Literature Review on Glacial Refugia

History of Earth describes the most important events in its development till the present day where
organisms have persisted over past through long periods of climate change like glacial and inter
glacial periods. Among many geological time, Pleistocene is considered to be one of the most
important time for evaluating the impact of climate change on biodiversity because the recent
glaciation known as Latest Glacial Maximum (LGM) occurred around 21,000 years ago
(Svenning, Normand and Kageyama, 2008). This led to serious argument about what happened to
the species of plants and animals that were living during this glaciated time? Two different types
of circumstances were considered, first circumstance suggest that all organisms were migrated
towards north of the southern boundary into completely uncovered land after the retreat of
Pleistocene glaciers i.e. tabula rasa hypothesis, and secondly, the organisms survived within
glaciated region i.e. glacial survival hypothesis or in glacial refugia i.e. nunatak hypothesis (Dahl,
1987).

Source: Crawford (2014)

Figure 1: Ariel View of Lake Hazen at high altitude providing support to 117 flowering plant.

Glacial refugia, defined as a geographical regions where various species of flora and fauna have
survived during the ice ages time and have recolonized after the post glacial events irrespective of
the geographical position or spatial extent of the location (Holderegger and Thiel-Egenter, 2009).
Glacial refugia is considered to be one of the major cause for the distribution of many species of
organisms because these were isolated areas suitable for habitat during unfavorable environmental
condition and likely have retained the environmental condition subsequent for recolonization when

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 Literature Review on Glacial Refugia

there is change in environmental condition to became more favorable (Aradhya et al., 2017).
Recently the study of glacial refugia has been increased due to the following reasons:
• Firstly, glacial refugia are the main source of knowledge for understanding postglacial
redistribution, so these regions studying glacial refugia and postglacial recolonization
pattern can help reveal the history and structure of different populations (Shen, Chen and
Li, 2018).
• Secondly, the isolation in glacial refugia may have led to the long-term evolution of modern
day biodiversity or new species or subspecies i.e. genetic divergence and natural selection
(Barnard-Kubow, Debban and Galloway, 2015).
• Lastly, glacial refugia are important areas or regions for maintaining the biodiversity for
long time as these locations have relatively high biodiversity (Aradhya et al., 2017).

Source: Crawford (2014)

Figure 2: View of West Facing Cliff on Prince Charles Foreland showing Semi Nunatak

The possible glacial refugia could be the mountain peaks and outcrops that are projected above the
ice-covered landscape, known as nunataks, and also can be found in the regions where there are
no nunataks and land is free of ice, known as thermal oases (Crawford, 2014). A good example for
glacial refuse be Lake Haze in Greenland, as we can see in figure 1 above, which has a flora of
about 117 different vascular species. A different and most frequent type of refugium is seen neat
the oceanic area, called semi nunatak, where the mountain face facing towards the sea has no or
little ice on it and the mountain face facing towards land is covered with ice. For example,
Spitsbergen on the west coast of Prince Charles Foreland as we can see in figure 2 above

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 Literature Review on Glacial Refugia

(Crawford, 2014). Nunatak are also known as glacial islands which are readily recognizable
landmark point in glaciers or ice fields which are isolated where only some animals and plants life
can survive. So, the nunatak hypothesis discuss about the concept of survival of plants and animal
species in formally glaciated areas which have harsh and inhospitable environment on areas such
as nunataks. This theory was first formally purposed by Sernander in 1896 and became widely
popular in mid-20th century when biologist working on flora of Greenland and Scandinavia
established the areas that were ice free during last glacial maximum (Dahl, 1987).

Source: Holderegger and Thiel-Egenter (2009)

Figure 3: Three different types of Glacial Refugia during Glaciation

There are basically three types of glacial refugia namely nunatak, peripheral refugia and lowland
refugia which are defined geographically with respect to the mountain range as shown in figure 3
above (Holderegger and Thiel-Egenter, 2009).

A. Nunatak Glacial Refugia:

A nunatak is typically an exposed area of mountains or a rocky ridge or a peak that was
once isolated from continental ice sheet or snow within an ice field or glacier or alpine type
ice cap as shown in figure 4 below (Holderegger and Thiel-Egenter, 2009). These are the
places that are projected above continental ice sheet where isolated colonies of various
animals and plants could survive the harsh condition and later can serve as the center for
rapid reoccupation of the land later deglaciated landscape (Crawford, 2014).

