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in zhe Alps

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an endangered authochtonous breed

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diffusion and economic and morphological characteristics

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Italian breeds of cattle: Pustertaler - Origins, diffusion and economic and morphological characteristics

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I know, off topic, but I just couldn't resist ;)

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And the marmot, it puts the chocolate in the alu-foil

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Alpine marmot videos, photos and facts - Marmota marmota | Arkive

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See videos and photos of the Alpine marmot on Arkive. Classified as Least Concern (LC) on the IUCN Red List. Subspecies Marmota marmota marmota, M. m. latirostris .

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Typical Alpine Animal: Apline Capricorn - habitat, distribution and ecological niche

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Typical Alpine Animal: Apline Capricorn - habitat, distribution and ecological niche

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Alpine Ibex - Facts, Diet, Habitat & Pictures on Animalia.bio

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Basic facts about Alpine Ibex: lifespan, distribution and habitat map, lifestyle and social behavior, mating habits, diet and nutrition, population size and status.

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A tapeworm that can be fatal to humans has been discovered in a Swedish fox, marking the first time the parasite has been found in Sweden.

If more animals are found to be infected by the Echinococcus multilocularis parasite, Swedes may have to stop eating fresh berries picked right from the forest.

"This is very serious. It has never before been found in Sweden," Carl Hård af Segerstad, division head at the National Veterinary Institute (Statens veterinärmedicinska anstalt, SVA), told news agency TT late on Sunday.

Up to 400 foxes are tested each year in an effort to detect the parasite. Last week, samples were taken from a suspected infected fox in northern Bohuslän in southwestern Sweden. On Friday, the infection was confirmed.

Tapeworm eggs can end up on berries and mushrooms through animal faeces. For dogs, cats and foxes, the parasite is relatively harmless. However, people must take precautions to prevent infection.

"It can form cysts in internal organs. If someone becomes infected, he or she will have to undergo lifelong anti-parasite treatment and survival is not certain," said Hård af Segerstad.

The worm is one of the reasons that dogs and cats entering Sweden must be dewormed. The parasite is found in many parts of Europe and is most common in the Alps, where people are discouraged from picking and eating mushrooms and berries in the countryside.

"If we ingest the eggs that the fox has crapped out in the woods, there is a risk that we'll get infected. As a result, we may need to change our habits in the forest, for example not picking berries from a stem and eating them," said Hård af Segerstad.

He emphasised that the worm has only been found in one animal. As such, he estimated that the infection has not been present here for very long.

"Now we have to start analysing as many foxes and small rodents as we can from the area to see how widespread the infection may be. There is no risk of infection now. There is nothing in the woods that one can put in one's mouth right now," he said.

A number of agencies, including the National Board of Health and Welfare (Socialstyrelsen) and the Swedish Institute for Communicable Disease Control ( Smittskyddsinstitutet), are expected to meet soon to develop an action plan for how to manage the parasite's entry into Sweden.

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A tapeworm that can be fatal to humans has been discovered in a Swedish fox, marking the first time the parasite has been found in Sweden. If more animals are found to be infected by the Echinococcus multilocularis parasite, Swedes may have to stop eating fresh berries picked right from the forest.

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Your BioArea needs some elaboration and enrichment.

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Your BioArea needs some elaboration and enrichment.

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Great pictures and ideas!

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I find this aspect of biodiversity very interesting...

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SAN FRANCISCO - The future of food is often defined by cutting-edge technologies like lab-grown meat or blockchain. Mike Lee, a Brooklyn, N.Y.-based food product developer and co-founder of Alpha Food Labs, argues for a simpler solution - we need to grow and eat different plants.

"Seventy-five per cent of our food comes from just 12 plants and 5 animals," Mr. Lee said during a presentation at the Winter Fancy Food Show in San Francisco. "We only eat about 150 out of 30,000 edible plant species in the world ...

"You're literally leaving on the table a whole variety of interesting flavors, tastes and food experiences."

Biodiversity, he said, is the intersection of taste and sustainability. The variety and variability of plants, animals and organisms within an ecosystem supports a healthier planet and people.

"Even if you don't care about the future of the planet or soil health or climate change - all of which biodiverse food systems support - everyone, I think, cares about finding delicious, tasty food," Mr. Lee said. "The power of biodiversity is that it links together a very altruistic need - saving the planet, reducing climate change - with a very selfish need - I want tasty stuff."

A display at the Winter Fancy Food Show spotlighted two dozen crops cultivated around the world, including grains and seeds such as teff, Kernza, amaranth and fonio; tubers, including yacon and ulluco; and beans such as cowpea, lupin and lablab bean. Jackfruit, moringa, baobob and quinoa also were included.

"These are 24 opportunities for you to differentiate your assortment, wow your customer and save the planet at the same time," Mr. Lee said. "It's a win-win-win."

General Mills, Inc.; Starbucks Corp.; Unilever's Sir Kensington's business; Thrive Market; and Back to the Roots, Inc. are among the companies that are exploring opportunities to incorporate more biodiversity in their businesses, Mr. Lee said.

"This is a conversation that has teeth that the industry is really looking at," Mr. Lee said.

Suppliers and producers exhibiting at the Winter Fancy Food Show highlighted products featuring underutilized ingredients. A brand named Wild For makes teff chips in sea salt, sour cream and onion, and sweet and smoky barbecue flavors. The products contain twice the protein and half the fat of other snack chip brands, according to the San Francisco-based company.

The Jackfruit Co., Boulder, Colo., produces a range of meal bowls and starters featuring jackfruit, a large tree fruit with a meaty texture and neutral flavor.

Yacon root adds sweetness and prebiotic fiber to Rowdy Bar, a Sparks, Nev.-based energy bar brand with three flavor varieties: chocolate coconut cashew, peanut dark chocolate, and sunflower butter and berries. Patagonia Provisions, Sausalito, Calif., offers a craft beer made with Kernza, a perennial long-rooted grain that grows in Kansas. Kuli Kuli, Oakland, Calif., makes beverages, smoothie mixes and energy bars with moringa, a nutrient-dense, sustainably grown tree sourced from Ghana.

A supplier at the show hopes to bring the bambara bean, another underutilized crop, into more American diets. Believe in Bambara sources the versatile and nutritious legume from a women's farming cooperative in Ghana. Bambara beans may be processed as a gluten-free flour or protein isolate, said founders Holly Tassi and Tamara Cohen, who are based in Los Angeles and New York, respectively.

TerViva, an Oakland start-up, is cultivating pongamia, a tree that is native to Australia and India, in the United States. Pongamia may be used in various applications, including vegetable oil, flour, protein and animal feed. Its beans produce up to 10 times more oil and up to 5 times more protein per acre than soybeans, according to the company.

"We're all here to look at what the next big thing is going to be," Mr. Lee said. "Innovate your way to a more sustainable food system."

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Forget lab-grown meat and blockchain. This is the future of food

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SAN FRANCISCO - The future of food is often defined by cutting-edge technologies like lab-grown meat or blockchain. Mike Lee, a Brooklyn, N.Y.-based food product developer and co-founder of Alpha Food Labs, argues for a simpler solution - we need to grow and eat different plants.

