Download Mountain Biomes - Ecology - Oxford

Mountain Biomes
Christoph Kueffer
Introduction
Mountains are landforms that rise prominently above the surrounding landscape. They are topographic
features that are defined by a high relief, and their most general features are that they encompass a certain elevational range, have
steep slopes, and converge toward small summit areas. Depending on the definition, up to a quarter of the global land area can be
considered mountainous. Ecosystem services in mountains directly support about a quarter of the world population that lives in or
near mountains, and many more humans depend on mountains, in particular as a source of water and minerals, and as a tourist
destination. Many mountains have special cultural or spiritual significance. Mountains have attracted the interest of scientists since
ancient times and have been of great importance to modern geography, biogeography, evolutionary biology, and ecology since the
18th century. Mountains are characterized by high species and habitat diversity that result from high environmental heterogeneity
due to elevational gradients, exposure, orographic effects, and natural disturbances such as landslides, avalanches, and floods.
The islandlike isolation of many mountains contributes to unique and diverse mountain biotas. At the same time mountains occur
mostly as part of elongated ranges or chains that connect biotas across large geographic distances (e.g., the North and South
American Cordilleras). Mountain ecosystems are increasingly exposed to human pressures including land-use changes (e.g.,
agriculture, livestock farming, tourism, urbanization, or abandonment), pollution, and climate change (e.g., loss of glaciers, melting
of permafrost, desertification, increased frequency of natural hazards such as floods, landslides and avalanches, and spread of
diseases and invasive species).
General Overviews
The diversity of mountains is limitless. They are found in all climate zones and biogeographic regions, include along their
elevational gradients strongly contrasting vegetation zones, and they differ widely in their maximal elevation, geology, or human
land use. Therefore, for introductory information on general features of mountains it is necessary to also consult treaties of
particular vegetation zones (see Vegetation Zones) and geographic regions (see Mountain Regions of the World). Some of the
classical historic work gives excellent overviews of mountains (see Historical Accounts and Foundational Works). A recent synthetic
and evidence-based assessment of mountain ecosystems and their threat status is Körner, et al. 2005. An excellent and
comprehensive textbook is Price, et al. 2013; in German, Franz 1979 is an available work. An Ambio Special Report on mountains
includes, among others, two complementary overviews of biodiversity in mountains, Körner 2004 and Molau 2004. Körner and
Spehn 2002 includes a representative collection of regional accounts of mountain biodiversity.
Franz, H. 1979. Ökologie der Hochgebirge. Stuttgart: Ulmer.
A comprehensive compilation of information on all aspects of the ecology of high-elevation ecosystems—including animal ecology,
human biology, and aquatic ecosystems—rooted in natural history. In German. “Hochgebirge der Erde” by the same author (1989,
Urania-Verlag, Leipzig, Berlin) introduces the major mountain systems of the world.
Körner, C. 2004. Mountain biodiversity, its causes and function. Ambio 13:11–17.
This general overview of mountain biodiversity gives estimates of species numbers in mountains, and discusses why mountains are
highly biodiverse and in particular the importance of plant functional diversity for mountain ecosystems. It concludes with an outlook
on the impacts of land use and climate change on mountain vegetation.
Körner, C., M. Ohsawa, and E. Spehn. 2005. Mountain systems. In Ecosystems and human well-being: Current state and
trends. Findings of the conditions and trends working group of the Millennium Ecosystem Assessment. Edited by R.
Hassan, R. Scholes, and N. Ash, 681–716. Washington, DC: Island Press.
Chapter 24 on “mountain systems” of the Millennium Ecosystem Assessment assessed the available knowledge at the time on
physical, biological, economic, and social conditions in the world’s mountains. It defines mountains; gives an overview of the global
distribution of mountain areas; characterizes mountain climates, biotas, ecosystems and ecosystem services; and in particular
reviews human pressures on mountain biodiversity and ecosystems. Available online.
Körner, C., and E. M. Spehn, eds. 2002. Mountain biodiversity: A global assessment. London: Parthenon.