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 Literature Review on Glacial Refugia

Source: Encyclopedia Britannica (2018)

Figure 4: Ariel View of Nunataks seen on the Eastern Coast of Greenland

B. Peripheral Glacial Refugia:

Generally, the Glacial Refugia that is location at the border of the mountain system but still
within that mountain system and still above the glaciers where the species have survived
for postglacial recolonization is known as Peripheral Glacial Refugia (Holderegger and
Thiel-Egenter, 2009). According to Holderegger and Thiel-Egenter (2009) “the peripheral
glacial refugia were located along the border of formally glaciated mountain systems such
as the Carpathians, Pyrenees or European Alps”.

C. Lowland Glacial Refugia:

The Glacial refugia that provided suitable habitat for species and organisms outside the
mountain system, i.e. in the adjacent mountain system, before the beginning of glaciation
and beyond the limit of the ice shields is known as Lowland Glacial Refugia (Holderegger
and Thiel-Egenter, 2009). These are the places that are considered to be outside of the
glaciation or continental ice layer but are still directly influenced by the glaciation or ice
layer which makes it hard to detect by the means of biogeography or phylogeographical
method. According to Holderegger and Thiel-Egenter (2009) “the mountain plant,
Oxytropis campestris, currently growing throughout the European Alps, probably survived
the ice ages in refugia of the adjacent lowlands”. Also, the arctic alpine plant Arabis alpin

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 Literature Review on Glacial Refugia

are located in lowland glacial refugia between the ice shields of the European Alps and
Scandinavian glaciers.

The Last Glacial Maximum (LGM) suitably refers to that coldest time period of the earth history
between 26,500 and 20,000 years ago when large glaciers and global ice sheet formed and reached
their extreme size (Svenning, Normand and Kageyama, 2008). According to Birks and Willis
(2008), during the Last Glacial Maximum (LGM) the alpine grew beyond the ice sheet and spread
and colonized rapidly to open habitats as they were made available by ice retreat. The two rare
arctic species of arctic flora, Arenaria humifusa and Sagina caespitosa, found on both sides of
Atlantic from North-Eastern America via Greenland to Scandinavia and Spitsbergen with no
existence in Southern and Central European mountain ranges provides a genetic evidence for
glacial survival within the area of glacial cover at the Last Glacial Maximum (Crawford, 2014).
These two species are known for their ability to survive in cold environment and have similar
distribution as shown in figure 5 below. According to Holderegger and Thiel-Egenter (2009), the
lowland glacial refugia, located between the ice shields of European Alps and Scandinavian
glaciers, has helped survive the arctic plant called Arabis alpina which was already occupied by
A. alpine before the last glaciation.

Source: Crawford (2014)

Figure 5: Distribution pattern of two different arctic species of flora (a) Arenaria humifusa and (b) Sagina
caespitosa providing evidence for glacial refugia

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There are many studies revealing that flora and fauna of North America have Bering Sea region,
high-altitude zone of North Pole, the southern Appalachian Mountains and the southern Rocks
Range as the main refugia with species namely Poke milkweed (Asclepias exaltata), Dryas
integrifolia, Microtus longicawdus, and Arctic grayling (Thymallus arcticus) representing four
types of glacial refugia and recolonization routes in Northern America. Similarly, there are three
southern peninsulas namely Iberian, Italion and Balkan, important refugia for terrestrial flora and
fauna in Europe during Pleistocene glaciation (Shen, Chen and Li, 2018). Recently, with the
development of molecular markers and allozyme markers, used to identify the refugium and track
the colonization route and from the studies of palynology, paleoclimatology, phylogeographic,
palaeobotany and fossils have provided some evidences for identifying the glacial refugia far north
of the southern European peninsulas and the Alps, also known as cryptic refugia (Homburg et al.,
2013).

In conclusion, glacial refugia are the places that were isolated during the glaciation time providing
suitable habitat for plant and animal species which in early post glaciation period descends to
milder climate and helped rapid spread of species and may also led to genetic divergence among
the population. The discussion above in a simple way helps clarifying the three different types of
glacial refugia, namely nunatak, peripheral and lowland refugia, in biogeography and
phylogeography of mountain. There are evidences presented above are sufficient to show some
possible cases of glacial refugia of plant species with respect to genetic structure as well as the
pattern of flora and fauna distribution.