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Alpine Convention - The Convention - The Alpine Convention in a nutshell - The Alps - Home

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Food partitioning of 3 shrew species from European Alps

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Food partitioning of 3 shrew species from European Alps

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Leafcutter ants and their antibiotics at the Royal Society Summer Science Exhibition 2014

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Theme 2, Q1: Reason  for the decision to reintroduce grey wolves into the Yellowstone National Park

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Theme 2, Q1: Reason  for the decision to reintroduce grey wolves into the Yellowstone National Park

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The wolves were brought in because the increased elk population was overgrazing the deciduous, woody species such as aspen and cottonwood

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Alparc - the Alpine Network of Protected Areas

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It is difficult to visit the Alps without becoming curious as to how the majestic mountains were formed. The Alps rose up due to the collision of the African and European tectonic plates, but the present-day landscape with iconic jagged mountains such as the Matterhorn are the result of millions of years of glacial activity and erosion. Here is a short guide to the geology of the Alps from the big bang to the present day:
15bn years ago: The Big Bang 4.5bn years ago: The early earth was formed 250 million years ago: Pangea

The earth is very hot and dry with a lot of volcanic activity.

The world's land mass is comprised of one supercontinent called Pangea.

At this stage the rocks which will later form Mont Blanc and the Matterhorn are lying dormant under sandy coastal dunes.

240 million years ago: First Dinosaurs begin to walk on earth 220 million years ago: Pangea splits into Laurasia and Gondwanaland

Convection currents in the earths mantle cause the continental crust of Pangea to split forming 2 continents - Laurasia and Gondwanaland - with an ocean called the Tethys sea between them.

Tectonic activity cause Laurasia and Gondwanaland to move further and further apart and the Tethys sea grows by 2cm per year.

The rocks that will form the future Mont Blanc are part of the Laurasian continent and the rocks that will form the Matterhorn are part of the Gondwanaland continent. The geology of the Alps is becoming more and more recognizable (at least for modern hikers).

160-120 million years ago: The Tethys sea continues to expand, and the Atlantic Ocean begins to form 130 million years ago: Continents continue to split apart
Convectional currents in the earth's mantle cause the continents to begin to split apart. Most notably modern-day India detaches from the African plate and begins its long migration northwards. It will later collide with Asia and this collision will form the Himalaya.
60 million years ago: Mass extinction

A 10km wide asteroid hits the earth with a force that is 1 million times more powerful than a nuclear bomb. This leads to the extinction of all dinosaurs and ammonites.

At this stage the Alps is still under water, which is the reason why so much of the rock found in the Alps is formed of sedimentary limestone.

35 - 20 million years ago: Continents collide, and the Alps begin to form

The North part of the African plate collides with the European plate and is thrust over the European plate. This is the start of the formation of the Alps which grows by 1000m per million years.

The Matterhorn is an interesting example of this collision:

The base of the Matterhorn contains rocks which were formed under the Tethys sea. As the African plate moved into the Eurasian plate the denser oceanic crust was subducted under the lighter, continental crust.

The centre of the Matterhorn contains rocks from the Eurasian plate.

The top of the Matterhorn comprises of rocks from the African plate which was thrust up on top of the Eurasian plate when it collided together.

5 million years ago: Early humans are evolving in Africa

Favourable climatic conditions in the great rift valley of Africa leads to the evolution of our early ancestors.

The earliest footprints of our ancestors can be found preserved in volcanic ash in Tanzania and show an adult and child walking upright on two legs side by side - they are believed to date back to 3.5 million years ago.

The Arabian plate joins the Eurasian plate and early hominids can migrate to Asia and Europe.

2.4 million - 1.6 million years ago: The Quaternary Period

The Quaternary period is a geological time period characterised by progressive cooling.

There were cycles of 100,000 year glacials (cooler periods in which glaciers advance), interspersed with 15-20,000 year inter-glacials (warmer periods in which glaciers retreat).

100,000 years - 40,000 years ago: The last great ice age

The Alps, the Swiss Plateau and the Jura mountains were completely covered in Ice.

The Rhone Glacier reached all the way to where the modern-day city of Lyon stands.

100,000 years ago - the first homo sapiens evolved.

40,000 years ago - homo sapiens displaced Neanderthals at the end of the last great ice age.

18,000 years ago: Climate begins to warm up again

The sea level which was 120 metres lower than today, begins to rise.

The Rhone Glacier retreats to Geneva but is still immense; reaching 700m above the present-day cities of Lausanne and Evian.

10,000 years ago - Present day: Holocene Period

At lower altitudes the warming climate means allows pine and birch forests to give way to hazel, oak, ash and linden trees.

The absence of ice on mountain sides causes huge amounts of rock fall, forming the iconic steep sided faces of the Alps that we see today.

If you would like to find out more about the geology of the Alps, an excellent resource is the book 'The Alps and our Planet, The African Matterhorn: A Geological Story' by Michel Marthaler.

If you are interested in seeing the magnificent geology of the Alps firsthand, you may want to consider a trekking tour of the Alps.

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How were the Alps formed? A guide to the geology of the Alps

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Curious about how the Alps were formed? They rose up due to the collision of the African and European tectonic plates, but the present-day landscape is the result of millions of years of glacial activity and erosion. Here is a short guide to the geology of the Alps, from the Big Bang to the present.

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Alpine Seed Conservation and Research Programme

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Alpine Seed Conservation and Research Programme

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by the Alpine Plant Conservation and Research Programme
The Alps - a European hotspot of phytodiversity

The European Alps harbour about 4450 vascular plant taxa, with a density of 2200 taxa per 10,000 km² (Aeschimann et al. 2011a). Together with some Mediterranean areas and the Pyrenees, we find the highest phytodiversity in Europe in this area. The most species rich areas of the Alps are in the west and the south, while the most endemic-rich areas are those that were glacier-free during the Pleistocene, as well as the south-west and the very eastern parts of the Alps (Tribsch & Schönswetter 2003, Tribsch 2004, Aeschimann et al. 2011b). While most of the central European grassland at low altitudes results from cattle grazing or mowing, we find in the alpine belt "native grassland" as an ancestral habitat of many of the widely cultivated grassland species found today.

The ecological conditions for life are extreme here. With every 100 m increase in altitude the mean annual temperature decreases 0.7 degrees, leading to the characteristic vegetation zonation of high-montane forests, subalpine Krummholz with shrubs like Pinus mugo or Alnus alnobetula, and alpine grassland above the timberline. Elevation elongates the frost period, and reduces the vegetation period - the time during the year when plant growth is possible. In the same way, bedrock weathering, soil and humus development and especially microbial activities providing mineral nutrients for plant growth are reduced. Unlike arctic regions with similar conditions, solar radiation is high with an increasing amount of dangerous UV radiation and nocturnal back-up radiation with altitude. All life forms have to face huge differences between day and night temperatures and radiation. Precipitation is high, but the thin soils do not steadily provide water and nutrients to plant roots. Strong rain or snow melt cause floods that float the few nutrients away, while the soils dry out quickly in the hot radiation of sunny days.