This edited volume is the outcome of an international conference organized in 2000 by the Global Mountain Biodiversity
Assessment (GMBA; see Online Databases and Web Pages). Apart from an almost exclusive focus on plant diversity, it covers
different aspects of mountain biodiversity: the main mountain regions of the world; vegetation zones from montane forest to alpine
vegetation; genetic, species, functional, and habitat diversity; and land use, climate change, and biodiversity conservation.
Molau, U. 2004. Mountain biodiversity patterns at low and high latitudes. Ambio 13:24–28.
This concept paper addresses mountain biodiversity at different spatial scales, considering both genetic and species diversity. In
particular it discusses how mountain biodiversity patterns differ between high latitudes and the tropics.
Price, M. F., A. C. Byers, D. A. Friend, T. Kohler, and L. W. Price, eds. 2013. Mountain geography: Physical and human
dimensions. Berkeley: Univ. of California Press.
This is a major revision of Mountains and Man (1981) by Larry Price and will likely become the general reference book on all
aspects of mountain biomes like its predecessor. The multiauthored book is of global scope and includes chapters on origins of
mountains, climate, geomorphology, soils, vegetation, wildlife, people, and sustainable development.
Historical Accounts and Foundational Works
Mountains have attracted the interests of scientists for centuries. Chapter 9 in Price, et al. 2013 (cited under General Overviews)
introduces early perceptions of mountains in different cultures. Mathieu 2011 recounts the history of mountain research and
perceptions since the 16th century. In particular, mountains played a central role in the development of early plant ecology and
biogeography including through the work of key figures such as Alexander von Humboldt. Since the 1970s a global mountain
research and management community has emerged (Messerli 2012, Debarbieux and Price 2008, see both under Rise of a Global
Mountain Research Community).
Mathieu, J. 2011. The third dimension: A comparative history of mountains in the modern era. Cambridge, UK: White
Horse.
A concise and easily accessible global history of mountains. The book has two main foci: the history of the scientific and cultural
perception of mountains (including Humboldt and the rise of a global research and policy community since the 1970s), and a
comparison of the socioeconomic development in different mountain regions of the world.
EARLY PLANT ECOLOGY AND BIOGEOGRAPHY
Particularly important for early modern mountain science was the work of Alexander von Humboldt (Buttimer 2012). Hart Merriam’s
work on life zones built on work along elevational gradients (Merriam and Steineger 1890) was central to the development of the
concept of biomes. Schroeter 1926 and Daubenmire 1943 illustrate the rich understanding of alpine ecology by early plant
ecologists. The gradient analysis work of Whittaker challenged a static view of vegetation zones (Whittaker 1956). Elevational
gradient studies were also important for an early understanding of population differentiation into different ecotypes (Nunez-Farfan
and Schlichting 2001). For a short overview of early plant ecological work in mountains see also Körner 2003 (cited under Alpine
Vegetation) and Holtmeier 2009 (cited under Treelines).
Buttimer, Anne. 2012. Alexander von Humboldt and planet earth’s green mantle. Cybergeo: European Journal of
Geography.
Introduces Alexander von Humboldt’s work with a special focus on its relevance for mountain research and puts it in a historical
context. The open-access article also includes a complete English translation of Humboldt’s “An Essay on the Geography of
Plants.”
Daubenmire, R. F. 1943. Vegetational zonation in the Rocky Mountains. Botanical Review 9.6: 325–393.
A classical account of vegetation zonation in mountains.
Merriam, C. H., and L. Steineger. 1890. Results of a biological survey of the San Francisco mountain region and the desert
of the Little Colorado, Arizona. North American Fauna Report 3. Washington, DC: Department of Agriculture, Division of
Ornithology and Mammalia.
Hart Merriam’s concept of life zones based on the relationship between climate and vegetation was central to the development of
the concept of the biome. His work extensively builds on research along elevational gradients as illustrated by this report. Influential
was his idea to classify elevational vegetation zones in correspondence to latitudinal vegetation zones (compare Holdridge 1967,
Vegetation Zones). Available online.
