
In biology, a refugium (plural: ''refugia'') is a location which supports an isolated or
relict 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 ...
of a once more widespread species. This isolation (
allopatry) can be due to climatic changes, geography, or human activities such as
deforestation
Deforestation or forest clearance is the removal and destruction 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. Ab ...
and overhunting.
Present examples of refugial animal species are the
mountain gorilla, isolated to specific mountains in central Africa, and the
Australian sea lion
The Australian sea lion (''Neophoca cinerea''), also known as the Australian sea-lion or Australian sealion, is a species of sea lion that is the only endemic pinniped in Australia. It is currently Monotypic taxon, monotypic in the genus ''Neopho ...
, isolated to specific breeding beaches along the south-west coast of Australia, due to humans taking so many of their number as game. This resulting isolation, in many cases, can be seen as only a temporary state; however, some refugia may be longstanding, thereby having many
endemic species, not found elsewhere, which survive as relict populations. The
Indo-Pacific Warm Pool has been proposed to be a longstanding refugium, based on the discovery of the "living fossil" of a marine
dinoflagellate called ''
Dapsilidinium pastielsii'', currently found only in the Indo-Pacific Warm Pool.
For plants,
anthropogenic climate change propels scientific interest in identifying refugial species that were isolated into small or
disjunct ranges during glacial episodes of the
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 ...
, yet whose ability to expand their ranges during the warmth of
interglacial periods (such as the
Holocene
The Holocene () is the current geologic time scale, geological epoch, beginning approximately 11,700 years ago. It follows the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene to ...
) was apparently limited or precluded by
topographic,
streamflow, or
habitat
In ecology, habitat refers to the array of resources, biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species' habitat can be seen as the physical manifestation of its ...
barriers
—or by the extinction of
coevolved animal dispersers. The concern is that ongoing warming trends will expose them to
extirpation
Local extinction, also extirpation, is the termination of a species (or other taxon) in a chosen geographic area of study, though it still exists elsewhere. Local extinctions are contrasted with extinction, global extinctions.
Local extinctions ...
or
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 ...
in the decades ahead.
In
anthropology
Anthropology is the scientific study of humanity, concerned with human behavior, human biology, cultures, society, societies, and linguistics, in both the present and past, including archaic humans. Social anthropology studies patterns of behav ...
, ''refugia'' often refers specifically to
Last Glacial Maximum refugia, where some ancestral human populations may have been forced back to
glacial refugia (similar small isolated pockets on the face of the continental
ice sheet
In glaciology, an ice sheet, also known as a continental glacier, is a mass of glacier, glacial ice that covers surrounding terrain and is greater than . The only current ice sheets are the Antarctic ice sheet and the Greenland ice sheet. Ice s ...
s) during the
last glacial period. Going from west to east, suggested examples include the
Franco-Cantabrian region (in northern
Iberia
The Iberian Peninsula ( ), also known as Iberia, is a peninsula in south-western Europe. Mostly separated from the rest of the European landmass by the Pyrenees, it includes the territories of peninsular Spain and Continental Portugal, compri ...
), the
Italian and
Balkan peninsulas, the
Ukrainian LGM refuge, and the
Bering Land Bridge. Archaeological and genetic data suggest that the source populations of Paleolithic humans survived the glacial maxima (including the
Last Glacial Maximum
The Last Glacial Maximum (LGM), also referred to as the Last Glacial Coldest Period, was the most recent time during the Last Glacial Period where ice sheets were at their greatest extent between 26,000 and 20,000 years ago.
Ice sheets covered m ...
) in sparsely wooded areas and dispersed through areas of high primary productivity while avoiding dense
forest cover
Forest cover is the amount of trees that covers a particular area of land. It may be measured as relative (in percent) or absolute (in square kilometres/ square miles). Nearly a third of the world's land surface is covered with forest, with clos ...
.
Glacial refugia, where human populations found refuge during the last glacial period, may have played a crucial role in shaping the emergence and diversification of the language families that exist in the world today.
[
]
More recently, ''refugia'' has been used to refer to areas that could offer relative climate stability in the face of modern
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 ...
.