Seeds can be easily dispersed by wind in the Alps, but the winds are too strong for pollination and insect pollination predominates. Insects have evolved in conjunction with the most species-rich group in the Alps, with high rates of speciation and endemism, but with low individual numbers due to the harsh ecological conditions. Plants have to produce conspicuous flowers to attract insects in this harsh environment; one of the reasons we consider the alpine flora extraordinarily beautiful. Plants have also had to evolve a set of vegetative and regenerative adaptations to survive here on the physiological border of the Earth's biosphere.

This short description of the natural conditions in alpine environments makes it clear that alpine life is a highly complex interactive system that has evolved over millions of years. Every link in the system has its own role in this ecological interplay, and the system is highly endangered if one link is lost - making Alpine belt ecosystems more sensitive to disturbance than most other ecosystems in the world.

The endangerment of the flora of the Alps by habitat loss and tourism

The European Alps are a long-lasting region of pre-industrial land use. The pasture-like alpine grasslands invited ancestral people to bring cattle (mainly sheep and cows) to the alpine zone during the summer, and hey from the meadows was taken down to the valleys for winter feed. As result a diverse, mosaic landscape with montane meadows developed. While the alpine grassland expanded, the timberline and Krummholz-belt were relegated dozens of meters to lower altitudes. The impact on the native flora was low; many species of the alpine belt were adapted to light grazing by the natural pressure of wild chamois and Capricorns.

After the development of industrial agriculture in the second half of the 20th century the picture changed. Pre-industrial agriculture was not competitive any more, and tourism - notably winter sports - developed as a new source of income for the local population. This altered whole landscapes, robbed them of their grandiosity, and depreciated the Alps to the backdrop (Niedrist et al. 2009). The vegetation in winter sport areas is replaced, or at least strongly altered, with a clear decline in the sensitive alpine flora (Rusterholz et al. 2004, Wipf et al. 2005, Negro et al. 2010).

Winter tourism is, however, highly endangered by global warming. The answer of the tourism branch is to build up a giant and expensive infrastructure for artificial snow production (e.g. water basins above ski slopes), and move the ski slopes to higher altitudes or to northern exposures. The impact of the tourism industry to the nature of the Alps has reached a new level at the 21th century.

As a result, farmers concentrate their cattle in the remaining areas, or give up completely, losing their good, accessible pastures and find new jobs in tourism. Both have negative effects on the alpine flora. High densities of cattle facilitate the negative effects of grazing on the vegetation, preventing many species from producing seeds, with long-term negative effects on population development. Threatened species are more sensitive to the impacts of over-grazing e.g. dwarf shrubs, cushion and wetland plants. In areas were livestock are completely removed, high montane forest and Krummholz recolonise from lower altitudes, reducing the habitat of alpine grassland species.

Climate change - the main threat of the alpine flora

The example of winter sports areas show the indirect impact of global warming on alpine plants through exacerbated habitat loss. The direct effects of climate change on alpine ecosystems are even greater, and will be the much bigger challenge for nature conservation in the 21 st century. Since the 1990s, the mean temperature of the Alps has been well above the long-term mean, and far beyond the global mean (Beniston 2006). Climate scenarios predict a temperature increase of 4 to 4.5 degrees by 2100, with less precipitation in the Western Alps and slightly more precipitation in the Eastern Alps (Heimann & Sept 2000, Beniston 2005). In addition, there will be less snow cover and a reduction of permafrost across the Alps (Beniston 2006, Harris et al. 2009).

Climate change will affect global biodiversity (Malcolm et al. 2006). According to the vegetation sensitivity index (Seddon et al. 2016), arctic and alpine ecosystems are most sensitive to climate change. For a long time, the shrinking of alpine glaciers was used as the most visible and spectacular indicator of climate change (Haeberli et al. 2007). The reaction of plant cover is different (Dirnböck et al. 2003, Erschbamer 2007) and not easy to explain (Theurillat & Guisan 2001, Rixen et al. 2014). According to most predicted scenarios, a general up-shift of plant populations (Pauli et al. 1996, Erschbamer et al. 2009), vegetation units (Brzeziecki 1995) and the timberline (Gehrig-Fasel et al. 2007) can be expected, as well as exceptions for individuals and populations that use microclimatic heterogeneity of a particular habitat to remain at the same altitude (Maclean et al. 2015). It is also known, that small range species with small populations are more affected by climate change (Ohlemüller et al 2008). Even if we consider that the retreat of the glaciers provide additional habitats for alpine and subnival species in a "cooled" glacier mesoclimate, all summits are finite and the cone shape of summits results in a permanent reduction of the available surface area at higher altitudes.

In our project, we want to get a deeper insight into plant reactions to these pressures in light of increasing future endangerment, from reproduction parameters like seed production and germination strategies to the ecological level.

Aeschimann, D., N. Rasolofo & J.-P. Theurillat (2011a). Analysis of the flora of the Alps. 1: historical account and biodiversity. Candollea 66: 27-55.

Aeschimann, D., N. Rasolofo & J.-P. Theurillat (2011b). Analysis of the flora of the Alps. 2: biodiversity and chorology. Candollea 66: 225-253.

Beniston, M. (2005). Mountain Climates and Climatic Change: An Overview of Processes Focusing on the European Alps. Pure and Applied Geophysics 162, 1587-1606.

Beniston, M. (2006): Mountain Weather and Climate: A General Overview and a Focus on Climatic Change in the Alps. Hydrobiologia 562: 3-16.

Brzeziecki, B., Kienast, F. & Wildi, O. (1995). Modelling potential impacts of climate change on the spatial distribution of zonal forest communities in Switzerland. Journal of Vegetation Science 6: 257-268.

Dirnböck, T., Dullinger, S., Grabherr, G., 2003. A regional impact assessment of climate and land-use change on alpine vegetation. Journal of Biogeography 30, 401-417.

Erschbamer, B. (2007). Winners and Losers of Climate Change in a Central Alpine Glacier Foreland. Arctic, Antarctic, and Alpine Research 39, 237-244.

Erschbamer, B., Kiebacher, T., Mallaun, M. & Unterluggauer, P. (2009). Short-term signals of climate change along an altitudinal gradient in the South Alps. Plant Ecology 202, 79-89.

Gehrig-Fasel, J., Guisan, A. & Zimmermann, N. E. (2007). Tree line shifts in the Swiss Alps: Climate change or land abandonment? Journal of Vegetation Science 18, 571-582.

Haeberli, W., Hoelzle, M., Paul, F. & Zemp, M. (2007). Integrated monitoring of mountain glaciers as key indicators of global climate change: the European Alps. Annals of Glaciology 46, 150-160.

Heimann, D. & Sept, V. (2000). Climate Change Estimates of Summer Temperature and Precipitation in the Alpine Region. Theoretical and Applied Climatology 66, 1-12.

Harris, C., et al. (2009). Permafrost and climate in Europe: Monitoring and modelling thermal, geomorphological and geotechnical responses. Earth-Science Reviews 92, 117-171

Maclean, I. M. D., Hopkins, J. J., Bennie, J. J. & Lawson, C. R. (2015). Microclimates buffer plant community responses to climate change. Global Ecology & Biogeography, 24, 1340-1350.