Nunez-Farfan, J., and C. Schlichting. 2001. Evolution in changing environments: The “synthetic” work of Clausen, Keck,
and Hiesey. Quarterly Review of Biology 76:433–457.
In the first half of the 20th century, Clausen, Keck, and Hiesey pioneered the study of population differentiation into ecotypes
through observational and experimental studies along elevational gradients. The same research approaches are still widely used in
plant ecology.
Schroeter, C. 1926. Das Pflanzenleben der Alpen. Eine Schilderung der Hochgebirgsflora. Zürich, Switzerland: Verlag von
Albert Raustein.
An exemplary book illustrating the rich knowledge of early plant ecologists of plant life and ecology of mountain ecosystems. Such
historical publications remain a valuable source of information but unfortunately are often not written in English and difficult to find.
Whittaker, R. H. 1956. Vegetation of the Great Smoky Mountains. Ecological Monographs 26:1–69.
Whittaker’s gradient analysis of species co-occurrence along environmental gradients challenged the view of vegetation
communities as coherent entities and represents some of the most influential work in ecology and biogeography. Whittaker’s
theoretical work was based on fieldwork in mountains including the Great Smoky Mountains.
THE RISE OF A GLOBAL MOUNTAIN RESEARCH COMMUNITY
Since the 1970s a global mountain research and management community has emerged (Messerli 2012, Debarbieux and Price
2008). In 1968, the International Geographical Union’s Commission on high-altitude geoecology was founded thanks to the initiative
of Carl Troll. A special issue published in 1973 in Arctic, Antarctic, and Alpine Research (cited under Journals) illustrates efforts to
standardize definitions and promote global-scale comparative studies on mountain ecology, including on treelines (Troll 1973). In
the 1970s UNESCO’s Man and Biosphere (MaB) program on mountains became an important stimulus of mountain research (Price
1995). In the same decade a theory of Himalayan degradation became prominent among international policymakers that stated that
poverty and overpopulation in the Himalayas caused degradation of highland forests, erosion, and flooding in downstream regions
such as Bangladesh. These debates were a testing ground for the capacity of the international mountain research to assist
policymaking, with leading mountain scientists challenging the scientific validity of the prevailing thinking (Ives 2004). Since the
early 1990s, the mountain research community has successfully used the UNCED Earth Summit process to promote mountains as
a global environmental issue. In preparation of the first summit in Rio de Janeiro (in 1992), a global assessment was produced by a
small expert group known as “mountain agenda” (Stone 1992). The report contributed substantially to the formulation of the
mountain-related Chapter 13 of the Agenda 21, which led to a more comprehensive global assessment of mountain issues
(Messerli and Ives 1997). Körner, et al. 2005 (cited under General Overviews) is a global expert-based assessment of environment
issues related to mountains.
Debarbieux, B., and M. F. Price. 2008. Representing mountains: From local and national to global common good.
Geopolitics 13:148–168.
An analysis of the emergence of a global awareness of environmental issues in mountains from a theoretical stance.
Ives, J. 2004. Himalayan perceptions: Environmental change and the well-being of mountain peoples. Himalayan Journal
of Sciences 2:17–19.
In 1989 a book entitled The Himalayan Dilemma by Ives and Messerli (New York: Routledge) questioned the scientific basis of
popular thinking among policymakers about environmental degradation in the Himalayan region. In 2004 Ives published an updated
assessment of the issue entitled Himalayan Perceptions (New York: Routledge). This article (freely available online) summarizes
the main conclusions of the two books and puts the debate into a broader perspective.
Messerli, B. 2012. Global change and the world’s mountains: Where are we coming from, and where are we going to?
Mountain Research and Development 32:S55–S63.
A personal account of the rise of a global mountain research community since the 1970s written by one of the involved key
persons. The article is open access.
Messerli, B., and J. D. Ives, eds. 1997. Mountains of the world: A global priority. New York: Parthenon.
This multiauthored global assessment of environmental issue is a result of Chapter 13 of Agenda 21, and in contrast to Stone 1992
it is not structured by region but by topics that are assessed across regions. In the first two parts the cultural, economic, and
political dimensions of mountain development and environmental impacts are assessed. The third part sketches an agenda for
sustainable development in mountains.
