats, whose fossils are found in North America and Eurasia. Not considered to belong to the true cats (family Felidae), the nimravids are gen ...
, an extinct branch of
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they ar ...
s in the mammalian evolutionary tree, if said specimen came from Europe in the
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
epoch. If that was the case, the specimen would represent, not the main population, but a last surviving remnant of the nimravid lineage. These carnivores were common and widespread in the previous epoch, the
Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...
, and disappeared when the climate changed and woodlands were replaced by
savanna A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) biome and ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach th ...
. They persisted in Europe in the last remaining forests as a relict of the Oligocene: a relict species in a relict habitat. An example of divergent evolution creating relicts is found in the shrews of the islands off the coast of Alaska, namely the Pribilof Island shrew and the St. Lawrence Island shrew. These species are apparently relicts of a time when the islands were connected to the mainland, and these species were once conspecific with a more widespread species, now the cinereus shrew, the three populations having diverged through
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
. In
botany Botany, also called plant science, is the branch of natural science and biology studying plants, especially Plant anatomy, their anatomy, Plant taxonomy, taxonomy, and Plant ecology, ecology. A botanist or plant scientist is a scientist who s ...
, an example of an ice age relict plant population is the Snowdon lily, notable as being precariously rare in
Wales Wales ( ) is a Countries of the United Kingdom, country that is part of the United Kingdom. It is bordered by the Irish Sea to the north and west, England to the England–Wales border, east, the Bristol Channel to the south, and the Celtic ...
. The Welsh population is confined to the north-facing slopes of
Snowdonia Snowdonia, or Eryri (), is a mountainous region and National parks of the United Kingdom, national park in North Wales. It contains all 15 mountains in Wales Welsh 3000s, over 3000 feet high, including the country's highest, Snowdon (), which i ...
, where climatic conditions are apparently similar to ice age Europe. Some have expressed concern that the warming climate will cause the lily to die out in
Great Britain Great Britain is an island in the North Atlantic Ocean off the north-west coast of continental Europe, consisting of the countries England, Scotland, and Wales. With an area of , it is the largest of the British Isles, the List of European ...
. Other populations of the same plant can be found in the Arctic and in the mountains of Europe and North America, where it is known as the common alplily. While the extirpation of a geographically disjunct population of a relict species may be of regional conservation concern, outright extinction at the species level may occur in this century of rapid
climate change Present-day climate change includes both global warming—the ongoing increase in Global surface temperature, global average temperature—and its wider effects on Earth's climate system. Climate variability and change, Climate change in ...
if geographic range occupied by a relict species has already contracted to the degree that it is narrowly
endemic Endemism is the state of a species being found only 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 foun ...
. For this reason, the traditional conservation tool of translocation has recently been reframed as assisted migration of narrowly endemic, critically endangered species that are already (or soon expected) to experience climate change beyond their levels of tolerance. Two examples of critically endangered relict species for which assisted migration projects are already underway are the western swamp tortoise of Australia and a subcanopy conifer tree in the United States called Florida Torreya. A well-studied botanical example of a relictual taxon is ''
Ginkgo biloba ''Ginkgo biloba'', commonly known as ginkgo or gingko ( ), also known as the maidenhair tree, is a species of gymnosperm tree native to East Asia. It is the last living species in the order Ginkgoales, which first appeared over 290 million year ...
'', the last living representative of Ginkgoales that is restricted to China in the wild. ''Ginkgo'' trees had a diverse and widespread northern distribution during the
Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...
, but are not known from the fossil record after the
Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58 The Saimaa ringed seal (''Phoca hispida saimensis'') is an endemic subspecies, a relict of last ice age that lives only in
Finland Finland, officially the Republic of Finland, is a Nordic country in Northern Europe. It borders Sweden to the northwest, Norway to the north, and Russia to the east, with the Gulf of Bothnia to the west and the Gulf of Finland to the south, ...
in the landlocked and fragmented Saimaa freshwater lake complex. Now the population has less than 400 individuals, which poses a threat to its survival. Another example is the relict leopard frog once found throughout
Nevada Nevada ( ; ) is a landlocked state in the Western United States. It borders Oregon to the northwest, Idaho to the northeast, California to the west, Arizona to the southeast, and Utah to the east. Nevada is the seventh-most extensive, th ...
,
Arizona Arizona is a U.S. state, state in the Southwestern United States, Southwestern region of the United States, sharing the Four Corners region of the western United States with Colorado, New Mexico, and Utah. It also borders Nevada to the nort ...
,
Utah Utah is a landlocked state in the Mountain states, Mountain West subregion of the Western United States. It is one of the Four Corners states, sharing a border with Arizona, Colorado, and New Mexico. It also borders Wyoming to the northea ...
, and
Colorado Colorado is a U.S. state, state in the Western United States. It is one of the Mountain states, sharing the Four Corners region with Arizona, New Mexico, and Utah. It is also bordered by Wyoming to the north, Nebraska to the northeast, Kansas ...
, but now only found at Lake Mead National Recreation Area in Nevada and Arizona.