Malcolm, J.R., Liu, C., Neilson, R.P., Hansen, L., Hannah, L., 2006. Global warming and extinctions of endemic species from biodiversity hotspots. Conservation Biology 20, 538-548.

Negro, M., Isaia, M., Palestrini, C., Schoenhofer, A. & Rolando, A. (2010). The impact of high-altitude ski pistes on ground-dwelling arthropods in the Alps. Biodiversity and Conservation 19, 1853-1870.

Niedrist, G., Tasser, E., Lüth, C., Dalla Via, J. & Tappeiner, U. (2009). Plant diversity declines with recent land use changes in European Alps. Plant Ecology 202, 195-210.

Ohlemüller, R., Anderson, B.J., Araujo, M.B., Butchart, S.H.M., Kudrna, O., Ridgely, R.S., Thomas, C.D., 2008. The coincidence of climatic and species rarity: high risk to small-range species from climate change. Biology Letters 4: 568-572.

Pauli, H., Gottfried, M. & Grabherr, G. (1996). Effects of climate change on mountain ecosystems - upward shifting of alpine plants. World Resource Review 8, 382-390.

Rixen, C., Wipf, S., Frei, E. & Stöckli, V. (2014). Faster, higher, more? Past, present and future dynamics of alpine and arctic flora under climate change. Alpine Botany 124, 77-79.

Rusterholz, H.-P., Müller, S. W. & Baur, B. (2004). Effects of rock climbing on plant communities on exposed limestone cliffs in the Swiss Jura Mountains. Applied Vegetation Science 7, 35-40.

Seddon, A. W. R., Macias-Fauria, M., Long, P. R., Benz, D. & Willis, K. J. (2016). Sensitivity of global terrestrial ecosystems to climate variability. Nature 531, 229-232.

Theurillat, J. & Guisan, A. (2001). Potential Impact of Climate Change on Vegetation in the European Alps: A Review. Climatic Change 50, 77-109.

Tribsch, A. 2004. Areas of endemism of vascular plants in the Eastern Alps in relation to Pleistocene glaciation. Journal of Biogeography 31, 747-760.

Tribsch, A. & Schönswetter, P. 2003. Patterns of endemism and comparative phylogeography confirm palaeo-environmental evidence for Pleistocene refugia in the Eastern Alps. Taxon 52, 477-497.

Wipf, S., Rixen, C., Fischer, M., Schmid, B. & Stoeckli, V. (2005). Effects of ski piste preparation on alpine vegetation. Journal of Applied Ecology 42, 306-316.
Corresponding author: christian.berg@uni-graz.at

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Alpine Seed Conservation and Research Network

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by the Alpine Plant Conservation and Research Programme The Alps - a European hotspot of phytodiversity The European Alps harbour about 4450 vascular plant taxa, with a density of 2200 taxa per 10,000 km² (Aeschimann et al. 2011a). Together with some Mediterranean areas and the Pyrenees, we find the highest phytodiversity in Europe in this area.

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The Alps are one of the regions with the richest flora and fauna in Europe, second only to the Mediterranean

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The Alps are one of the regions with the richest flora and fauna in Europe, second only to the Mediterranean

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Of about 4,500 vascular plant species in the Alps, nearly 8% are endemic, occurring in the Alps and nowhere else in the world. The uniqueness of Alpine flora stems from the variety of habitats in the Alps and the "harsh" environmental conditions that drive species to change and adapt.

Plants above the forest line, for example, have evolved different growth forms that allow them to survive harsh winds and cold winters. Some plants are small and grow in the form of flat cushions, rosettes, or carpets to protect themselves from wind and to resist the pressure of heavy snow layers.

Others have large root structures and ample underground organs that both anchor the plant and act as a storage for water and nutrients. For example, the net-leaved willow ( Salix reticulata) barely reaches 10-15 cm in height, but has roots several meters long.

The forests below the timberline are considered relatively natural today and serve as important refuges for rare species as well as corridor areas for many others (e.g., large herbivores or large carnivores).

Alpine fauna

The exact number of animal species in the Alps is unknown, though estimates place that number at about 30,000.

With increasing altitude, living conditions become too harsh for animals that have adapted to lower altitude conditions. Alpine species are special in that they have found ways to survive in these difficult climatic conditions, resulting in a variety of unique adaptations.

The Alpine salamander ( Salamandra atra), for example, is the only European amphibian to give birth to fully developed young. Birds and mammals have thicker feathers or pelts and their feet or paws are perfectly adapted for treading on snowy surfaces. The mountain hare ( Lepus timidus), the stoat ( Mustela erminea), and the ptarmigan ( Lagopus mutus) change their brown coats to white in winter to avoid predation.

Many animals hibernate during the coldest months of the year, including the famous Alpine marmot ( Marmota marmota). Others, like the Alpine chamois ( Rupicapra rupicapra) and rock partridge ( Alectoris graeca), migrate over relatively long distances or descend to lower altitudes to avoid the cold of winter. The Alpine ibex ( Capra ibex), on the other hand, climbs to very steep slopes where the snow slides off and food can still be found.

In the aquatic world of the Alps, highly specialized species found in small Alpine lakes and glacial streams contribute to a diverse and characteristic freshwater fauna. But the high sensitivity of these organisms to global warming and human activities threatens their survival.

Though typical Alpine animals continue to exist in the Alps, many of their populations have been reduced in size and cut off from each other. After near extinction, the Alpine ibex survived as a small population in Gran Paradiso National Park in the Italian Alps. Individuals from this remnant population have since been re-introduced to other parts of the Alps allowing the species to reclaim much of its Alpine range.

The plight of the three large carnivores in the Alps - the lynx, bear, and wolf - is another example of how close some Alpine mammals have come to being wiped out from the Alpine arc. Fortunately, through policy changes and reintroduction projects, some of these species are making a come-back in the Alps.

Threats to Alpine nature

The natural richness of the Alps is at risk of disappearing. Habitat loss and fragmentation are the major threats to Alpine biodiversity. Spreading settlements, unsustainable farming, road networks, and river dams are the main culprits.

What WWF is doing

The European Alpine Programme coordinates WWF's conservation work across the Alps. Check out Our Solutions page to read more about what WWF is doing to conserve Alpine nature.

Facts and Figures

With over 30,000 animal species and 13,000 plant species, the Alps are one of the richest biodiversity "hotspots" in Europe.

The number of vascular plants in the Alps represents 39% of the total European flora.

About 8% of the total number of vascular plant species in the Alps are endemic to the region.

The Alpine salamander ( Salamandra atra) is the only amphibian in Europe to give birth to fully developed young. For those living above 1,400 m, the gestation period rivals that of elephants, with young remaining inside their mother for up to 3 years!

Source: Wikipedia

The Alpine marmot ( Marmota marmota) is found throughout the Alps and has adapted to the harsh winters by hibernating in burrows.

Source: Wikipedia

The Alpine ibex ( Capra ibex) has evolved to traverse near vertical cliffs in order to avoid deep snow and find food during the winter.