Price, M. F. 1995. Mountain research in Europe: An overview of MAB research from the Pyrenees to Siberia. Man and the
Biosphere series 14. New York: Parthenon.
Reviews mountain research undertaken within the MAB framework in Europe (France to Russia). The European MAB program had
a strong influence on mountain research in Europe and beyond, especially on interdisciplinary research addressing the interplay of
mountain ecology and land use.
Stone, P. B., ed. 1992. The state of the world’s mountains: A global report. London: Zed Books.
A global and expert-based assessment of mountain ecology and sustainability. The book includes regional case study chapters
covering African mountains, European Alps, Himalaya, Andes, former Soviet Union, Appalachians, and shorter treatments of
mountains that were considered neglected at the time: islands, arctic mountain systems, New Guinea, northwestern Thailand,
Yunnan, and Sierra Nevada (United States).
Troll, C. 1973. The upper timberlines in different climatic zones. Arctic and Alpine Research A3–A18.
Carl Troll was a pioneer of global comparative studies of alpine ecology, including treelines. Troll initiated in 1968 the International
Geographical Union’s Commission on high-altitude geoecology, which was important for initiating the networking and enabling of a
global mountain research community.
Journals
Research on mountains is published in a wide range of journals including those in ecology, conservation, and geography; and
journals specialized in arctic and polar sciences (see entries on Polar Regions and Tundra Biome in Oxford Bibliographies in
Ecology). There are a number of scientific journals specialized in mountain research. A broad interdisciplinary scope have Arctic,
Antarctic, and Alpine Research and Mountain Research and Development, and—with a main focus on the social sciences:
—Journal of Alpine Research. Plant Ecology and Diversity and Alpine Botany focus on plants. Eco.mont publishes work from
mountain protected areas. Journal of Mountain Science is a journal published by the Chinese Academy of Sciences. There are also
specialized mountain science journals with a regional focus that are not listed among the periodicals below: Sustainable
Development of Mountain Territories (Russia), Oecologia Montana (Eastern Europe), Pirineos (Spain), Himalayan Journal of
Sciences, and different institutions produce high-quality newsletters (see Online Databases and Web Pages).
Alpine Botany.
Since its new launch in 2011, the journal of the Swiss Botanical Society (formerly Botanica Helvetica) publishes botanical studies
with a particular interest in plant ecology, vegetation, and flora of mountain regions. Listed in Web of Science.
Arctic, Antarctic, and Alpine Research.
An interdisciplinary, quarterly, peer-reviewed journal founded in 1969; until 1998 it was published as Arctic and Alpine Research.
Listed in Web of Science.
Eco.mont.
Publishes peer-reviewed articles on research within protected mountain areas or of interest to protected area management. Listed
in Web of Science.
Journal of Mountain Science.
An international, peer-reviewed, English-language mountain sciences journal started in 2004 by the Chinese Academy of Sciences
with the support of the United Nations University.
Mountain Research and Development.
An interdisciplinary and policy-oriented, peer-reviewed, open-access journal founded in 1981. Main focus is sustainable
development in mountains. Listed in Web of Science.
Plant Ecology and Diversity.
The journal of the Botanical Society of Scotland publishes peer-reviewed articles on all aspects of plant ecology but particularly
welcomes submissions concerning cold environments. Listed in Web of Science.
Revue de Géographie Alpine/Journal of Alpine Research.
A human geography journal that publishes articles on spatial and environmental issues in mountains from the social sciences
(geography, anthropology, sociology, history, political science, etc.). Articles are published in two languages (French, Italian,
Spanish, or German, and English). Published since 1913 by the Institute de Geographie Alpine in Grenoble, France. Listed in Web
of Science.
Online Databases and Web Pages
Thanks to a well-connected mountain research community, there are a number of excellent information gateways available online.
Mountain ecoregions of high- conservation priority are documented on the WWF Ecoregions and Biodiversity Hotspot webpages.