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References:

Aradhya, M., Velasco, D., Ibrahimov, Z., Toktoraliev, B., Maghradze, D., Musayev, M., Bobokashvili, Z.
and Preece, J. (2017). Genetic and ecological insights into glacial refugia of walnut (Juglans regia
L.). PLOS ONE, [online] 12(10), p.e0185974. Available at:
http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0185974 [Accessed 18 Jan. 2018].

Ashcroft, M. (2010). Identifying refugia from climate change. Journal of Biogeography, [online] p.no-no.
Available at: http://citeseerx.ist.psu.edu/viewdoc/download?doi=10.1.1.459.5428&rep=rep1&type=pdf
[Accessed 17 Jan. 2018].

Barnard-Kubow, K., Debban, C. and Galloway, L. (2015). Multiple glacial refugia lead to genetic
structuring and the potential for reproductive isolation in a herbaceous plant. American Journal of Botany,
[online] 102(11), pp.1842-1853. Available at: http://www.amjbot.org/content/102/11/1842.full [Accessed
18 Jan. 2018].

Birks, H. and Willis, K. (2008). Alpines, trees, and refugia in Europe. Plant Ecology & Diversity, [online]
1(2), pp.147-160. Available at: http://www.tandfonline.com/doi/pdf/10.1080/17550870802349146
[Accessed 18 Jan. 2018].

Crawford, R. M. M. (2014). Tundra-taiga biology. 1st ed. New York: Oxford University Press, pp.23-28.

Dahl, E. (1987). The Nunatak Theory Reconsidered. Ecological Bulletins, [online] 28, pp.77-94. Available
at: http://www.jstor.org/stable/20112974 [Accessed 18 Jan. 2018].

Encyclopedia Britannica. (2018). Nunatak | geology. [online] Available at:

https://www.britannica.com/science/nunatak [Accessed 17 Jan. 2018].

Holderegger, R. and Thiel-Egenter, C. (2009). A discussion of different types of glacial refugia used in
mountain biogeography and phylogeography. Journal of Biogeography, [online] 36(3), pp.476-480.
Available at: http://onlinelibrary.wiley.com/doi/10.1111/j.1365-2699.2008.02027.x/epdf [Accessed 17 Jan.
2018].

Homburg, K., Drees, C., Gossner, M., Rakosy, L., Vrezec, A. and Assmann, T. (2013). Multiple Glacial
Refugia of the Low-Dispersal Ground Beetle Carabus irregularis: Molecular Data Support Predictions of
Species Distribution Models. PLoS ONE, [online] 8(4), p.e61185. Available at:
http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0061185 [Accessed 18 Jan. 2018].

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Naczk, A. and Kolanowska, M. (2015). Glacial Refugia and Future Habitat Coverage of Selected
Dactylorhiza Representatives (Orchidaceae). PLOS ONE, [online] 10(11), p.e0143478. Available at:
http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0143478#sec005 [Accessed 17 Jan.
2018].

Pennington, W. (1965). The Interpretation of Some Post-Glacial Vegetation Diversities at Different Lake
District Sites. Proceedings of the Royal Society B: Biological Sciences, [online] 161(984), pp.310-323.
Available at: http://rspb.royalsocietypublishing.org/content/royprsb/161/984/310.full.pdf [Accessed 18
Jan. 2018].

Roberts, D. and Hamann, A. (2015). Glacial refugia and modern genetic diversity of 22 western North
American tree species. Proceedings of the Royal Society B: Biological Sciences, [online] 282(1804),
pp.20142903-20142903. Available at: http://rspb.royalsocietypublishing.org/content/282/1804/20142903
[Accessed 17 Jan. 2018].

Shen, L., Chen, X. and Li, Y. (2018). Glacial Refugia and Postglacial Recolonization Patterns of
Organisms. [online] Europepmc.org. Available at: http://europepmc.org/abstract/cba/390842 [Accessed 17
Jan. 2018].

Svenning, J., Normand, S. and Kageyama, M. (2008). Glacial refugia of temperate trees in Europe: insights
from species distribution modelling. Journal of Ecology, [online] 96(6), pp.1117-1127. Available at:
http://onlinelibrary.wiley.com/doi/10.1111/j.1365-2745.2008.01422.x/full [Accessed 17 Jan. 2018].

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