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Of about 4,500 vascular plant species in the Alps, nearly 8% are endemic, occurring in the Alps and nowhere else in the world. The uniqueness of Alpine flora stems from the variety of habitats in the Alps and the "harsh" environmental conditions that drive species to change and adapt.

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I am living in an "biodiversity hotspot" area

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The Alps - a biodiversity hotspot - Website

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The Alps span eight countries, from the Mediterranean shores of Southern France to Slovenia. With summits above 4000 meters and an area of 250 000 km² the Alps are the most important mountain chain in Central Europe.

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Agricultural Genetic Resources in the Alps

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Agricultural Genetic Resources in the Alps

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A very interesting book on plants and animals of agricultural importance in the european Alps

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Biogeographical and evolutionary importance of the European high mountain systems

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Biogeographical and evolutionary importance of the European high mountain systems

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Abstract

Europe is characterised by several high mountain systems dominating major parts of its area, and these structures have strongly influenced the evolution of taxa. For species now restricted to these high mountain systems, characteristic biogeographical patterns of differentiation exist. (i) Many local endemics are found in most of the European high mountain systems especially in the Alps and the more geographically peripheral regions of Europe. Populations isolated in these peripheral mountain ranges often have strongly differentiated endemic genetic lineages, which survived and evolved in the vicinity of these mountain areas over long time periods. (ii) Populations of taxa with wide distributions in the Alps often have two or more genetic lineages, which in some cases even have the status of cryptic species. In many cases, these lineages are the results of several centres of glacial survival in the perialpine areas. Similar patterns also apply to the other geographically extended European high mountain systems, especially the Pyrenees and Carpathians. (iii) Populations from adjoining high mountain systems often show similar genetic lineages, a phenomenon best explained by postglacial retreat to these mountains from one single differentiation centre between them. (iv) The populations of a number of species show gradients of genetic diversity from a genetically richer East to a poorer West. This might indicate better glacial survival conditions for this biogeographical group of species in the more eastern parts of Europe.

Background

The evolution within species and sibling species complexes in Europe is closely connected to the range shifts caused by the cyclic changes between relatively short and warm, mostly humid and longer cold and predominantly dry periods [ 1, 2]. These climatic shifts resulted in large scale range shifts in many species often resulting in disjunct distribution pattern during at least one of these phases [ 3, 4]. However, the effects of these climatic fluctuations vary considerably in different ecological and distributional groups [ 5, 6], and three major groups can be distinguished: Mediterranean, continental and arctic/alpine species [ 2], which are all to some degree influenced in their biogeography and evolution by the complexity of the European high mountain systems.

The Mediterranean species survived the cold stages of the Pleistocene in the Mediterranean Basin and expanded northwards during the postglacial [ 7, 8]. These expansions are largely shaped by the high mountain systems of the Alps and Pyrenees acting as dispersal barriers of different strength so that four different paradigms of postglacial range expansions from the three Mediterranean peninsulas can be distinguished [ 9, 10]. In contrast to the assumptions made by de Lattin [ 6], the continental species, which in most cases have not immigrated into Europe from Asiatic core areas during the postglacial, had so called extra-Mediterranean refuge and differentiation centres north of the typical Mediterranean refugia. These were also strongly influenced by the European mountain systems because they offered suitable refuge areas in their non-glaciated parts during ice ages and acted as dispersal barriers during postglacial range shifts [e.g. [ 11- 19]].

The group of alpine and arctic-alpine species is most strongly shaped in their biogeographical and evolutionary history by the geographic location and complexity of the different European high mountain systems because (i) these species are now restricted to these mountain ranges and (in some cases) the arctic and (ii) most of their actual distribution areas were covered by extensive ice-shields during the glacial periods [ 5, 20]. In general, we can distinguish between several different distributional types in these mountain species. Thus, Varga and Schmitt [ 20] distinguish between five major patterns: (i) endemics of the Alps restricted to some parts or the entire range, (ii) the "Alpine archipelago" with the species' strongholds in the Alps and other parts of the range in adjoining mountain ranges, but without long distance disjunctions, (iii) mountain species with long distance disjunctions in Europe (e.g. in the mountains of the Balkan Peninsula, Italy or Iberia), (iv) oro-Mediterranean species with their ranges mostly restricted to the mountains of the summer-arid high mountain systems of southern Iberia, southern Italy and the southern Balkan Peninsula and (v) arctic-alpine species with a disjunction between mountain populations in the South (Alps, often also Pyrenees, Carpathians, Balkan high mountain systems) and a large zonal range in the Arctic.

However, the distribution histories and evolutionary processes underlying these different distributional types in mountain species are still poorly understood. Therefore, I summarise the known examples and extract the repetitive pattern. Examples are preferably taken from the butterflies because this group is well studied and suitable for biogeographical analyses [e.g. [ 21- 29]], but where necessary other animal groups and even plants are referenced. In this article, I especially focus on two questions:

(1) Which repetitive biogeographical and evolutionary patterns are characteristic for European mountain species?

(2) What is the influence of the climatic cycles on their differentiation?

Endemics and endemic genetic lineages of high mountain systems

Endemics of the Alps

Most of the larger European high mountain systems possess endemic species (or at least subspecies) confined to just one of them. As the Alps are by far the most extensive and highest mountain system of Europe, it is not surprising that the highest number and amount of endemics of typical high mountain species is found in this mountain range, as for example demonstrated by lepidopterans and especially in the little mobile group of the micro-moths [ 20]. Some of these alpine endemics (in animals as well as in plants) are widespread all over this mountain area, stretching almost the entire range from Nice in the southwest to Vienna in the northeast (in many cases showing remarkable genetic substructure, see below); while others are more or less local endemics of some parts of the Alps.

Two types of hotspots for local endemics of the Alps can be distinguished:

(i) Species with their ranges restricted to peripheral regions of the Alps and mostly confined to lower and intermediate elevations (e.g. observed in some Erebia species, Lycaenids etc.; for details see Varga & Schmitt [ 20]). The largest concentrations of these local endemics are in the regions of the southwestern and the southeastern Alps, two regions with larger areas at lower altitudes not covered by ice during glaciations. These ice-free areas most probably served as centres of glacial survival, from where these species only performed altitudinal shifts, but no major range expansions over large parts of the deglaciating Alps. As these taxa might have performed these altitudinal shifts repeatedly through the climatic fluctuations of the Pleistocene, these refuge areas might also be the locations of the evolutionary process of speciation [ 1]. However, genetic analyses in this group of species are scarce and in butterflies restricted to the three taxa Coenonympha darwiniana [ 30], C. macromma [ 30] and Erebia sudetica inalpina (Figure ) [ 31]. The latter is today confined to the region of Grindelwald (Berner Oberland, northwestern Alps) and most probably survived at least the last glaciation in a geographically restricted refuge area northwest of the Alpine ice-shield, likely suffering population bottlenecks and subsequent genetic erosion [ 31]. The two Coenonympha taxa are restricted to the south-central Alps ( C. darwiniana) and the Alpes Maritimes ( C. macromma). Both might have survived also at least the last ice age at lower elevations close to their recent distributions [ 30].