Data on mountain biodiversity can be found on the GMBA Mountain Biodiversity Portal, GLORIA collects floristic data from
mountaintops worldwide. Global expert networks are the Global Mountain Biodiversity Assessment (GMBA), Mountain Research
Initiative (MRI), Mountain Forum, and Mountain Partnership. Institutions with a regional focus include ICIMOD (Hindu KushHimalaya-Tibet), CONDESAN (Andes), CIRMOUNT (western North America), and EUROMONTANA (Europe).
Biodiversity Hotspots.
The biodiversity hotspot concept of Conservation International identifies world regions with a particularly important representation of
world biodiversity, which include many mountain regions. These biodiversity hotspot areas are portrayed on the web page and also
in a coffee table book. See Mittermeier, Russell. Hotspots Revisited (Mexico City: Cemex, 2004).
CIRMOUNT.
The Consortium for Integrated Climate Research in Western Mountains (CIRMOUNT) is an interdisciplinary consortium dedicated
to understanding climates and ecosystems of western North American mountains. It produces the newsletter Mountain Views.
CONDESAN.
A network and information portal for the Andes that runs different monitoring and research activities: ranging from vegetation to
socioeconomic aspects. In Spanish.
EUROMONTANA.
Founded in 1974, this is an early example of a mountain network. Originally focused on mountain agriculture, it is today a
thematically broad network with nearly seventy member organizations from fifteen European countries. The web page includes a
database of upcoming events.
Global Mountain Biodiversity Assessment (GMBA).
The Global Mountain Biodiversity Assessment (GMBA) is a cross-cutting network of DIVERSITAS with the task of exploring and
explaining the biological richness of the mountains of the world. Among others GMBA promotes a global mountain LTER network
(with information of existing ones included on the web page) and maintains the mountain biodiversity portal.
GLORIA.
The Global Observation Research Initiative in Alpine Environments (GLORIA) is a research network that monitors vegetation
change at mountaintops in over a hundred regions worldwide.
GMBA mountain biodiversity portal.
The mountain biodiversity portal allows searching GBIF’s species distribution data for particular mountain regions or vegetation
belts (e.g., alpine).
ICIMOD.
The International Centre for Integrated Mountain Development (ICIMOD) is a regional learning and knowledge-sharing center
based in Kathmandu (Nepal) serving the eight regional member countries of the Hindu Kush Himalayas: Afghanistan, Bangladesh,
Bhutan, China, India, Myanmar, Nepal, and Pakistan. A rich and up-to-date information source about the region.
Mountain Forum.
The Mountain Forum is a knowledge repository, social network (with thematic forums), and information portal for people interested
in sustainable mountain development around the world.
Mountain Partnership.
The Mountain Partnership is a United Nations voluntary alliance dedicated to improving the lives of mountain people and protecting
mountain environments around the world. Members are governments, intergovernmental organizations, and interest groups (e.g.,
civil society, NGOs, and the private sector). Includes an events database.
Mountain Research Initiative (MRI).
MRI promotes and coordinates global change research in mountain regions all over the world, and among others runs regional
networks for Africa, the Americas, Europe, the Carpathians, and southeastern Europe.
WWF Ecoregions.
The web page includes descriptions of many unique mountain ecosystems around the world.
Defining the Mountain Biome
Everyone has an intuitive understanding of what a mountain is, but how to define mountains remains a contested issue. Mountains
are landforms that rise prominently above the surrounding landscape. They have a high relief (steep slope at a local scale), are
elevated, and converge toward a summit. There are, however, different measures of local relief, and opinions differ about whether a
minimum threshold altitude should be defined for mountains. Depending on definition, high-altitude plateaus such as the Tibetan
Plateau are sometimes considered mountainous and sometimes not, and equally small topographic features such as inselbergs
Plateau are sometimes considered mountainous and sometimes not, and equally small topographic features such as inselbergs
might be included in the definition. Importantly, mountain environments are not synonymous with alpine environments that are
above the tree line, and it must also be kept in mind that many mountains in the tropics are exposed to a warm climate. Mountain
definitions also vary, depending on cultural views. Depending on the definition, between approximately 10 percent and 25 percent
of global land area can be considered mountainous. Chapter 1 in Price, et al. 2013 (cited under General Overviews) nicely
summarizes different perspectives on defining mountains, and Mathieu 2011 (cited under Historical Accounts and Foundational
Works) provides a more detailed historical perspective. Kapos, et al. 2000 introduces a widely used definition of mountains at a
global scale, and Körner, et al. 2011 reports on a recent adaptation of it.