Relevance

The concept of relictualism is useful in understanding the ecology and conservation status of populations that have become insularized, meaning confined to one small area or multiple small areas with no chance of movement between populations. Insularization makes a population vulnerable to forces that can lead to
extinction Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
, such as disease,
inbreeding Inbreeding is the production of offspring from the mating or breeding of individuals or organisms that are closely genetic distance, related genetically. By analogy, the term is used in human reproduction, but more commonly refers to the genet ...
,
habitat destruction Habitat destruction (also termed habitat loss or habitat reduction) occurs when a natural habitat is no longer able to support its native species. The organisms once living there have either moved elsewhere, or are dead, leading to a decrease ...
, competition from
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 ...
, and
global warming Present-day climate change includes both global warming—the ongoing increase in global average temperature—and its wider effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes ...
. Consider the case of the white-eyed river martin, a very localized species of bird found only in Southeast Asia, and extremely rare, if not already extinct. Its closest and only surviving living relative is the African river martin, also very localized in central Africa. These two species are the only known members of the subfamily Pseudochelidoninae, and their widely disjunct populations suggest they are relict populations of a more common and widespread ancestor. Known to science only since 1968, it seems to have disappeared. Studies have been done on relict populations in isolated mountain and valley habitats in western North America, where the basin and range topography creates areas that are insular in nature, such as forested mountains surrounded by inhospitable desert, called sky islands. Such situations can serve as refuges for certain
Pleistocene The Pleistocene ( ; referred to colloquially as the ''ice age, Ice Age'') is the geological epoch (geology), epoch that lasted from to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fin ...
relicts, such as Townsend's pocket gopher, while at the same time creating barriers for
biological dispersal Biological dispersal refers to both the movement of individuals (animals, plants, fungi, bacteria, etc.) from their birth site to their breeding site ('natal dispersal') and the movement from one breeding site to another ('breeding dispersal' ...
. Studies have shown that such insular habitats have a tendency toward decreasing species richness. This observation has significant implications for conservation biology, because
habitat fragmentation Habitat fragmentation describes the emergence of discontinuities (fragmentation) in an organism's preferred environment (habitat), causing population fragmentation and ecosystem decay. Causes of habitat fragmentation include geological proces ...
can also lead to the insularization of stranded populations. So-called "relics of cultivation"Celka Z., Drapikowska M. 2008
Relics of cultivation in Central Europe: ''Malva alcea'' L. as an example.
Vegetation History and Archaeobotany. Volume 17, Supplement 1, 251-255,
are plant species that were grown in the past for various purposes (medicinal, food, dyes, etc.), but are no longer utilized. They are naturalized and can be found at archaeological sites.


See also

*
Living fossil A living fossil is a Deprecation, deprecated term for an extant taxon that phenotypically resembles related species known only from the fossil record. To be considered a living fossil, the fossil species must be old relative to the time of or ...


References

{{Reflist Biogeography