(ii) Species restricted to some parts of the Inner Alps, as for example a number of (often flightless) micro-moth species [ 32] and beetles [ 5], but only a single butterfly species (i.e. Erebia nivalis) [ 20]. These species in general are confined to high alpine habitats and may have survived glaciations in situ at so called Nunataks (i.e. small ice-free areas topping over the ice-shield of the Inner Alps). For animals, survival in these areas is only supported by chorological data sets (i.e. distribution pattern) [ 5, 20], but genetic data in plants give some support for these refuges (see Schönswetter et al. [ 33] for a recent review, [ 34, 35]).

Endemics of other high mountain systems of Europe

Although the Alps are the European mountain system with the highest number of endemic species, numerous endemic genetic lineages or taxa are known for almost all high mountain systems in parts of Europe not entirely being covered by ice during the glacial periods. However, there are peculiar differences between the more marginal (e.g. Cantabrian Mts., Bulgarian Mts.) and more central (e.g. Carpathians, Massif Central, Apennines) mountain systems.

Thus, the more marginal mountain areas of Europe have particularly high proportions of old evolutionary units (often with relict status) when compared to more geographically central mountain ranges. For example, cases of such old genetic lineages are known for the flora of the Cantabrian Mts. in northwestern Spain [e.g. [ 36- 38]]; the number of examples in animals is rather low, but the known cases support the patterns observed in plants well, e.g. the butterfly Proclossiana eunomia [ 39, 40] and the caddisfly Drusus discolor (Figure ) [ 41].

Another stronghold of strongly differentiated genetic lineages and endemic species is situated in the Bulgarian high mountain systems. Thus, the most distinct genetic lineages of the beetle Nebria rufescens and the butterfly Erebia pandrose, two arctic-alpine species, were found here [ 42]; a strongly differentiated genetic lineage was also found in the Proclossiana eunomia populations from the Stara Planina [ 39]. In the spiders of the Pardosa saltuaria species group, the populations from the Bulgarian mountains, and also from the Pyrenees, showed a strong differentiation from a northern clade including the Alps, Carpathians, some middle high mountains of Central Europe and Scandinavia (Figure ) [ 43]. The caddisfly Drusus discolor also showed strongly differentiated genetic lineages for all of the marginal mountain systems of Europe where the taxon occurs, including the high mountain systems of Bulgaria (Figure ) [ 41]. Also plants in the Balkan high mountain systems showed strongly differentiated genetic lineages from other regions of Europe [ 44].

Furthermore, the distribution pattern of some species and subspecies strongly support these observed genetic structures. Thus, the eastern Balkan mountains are the only mountain range of the southern part of Europe where the fairly widespread mountain ringlet Erebia epiphron is lacking, but the species is substituted by its sister taxon Erebia orientalis supporting the strong evolutionary independence of this region. Regarding this latter species, each of the three mountain ranges harbouring it (i.e. Rila, Pirin, Stara Planina) has its own subspecies with strong morphological differentiations among them [ 45, 46]. A similar situation is also observed for the only occurrences of Euphydryas cynthia outside the Alps: the male individuals of the Pirin population consistently have a considerably more extended white wing colouration than average populations from the Alps, while the white colouration is strongly reduced in the Rila individuals [ 45]. This also underlines the evolutionary independence among these mountain ranges of the eastern Balkan Peninsula on a fairly small geographic scale of sometimes only few tens of kilometres.

Such old lineages are fewer in the mountain systems with closer geographic connections with the Alps (e.g. Carpathians, Massif Central), and genetic differentiations of these mountain systems from the Alps are often low; most examples come from plants [e.g. [ 36, 38, 47, 48]], but also include the Pardosa saltuaria spider species group (Figure ) [ 43]. On the species level, these mountain ranges mostly share their species with the Alps. However, well distinguished subspecies are known for these mountain species with many examples in the genus Erebia (e.g. E. epiphron, E. euryale, E. gorge, E. manto, E. melas) [ 49], but the ages of these splits are mostly unknown and some even might be rather recent, even postglacial.

The Pyrenees have an intermediate position, in which they show some close connections to the Alps (see below), but also have numerous endemic species (e.g. in lepidopterans Erebia gorgone, three Sattleria species) [ 20, 49]; their number however is much lower than in the Alps, which are considerably higher and much more extended. Many mountain species with larger distributions show endemic lineages in the Pyrenees or parts of them, thereby supporting the idea of differentiation centres at the lower elevations of the Pyrenean area and recent (mostly postglacial) range shifts to higher altitudes of the mountains, as demonstrated for the butterflies Erebia epiphron (Figure ) [ 50], Erebia euryale (Figure ) [ 51], Erebia pandrose [ 42] and Erebia manto (unpublished data), but also other animal [ 41, 43] and plant species [ 36].

The summer-arid high mountain areas of the extreme south of Europe (especially southern Iberia and the southern Balkan Peninsula) and the Maghreb (here in particular the Atlas Mts.) do not have the typical alpine zonation [ 52], but show typical oro-Mediterranean characteristics. Some mountain species show local endemism to strongly confined mountain blocks in this area as e.g. known for a number of butterfly species [ 20, 45, 49]. These species in many cases represent relicts of often old invasions of species with their closed biogeographical connections to Central Asia [ 20, 53, 54].

Different genetic lineages within one high mountain system

The large high mountain systems of the southern part of Europe (i.e. Pyrenees, Alps and Carpathians) have in many cases been colonised after the postglacial deglaciation by several genetic lineages, which have survived at least the last glacial period in several allopatric refugial areas often adjoining these high mountain systems.

Different genetic lineages in the Alps

The Alps as the largest European high mountain system show a remarkable phylogeographic structure in many species. In plants, a differentiation into four genetic units along the Alps seems the most common feature with these lineages typically being confined to the southwestern, western central, eastern central and eastern Alps and the eastern lineages more often being of higher genetic diversity than the more western ones [ 2, 33, 42]. A quite similar pattern to that in many plant species was proposed for the leaf beetle Oreina elongata [ 55]. Also other animals frequently show different genetic lineages in the Alps, but their number is often less than in plants, and in many cases, only a western and an eastern lineage can be distinguished. This is the case in the two butterflies Erebia melampus (Figure ) [ 31] and Erebia euryale (Figure ) [ 51], the wolf spiders species group Pardosa saltuaria (Figure ) [ 43] and the caddisfly Drusus discolor (Figure ) [ 41]. In the first case, two strongly differentiated lineages (supported by allozyme polymorphisms [ 31] and differences in the male genitalia [ 56]) can be distinguished with a contact zone running north-south through Tyrol and along the Adige/Etsch valley (maybe with some hybridisation in a limited area of southeastern Switzerland and South Tyrol). The differentiation between these two lineages is rather advanced, and they most probably represent different species, with the eastern lineage showing a significantly higher genetic diversity than the western one [ 31]. These genetic and morphological differences are best explained by allopatric differentiation in two centres; and as the beginning of the first split is assumed to be older than the last ice age, the location of the initial differentiation centres must remain enigmatic. However, the most likely Würm glacial refuge centres are in the hilly areas south of the Upper Italian Lakes for the western lineage and in the lower elevation areas of the southeastern Alps (eastern Carinthia, Styria, Friul, parts of Slovenia) for the eastern lineage. Based on the higher genetic diversities of the populations belonging to the eastern lineage, we may assume that the survival conditions were more suitable in this eastern region and/or this refuge area was geographically considerably more extended than the more western refuge [ 31]. A very similar pattern was observed for Erebia euryale (Figure ) with the only difference that survival of the western lineage is more likely around the southwestern Alps [ 51]. The caddisfly Drusus discolor repeats this geographic structure of a western and an eastern Alpine lineage (Figure ) [ 41]. A much more complicated case is known in the butterfly Erebia epiphron [ 50]. This species comprises four strongly differentiated lineages in the Alps: western Alps, Aosta valley, northern Alps and eastern Alps (Figure ). These lineages most probably evolved in four differentiation centres west, south, southeast and north of the Alps, often linking the Alps with other mountain systems (see below).