Kapos, V., J. Rhind, M. Edwards, M. F. Price, and C. Ravilious. 2000. Developing a map of the world’s mountain forests. In
Forests in sustainable mountain development: A State of knowledge report for 2000. Edited by M. F. Price and N. Butt, 4–
9. Wallingford, UK: CABI.
Widely used and cited definition and global map of mountains based on a digital elevation model (DEM) with a resolution of thirty
arc seconds (approximately 1 km). The definition considers both elevation and slope.
Körner, C., J. Paulsen, and E. M. Spehn. 2011. A definition of mountains and their bioclimatic belts for global comparisons
of biodiversity data. Alpine Botany 121:73–78.
Körner, et al. 2011 is a recent adaptation of Kapos, et al. 2000, which is used for the GMBA mountain biodiversity portal (see
Online Databases and Web Pages). It defines mountains solely based on slope and combines this with a definition of elevational
vegetation belts based on bioclimatic variables.
Mountain Regions of the World
Mountains occur mostly as part of elongated mountains ranges or chains. Consequently, different mountain regions often have
distinctive features. For an overview of mountain habitats in different regions the WWF Ecoregions and Biodiversity Hotspot web
pages (see Online Databases and Web Pages) and Stone 1992 (cited under Rise of a Global Mountain Research Community) may
be consulted. Some books on particular vegetation zones in mountains also include regional accounts (e.g. Wielgolaski 1997, cited
under Alpine Vegetation). An excellent overview of mountain regions of the world is Burga, et al. 2004. The longest mountain range
is found along the west coast of both North and South America (the American Cordilleras, stretching from Alaska to southern
Chile). The largest and highest mountain area is in the Hindu Kush-Himalaya-Tibet region. Other important mountain areas are
located in the eastern Americas (e.g., Appalachians, Guayana Highlands), Asia (Caucasus, Irano-Anatolian mountains, Pamir, Tien
Shan, Urals, Altai, eastern Siberian mountains, Western Ghats, mountains of China), Europe (Alps, Pyrenees, Scandes,
Carpathians, Mediterranean mountain systems), Africa (Atlas, Ethiopian Highlands, east Africa, Great Escarpment in southern
Africa), and on islands (e.g., Southeast Asia, Japan, New Zealand, Hawaii). There is a wealth of regional accounts available. A
small selection of this literature is listed by region.
Burga, C. A., F. Klötzli, and G. Grabherr, eds. 2004. Gebirge der Erde. Landschaft, Klima, Pflanzenwelt. Stuttgart: Ulmer.
The main mountain systems of the world are examined here. For each region, information is given on geography, climate, and
human dimensions with a main focus on vegetation zones. Although in German, photographs, diagrams, and species and literature
lists are useful for non-German readers. An older but thematically broader overview of major mountain systems of the world is
Herbert Franz’s Hochgebirge der Erde (Berlin: Urania-Verlag, 1989).
AMERICAN CORDILLERAS
The “California floristic province” and “Mesoamerica” biodiversity hotspots include North American mountain ecosystems and
Mesoamerican montane forest, respectively (see Online Databases and Web Pages). The North American and Mesoamerican
mountain vegetation is extensively treated in Barbour and Billings 2000. Chapters 3, 4, and 15 in Körner and Spehn 2002 (cited
under General Overviews) cover plant and vertebrate diversity in alpine and tree-line habitat in the Rocky Mountains. Information
on global change impacts in the region can be found through CIRMOUNT (see Online Databases and Web Pages). Dale, et al.
2005 and Bowman and Seastedt 2001 comprehensively discuss two prominent North American mountain locations: Mount St.