Speciation
As an example of a locale refugia study,
Jürgen Haffer first proposed the concept of refugia to explain the
biological diversity
Biodiversity is the variability of life on Earth. It can be measured on various levels. There is for example genetic variability, species diversity, ecosystem diversity and phylogenetic diversity. Diversity is not distributed evenly on Eart ...
of bird populations in the
Amazonian river basin. Haffer suggested that climatic change in the late
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 ...
led to reduced reservoirs of habitable forests in which populations become allopatric. Over time, that led to
speciation: populations of the same species that found themselves in different refugia evolved differently, creating
parapatric sister-species. As the Pleistocene ended, the arid conditions gave way to the present humid rainforest environment, reconnecting the refugia.
Scholars have since expanded the idea of this mode of speciation and used it to explain population patterns in other areas of the world, such as
Africa
Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
,
Eurasia
Eurasia ( , ) is a continental area on Earth, comprising all of Europe and Asia. According to some geographers, Physical geography, physiographically, Eurasia is a single supercontinent. The concept of Europe and Asia as distinct continents d ...
, and
North America
North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
. Theoretically, current biogeographical patterns can be used to infer past refugia: if several unrelated species follow concurrent
range patterns, the area may have been a refugium. Moreover, the current distribution of
species
A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
with narrow ecological requirements tend to be associated with the spatial position of glacial refugia.
Simple environment examples of temperature
One can provide a simple explanation of refugia involving core temperatures and exposure to sunlight. In the
northern hemisphere
The Northern Hemisphere is the half of Earth that is north of the equator. For other planets in the Solar System, north is defined by humans as being in the same celestial sphere, celestial hemisphere relative to the invariable plane of the Solar ...
, north-facing sites on hills or mountains, and places at higher elevations count as ''cold sites''. The reverse are sun- or heat-exposed, lower-elevation, south-facing sites: ''hot sites''. (The opposite directions apply in the
southern hemisphere.) Each site becomes a refugium, one as a "cold-surviving refugium" and the other as a "hot-surviving refugium". Canyons with deep hidden areas (the opposite of hillsides, mountains, mesas, etc. or other exposed areas) lead to these separate types of refugia.
A concept not often referenced is that of "sweepstakes colonization":
when a dramatic ecological event occurs, for example a meteor strike, and global, multiyear effects occur. The sweepstake-winning species happens to already be living in a fortunate site, and their environment is rendered even more advantageous, as opposed to the "losing" species, which immediately fails to reproduce.
Consequences for disease
It is hypothesized that the creation of a refugium for bats and carnivores in
Thailand
Thailand, officially the Kingdom of Thailand and historically known as Siam (the official name until 1939), is a country in Southeast Asia on the Mainland Southeast Asia, Indochinese Peninsula. With a population of almost 66 million, it spa ...
may have been a necessary step in ''Beta-''
Coronavirus' evolution and
jumping from bats to
civets, and then to humans, which ultimately caused the
Covid-19 Pandemic
The COVID-19 pandemic (also known as the coronavirus pandemic and COVID pandemic), caused by severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2), began with an disease outbreak, outbreak of COVID-19 in Wuhan, China, in December ...
.
Past climate change refugia
Ecological understanding and geographic identification of climate refugia that remained significant strongholds for plant and animal survival during the extremes of past cooling and warming episodes largely pertain to the
Quaternary glaciation
The Quaternary glaciation, also known as the Pleistocene glaciation, is an alternating series of glacial period, glacial and interglacial, interglacial periods during the Quaternary period that began 2.58 Year#SI prefix multipliers, Ma (million ...
cycles during the past several million years, especially in the
Northern Hemisphere
The Northern Hemisphere is the half of Earth that is north of the equator. For other planets in the Solar System, north is defined by humans as being in the same celestial sphere, celestial hemisphere relative to the invariable plane of the Solar ...
. A number of defining characteristics of past refugia are prevalent, including "an area where distinct genetic lineages have persisted through a series of Tertiary or Quaternary climate fluctuations owing to special, buffering environmental characteristics", "a geographical region that a species inhabits during the period of a glacial/interglacial cycle that represents the species' maximum contraction in geographical range," and "areas where local populations of a species can persist through periods of unfavorable regional climate."
Future climate change refugia
In
systematic conservation planning, the term ''refugium'' has been used to define areas that could be used in
protected area
Protected areas or conservation areas are locations which receive protection because of their recognized natural or cultural values. Protected areas are those areas in which human presence or the exploitation of natural resources (e.g. firewood ...
development to protect species from
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 ...