Different genetic lineages in the Pyrenees

As in the Alps, more than one genetic lineage per species is also observed in mountain elements of the Pyrenees [ 20]. However, the Pyrenees are considerably less extended than the Alps, and the maximum number of different lineages found here is not as high as in the Alps; many species do not show marked differentiation over the Pyrenees. As an example, two different lineages are observed in the Pyrenees for Erebia epiphron (Figure ), one in the western and the other in the eastern parts of these mountains, with the first one most probably surviving the last ice age north and the second in the southeastern parts of the Pyrenees [ 50]. Also for the plant Cardamine alpina, AFLP patterns suggest survival in multiple Pyrenean refugia [ 57].

Different genetic lineages in the Carpathians

Although being the largest high mountain system of eastern/southeastern Europe, only a few genetic analyses include populations from more than one part of the Carpathian arc. However, compiling existing data for different plant species revealed a generally lower genetic diversity of the populations in the Carpathians than in the Alps, maybe due to higher topographic isolation of alpine habitats in the Carpathians [ 58].

One study on animals that included a considerable number of samples from the Carpathian region was performed on the caddisfly Drusus discolor and showed a remarkable differentiation between the northwestern and the southern Carpathians (Figure ). This thus supports the idea of a long lasting separation between these two parts of this high mountain system predating at least the last glaciation [ 41]. Similar cases are also reported for plants as Hypochaeris uniflora [ 59] and Campanula alpina [ 60].

Thus, multiple glacial survival centres exist along the Carpathian arc, as in the cases of the Pyrenees and especially Alps, but the number of known cases is still rather limited. Therefore, a research focus on the phylogeography of the Carpathians is needed.

Genetic links between different high mountain systems

As many of the glacial differentiation centres were not restricted to the foot-hills of the respective high mountain systems, but were located in the (hilly) areas between high mountain systems, many links (by species and genetic lineages) can be observed between adjoining high mountain systems, especially between (i) Pyrenees and southwestern Alps, (ii) northeastern Alps and northwestern Carpathians, (iii) southeastern Alps and western Balkan mountains and (iv) southern Carpathians and the eastern Balkan high mountain systems [ 20]. In the following, these connections are discussed using examples.

Links between Pyrenees and southwestern Alps

The biogeographical links between the Pyrenees and the Alps are seen in the occurrences of many taxa with restricted distributions in these two areas, e.g. in lepidopterans by taxa as Euchloe simplonia, Polyommatus g. glandon, Aricia n. nicias, Erebia o. oeme [ 49]. However, the same genetic lineages are also repeatedly found in these two regions, as in the case of Erebia cassioides [ 61] and Erebia epiphron (Figure ), which has genetically very similar populations in the western Pyrenees and the southwestern Alps, e.g. the Wallis in Switzerland [ 50]. The genetic similarity in this case is best explained by a Würm glacial link between the Pyrenees and the Alps, and thus a more widespread distribution in the hilly regions of France between these two high mountain systems during this time period. The postglacial disjunction has not been sufficiently long for the evolution of a clear differentiation between these now disjunct parts of the distribution. Similar patterns are also known for mountain plant species [ 47, 62- 66]. In the case of Carex curvula, the Pyrenean populations are nested within the western Alps group and show a low level of genetic diversity, probably due to recent long distance colonisation [ 67].

Links between northeastern Alps and northwestern Carpathians

As in the case of the Pyrenees-Alps connections, many biogeographical interactions exist between the northeastern Alps and the northwestern Carpathians (i.e. Tatra Mts.). Typical examples are found in distribution patterns of lepidopterans like Pieris bryoniae and Erebia pharte [ 49]. An mtDNA analysis of the wolf spider species complex Pardosa saltuaria also underlines this link as the same mtDNA lineage was detected in the Alps and Carpathians (Figure ) [ 43].

So far other genetic data for animals are lacking, but data on plants support this biogeographical link (e.g. for Pritzelago alpina [ 36] and Senecio carniolicus [ 48]). For Ranunculus alpestris, the Carpathians have been colonised stepwise from an eastern Alpine lineage [ 68], and the arctic-alpine Ranunculus pygmaeus colonised the Alps relatively recently from the east through the Tatra Mts. [ 69]. A relatively similar genetic link was observed between the northern Alps and the mountains of northern Moravia about 200 km west of the High Tatra Mts. for the butterfly Erebia epiphron (Figure ) [ 50].

Nevertheless, many examples also revealed long-lasting separations between the populations from the Alps and the Carpathians [ 57, 60, 65, 70].

Links between southeastern Alps and western Balkan mountains

The southeastern edge of the Alps is biogeographically strongly linked with the mountains of the western Balkan Peninsula, as demonstrated by many occurrences of identical mountain beetle species in the southeastern Alps and the northwestern Balkan mountains [ 5, 20]. While less in number, this distribution type is also known in the Lepidoptera for taxa like Coenonympha gardetta, Erebia ottomana, E. stirius, E. oeme spodia, E. epiphron aetheria and E. styx trentae [ 49]. These distribution patterns support the survival of mountain species in the southeastern parts of the Alps and adjoining areas (as demonstrated for Erebia melampus (Figure ) [ 31] and Erebia euryale (Figure ) [ 51]), with postglacial retreat into the Alps in the northwest and the northwestern Balkan mountains in the southeast.

Genetic data sets supporting these biogeographical links for animals are (to my knowledge) not available at the moment, but genetic data on plants support these connections e.g. in the species Pulsatilla vernalis [ 65] and Polygonatum verticillatum [ 38]. Considering morphological studies, the western Balkan populations of Erebia pandrose in the high mountain systems of Monte Negro, Serbia and Macedonia are closely related with populations existing in the southeastern Alps. This thus supports a recent (i.e. Würm glacial) biogeographical link between these two areas that are now separated by several hundreds of kilometres with no mountains over 2,000 m. On the contrary, the Erebia pandrose populations of the high mountain systems Rila and Pirin in Bulgaria are strongly differentiated from this group, and more closely related with the populations from the southern Carpathians [ 71], thus underlining a biogeographical east-west split throughout the central Balkan Peninsula [ 72].