Helens and Yellowstone National Park, respectively (see Mountain Ecosystems). The South American Andes are included in the
Colombia, Ecuador, Peru, and Venezuela biodiversity hotspots (see Online Databases and Web Pages). Borsdorf and Stadel 2014
is a comprehensive treatment of the Andes. High-altitude biodiversity in the Andes (in the Paramo vegetation zone) is covered in
many chapters in Vuilleumier and Monasterio 1986 (cited under Tropical Alpine Environments). Chapters 5 and 6 in Körner and
Spehn 2002 (cited under General Overviews) address plant diversity in the Andes along elevational gradients and across the whole
mountain system, respectively. Cuesta, et al. 2012 focuses on global change impacts on high-altitude biodiversity in the Andes.
Barbour, M. G., and W. D. Billings, eds. 2000. North American terrestrial vegetation. 2d ed. Cambridge, UK: Cambridge
Univ. Press.
A reference book for vegetation types in North America. Among others chapter 3 (“Forests and Meadows of the Rocky Mountains”
by R. K. Peet, pp. 75–121), chapter 5 (“Californian Upland Forests and Woodlands” by M. G. Barbour and R. A. Minnich, pp. 161–
202), chapter 7 (“Intermountain Valleys and Lower Mountain Slopes” by N. E. West and J. A. Young, pp. 255–284), and chapter 14
(“Alpine Vegetation” by W. D. Billings, pp. 537–572) address mountains.
Borsdorf, A., and C. Stadel. 2014. The Andes: A geographical portrait. Berlin: Springer.
An easily accessible and colorful textbook. The first three chapters introduce the geography, geology, climate, ecosystems, and
nature conservation issues of the Andes. The remaining chapters are focused on socioeconomic dimensions. The book is available
in electronic form and also in German.
Cuesta, F., P. Muriel, S. Beck, et al., eds. 2012. Biodiversidad y cambio climático en los Andes Tropicales. Conformación
de una red de investigación para monitorear sus impactos y delinear acciones de adaptación CONDESAN, Lima-Quito &
GLORIA-Andes.
Gives an overview of alpine biodiversity of the tropical Andes between Colombia and northern Argentina including global change
impacts. The book is available online in Spanish.
HINDU KUSH-HIMALAYA-TIBET
Himalaya is a biodiversity hotspot (see General Overviews). A thorough introduction and up-to-date scientific information give the
publication series, news blogs, and data portals of ICIMOD (see Online Databases and Web Pages). For instance, Singh, et al.
2011 summarizes for a general audience the science about climate change impacts in the region. Bawa and Kadur 2013 gives a
colorful overview of Himalayan biodiversity. Ives 2004 (cited under Rise of a Global Mountain Research Community) introduces the
decade-long debates about environmental degradation in the region. Schickhoff 2005 gives a cross-sectional view of the region by
following the tree-line ecotone from Afghanistan in the northwest to Yunnan in the southeast. Kindlmann 2012 introduces major
vegetation zones as well as their flora and fauna and conservation issues in Nepal in the center of the region.
Bawa, K., and S. Kadur. 2013. Himalaya: Mountain of life. Bangalore, India: Ashoka Trust for Research in Education and
the Environment (ATREE).
A coffee table book with beautiful photographs but also informative text that introduces the ecoregions, people, flora, fungi, and
wildlife (invertebrates to mammals) of the Himalaya.
Kindlmann, P., ed. 2012. Himalayan biodiversity in the changing world. Berlin: Springer.
Chapters 1 and 2 introduce major vegetation zones, their flora and fauna and conservation issues in Nepal in the center of the
Chapters 1 and 2 introduce major vegetation zones, their flora and fauna and conservation issues in Nepal in the center of the
region. Richly illustrated with photographs. The book is available in electronic form.
Schickhoff, U. 2005. The upper timberline in the Himalayas, Hindu Kush and Karakorum: A review of geographical and
ecological aspects. In Mountain ecosystems: Studies in treeline ecology. Edited by G. Broll and B. Keplin, 275–354. Berlin:
Springer.