.
The term has been used alternatively to refer to areas with stable habitats or stable climates.
More specifically, the term ''in situ'' refugium is used to refer to areas that will allow species that exist in an area to remain there even as conditions change, whereas ''ex situ'' refugium refers to an area into which
species distribution
Species distribution, or species dispersion, is the manner in which a biological taxon is spatially arranged. The geographic limits of a particular taxon's distribution is its range, often represented as shaded areas on a map. Patterns of distr ...
s can move to in response to climate change.
Sites that offer ''in situ'' refugia are also called ''
resilient sites'' in which species will continue to have what they need to survive even as climate changes.
One study found with downscaled
climate models that areas near the coast are predicted to experience overall less warming than areas toward the interior of the US
State of Washington. Other research has found that
old-growth forests are particularly insulated from climatic changes due to evaporative cooling effects from
evapotranspiration
Evapotranspiration (ET) refers to the combined processes which move water from the Earth's surface (open water and ice surfaces, bare soil and vegetation) into the Atmosphere of Earth, atmosphere. It covers both water evaporation (movement of w ...
and their ability to retain moisture. The same study found that such effects in the
Pacific Northwest
The Pacific Northwest (PNW; ) is a geographic region in Western North America bounded by its coastal waters of the Pacific Ocean to the west and, loosely, by the Rocky Mountains to the east. Though no official boundary exists, the most common ...
would create important refugia for bird species. A review of refugia-focused conservation strategy in the
Klamath-Siskiyou Ecoregion found that, in addition to old-growth forest, the northern aspects of hillslopes and deep gorges would provide relatively cool areas for wildlife and
seeps or
bog
A bog or bogland is a wetland that accumulates peat as a deposit of dead plant materials often mosses, typically sphagnum moss. It is one of the four main types of wetlands. Other names for bogs include mire, mosses, quagmire, and musk ...
s surrounded by mature and old-growth forests would continue to supply moisture even as water availability decreases.
Beginning in 2010 the concept of
geodiversity (a term used previously in efforts to preserve scientifically important geological features) entered into the literature of
conservation biologists as a potential way to identify climate change refugia and as a surrogate (in other words, a proxy used when planning for protected areas) for biodiversity.
While the language to describe this mode of conservation planning hadn't fully developed until recently, the use of geophysical diversity in conservation planning goes back at least as far as the work by Hunter and others in 1988, and Richard Cowling and his colleagues in
South Africa
South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
also used "spatial features" as surrogates for
ecological processes in establishing conservation areas in the late 1990s and early 2000s. The most recent efforts have used the idea of ''land facets'' (also referred to as ''geophysical settings'', ''enduring features'', or ''geophysical stages''
), which are unique combinations of
topographical features (such as slope steepness, slope direction, and
elevation
The elevation of a geographic location (geography), ''location'' is its height above or below a fixed reference point, most commonly a reference geoid, a mathematical model of the Earth's sea level as an equipotential gravitational equipotenti ...
) and
soil
Soil, also commonly referred to as earth, is a mixture of organic matter, minerals, gases, water, and organisms that together support the life of plants and soil organisms. Some scientific definitions distinguish dirt from ''soil'' by re ...
composition, to quantify physical features.
The density of these facets, in turn, is used as a measure of geodiversity.
Because geodiversity has been shown to be correlated with biodiversity,
even as species move in response to climate change, protected areas with high geodiversity may continue to protect biodiversity as
niches get filled by the influx of species from neighboring areas.
Highly geodiverse protected areas may also allow for the movement of species within the area from one land facet or elevation to another.
Conservation scientists, however, emphasize that the use of refugia to plan for climate change is not a substitute for fine-scale (more localized) and traditional approaches to conservation, as individual species and
ecosystem
An ecosystem (or ecological system) is a system formed by Organism, organisms in interaction with their Biophysical environment, environment. The Biotic material, biotic and abiotic components are linked together through nutrient cycles and en ...
s will need to be protected where they exist in the present.
They also emphasize that responding to climate change in conservation is not a substitute for actually limiting the causes of climate change.
See also
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Notes
References
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{{Extinction
Biogeography
Biomes
Habitat
Population ecology
pt:Teoria dos Refúgios