Links between southern Carpathians and eastern Balkan mountains

The Danube valley apparently has often acted as effective dispersal barrier between the Carpathians and the high mountain systems of the eastern Balkan Peninsula [ 20] with Erebia medusa being a well studied case in the Lepidoptera [ 18]. However, many connections between the mountain systems of these two areas are also known. Thus, a morphometric study of the male genitalia of Erebia pandrose demonstrated high similarities between these two high mountain areas [ 71] and hereby also supported a close biogeographical link between them.

An allozyme analysis of the mountain forest species Erebia euryale revealed by far the highest genetic diversity in this species in the populations from the Bulgarian Rila and the Romanian southern Carpathians, but no significant differentiation was detectable between these two areas (Figure ) [ 51]. This is most probably due to the fact that these two areas were linked by cool forested areas during at least the last ice age allowing massive gene flow between these two regions; thus, at least the region of the Iron Gate (passage of the Danube through the southern Carpathians) must have been extensively covered by this type of forest during this time period. Furthermore, this study supports the conclusion of other analyses based on fossil remains (e.g. pollen, remains of animals) [ 73- 78] that southeastern Europe was the most important glacial retreat of the representatives of the European mountain forest biome.

Furthermore, the distribution of many mountain species (e.g. Erebia melas, E. cassioides, Coenonympha rhodopensis) in the Bulgarian high mountain systems and the southern Carpathians underlines the biogeographical connections of the high mountain biota of both areas [ 20]. Genetic links between both areas are also known for plant species [ 67].

Genetic links between high mountain systems and the Arctic

Recent (i.e. late glacial or postglacial) genetic links between the high mountain systems in the South and the Arctic in the North are a rather common biogeographical characteristic of many species with typical arctic-alpine disjunctions as already postulated by Holdhaus in his landmark book in 1954 [ 5]. This proposal is largely supported by many genetic analyses on plants [ 65, 66, 70, 79- 83] and animals (Figure ) [ 42, 43]. This argues for the wide distribution of these species in the periglacial steppes between the northern glacier and the glaciated mountains in the South and postglacial retreat to higher altitudes in the South, and higher latitudes in the North.

In butterflies, the mtDNA sequences of Erebia pandrose indicate a very close genetic connection between the populations from the Alps, Pyrenees and Scandinavia thus supporting their recent (i.e. postglacial) disjunction. However, the individuals from the Rila in Bulgaria show a stronger genetic differentiation from these other regions, thus supporting the idea that the Balkan populations were not included in this large zonal distribution during the last glaciation, but survived in an area of southeastern Europe not linked to the main distribution [ 42]. A similar structure was also observed in the leaf beetle Nebria rufescens, but the differences against the Balkan haplotypes were even stronger in this case [ 42]. Another butterfly study with the lycaenid Aricia artaxerxes revealed a very weak genetic differentiation between the populations from the northern UK and Scandinavia of only one or two mutations in the mtDNA cytochrome-b gene thus supporting the idea of a common source and only postglacial isolation [ 84]. A rather similar genetic structure was observed for the diving beetle Hydroporus glabriusculus [ 85]. The mountain hare Lepus timidus showed genetically very similar populations in the Alps and in Fennoscandia analysed by allozymes [ 86] microsatellites and mtDNA [ 87] underlining a postglacial common source for both regions between the Alpine and the Fennoscandian ice-shields. The lack of genetic differentiation between the Alps and Pyrenees in the arctic-alpine burnet Zygaena exulans also supports the idea of a survival of this species in a large zonal distribution range in the periglacial area of Europe during the last ice age and postglacial retraction to the high mountain systems and the Arctic [ 88].

Genetic connections of middle high mountain systems

The middle high mountain systems of Europe show numerous biogeographical connections with the high mountain systems and among each other. Thus, the French Massif Central shows strong connections with the Pyrenees in many cases, e.g. by having the same melanistic subspecies Erebia manto constans [ 49] as well as showing rather weak genetic differentiation in Parnassius apollo [ 89] and Pulsatilla vernalis [ 65]. However, in other cases, close connections exist between the Massif Central and the Alps, as in the plant Polygonatum verticillatum [ 38]. However, the populations of the Massif Central are detected as even an independent phylogeographic group, as in the case of the caddisfly Drusus discolor (Figure ) [ 41].

Links with the Alps are known for the Bavarian and Bohemian Forest, e.g. by sharing the eastern Alpine lineage in the caddisfly Drusus discolor (Figure ) [ 41], and for the Jesenik mountains in northern Moravia, e.g. by striking genetic similarities with northern Alps E. epiphron populations (Figure ) [ 50]. Relicts of alpine species in the Vosges and (to a lesser degree) in the Black Forest (e.g. in butterflies Erebia epiphron, E. manto, Clossiana titania, C. thore) [ 49] underline their close biogeographical relation with the Alps, which is also supported by genetic studies e.g. in the plant Polygonatum verticillatum [ 38]. In Pulsatilla vernalis, Tatra and Sudeten Mts. share the same chloroplast haplotype indicating a close link between these two mountain ranges for this species, but isolation from the Alps [ 65]. These patterns are most likely due to glacial refuge areas in the vicinity of the Alps, Carpathians or Pyrenees and postglacial retreat into one of these high mountain systems, but also into at least one of the adjoining middle high mountain systems with not sufficient time for differentiation since then.

Close biogeographical connections were also revealed among these middle high mountain systems. One striking example is the caddisfly Drusus discolor showing the same mtDNA lineage in the COI gene in middle high mountain systems (Figure ) [ 41]. A similar pattern was also obtained in a morphometric study of another caddisfly species Rhyacophila aquitanica with very similar populations in the French Massif Central, the Vosges, the Black Forest and the Swiss canton of Fribourg (Figure ) [ 90]. These patterns most probably result from a glacial distribution in the lower altitudes between these middle high mountain areas and postglacial retreat into the higher altitudes of these adjoining mountains.

Conclusion

The topology of the European mountain systems had a strong influence on the biogeography and evolution of mountain and high latitude species. Due to the dynamic changes of these biota over time, a large variety of different evolutionary processes are responsible for the development of numerous biogeographical patterns. The most simple of these is the evolution of one genetic lineage or species endemic to one single mountain system due to vicariance with subsequent long processes of geographic isolation and evolution, as observed in almost all European high mountain systems. Especially in the Alps, but also in smaller high mountain systems, different genetic lineages exist beside each other as a result of evolution in geographically isolated areas, with often secondary contacts in these mountain systems. Where such centres of evolution and differentiation of lineages were located between two mountain systems, retraction during the postglacial into both of them is commonly resulting in one genetic lineage being found in two different mountain systems.

Competing interests

The author declares that he has no competing interests.

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# data[].attributes.attachment_link.title
Biogeographical and evolutionary importance of the European high mountain systems

# data[].attributes.attachment_link.description
Europe is characterised by several high mountain systems dominating major parts of its area, and these structures have strongly influenced the evolution of taxa. For species now restricted to these high mountain systems, characteristic biogeographical patterns of differentiation exist.