Based on an extensive literature review, the author gives a cross-sectional view of the region by following the treeline ecotone from
Afghanistan in the northwest to Yunnan in the southeast. The book is available in electronic form.
Singh, S. P., I. Bassignana-Khadka, B. S. Karky, and E. Sharma. 2011. Climate change in the Hindu Kush-Himalayas: The
state of current knowledge. Kathmandu, Nepal: ICIMOD.
An easily accessible synthesis of current knowledge about climate change impacts in the Hindu Kush-Himalayas for policymakers.
Available online.
ASIA
The biodiversity hotspots Caucasus, Irano-Anatolian, mountains of Central Asia (Pamir, Tien Shan), mountains of southwest China,
and the western Ghats and Sri Lanka cover many of the major Asian mountain systems (see Online Databases and Web Pages).
The Caucasus is sometimes treated in books on European mountains (see Europe). Chapters 9 and 10 in Körner and Spehn 2002
(cited under General Overviews) treat plant diversity in Central Asia, Caucasus, Siberia, and Karakorum. Borchardt, et al. 2010
introduces the vegetation of post-Soviet mountain landscapes in southwestern Tien Shan. Tang, et al. 2006 gives an account of
species numbers in China’s different mountain systems.
Borchardt, P., M. Schmidt, and U. Schickhoff. 2010. Vegetation patterns in Kyrgyzstan’s walnut-fruit forests under the
impact of changing forest use in post-Soviet transformation. Die Erde 141:255–275.
This research article gives a glimpse of the vegetation of post-Soviet mountain landscapes in southwestern Tien Shan, including
land use and socioeconomic factors.
Tang, Z., Z. Wang, C. Zheng, and J. Fang. 2006. Biodiversity in China’s mountains. Frontiers in Ecology and the
Environment 4:347–352.
The article gives an account of species numbers in China’s different mountain systems.
EUROPE
There is a broad literature on European mountains, although to a considerable extent in local languages (e.g., German, French).
Nagy, et al. 2003 is a comprehensive synthesis of biodiversity in the alpine zone, Price 2010 is a policy-oriented assessment of
biodiversity and sustainable development, Vogiatzakis 2012 covers Mediterranean mountains, and Balint, et al. 2011 highlights the
importance of the Carpathians for European mountain biodiversity.
Balint, M., L. Ujvarosi, K. Theissinger, S. Lehrian, N. Meszaros, and S. U. Pauls. 2011. The Carpathians as a major diversity
hotspot in Europe: Distribution and protection of conservation priority areas. In Biodiversity hotspots. Edited by F. E.
Zachos and J. C. Habel, 189–205. Berlin: Springer.
The Carpathians’ biogeography and importance for European biodiversity (as a source and refugium) is discussed based on
population genetics case studies on aquatic insects. The book is available in electronic form.
Nagy, L., G. Grabherr, C. Körner, and D. B. A. Thompson, eds. 2003. Alpine biodiversity in Europe. Berlin: Springer.
Includes regional accounts of the climate, geology, geography, and vegetation of the alpine zone of all major mountain systems in
Europe from Scotland and the Scandes in the north to the Mediterranean region in the south, and from the Pyrenees in the west to
the Caucasus in the east. Synthesis chapters give an overview of plant, invertebrate, and vertebrate diversity in the alpine zone
across the region.
Price, M. F. 2010. Europe’s ecological backbone: Recognising the true value of our mountains. Copenhagen: European
Environment Agency.
A multiauthored synthesis for policymakers of current knowledge about biodiversity, climate change, protected areas, land use,
water use, ecosystem services, and socioeconomical dimensions of sustainable development in the European mountains. The
report is available online.
Vogiatzakis, I. N., ed. 2012. Mediterranean mountain environments. Oxford: Wiley-Blackwell.
Provides an introduction to mountainous areas in the Mediterranean: from the Atlas and Sierra Nevada to the Lebanese mountains
and Caucasus. Topics covered include physical geography, paleoecology, biogeography, cultural geography, land-use changes,
and climate change impacts. The book is available in electronic form.
AFRICA
Biodiversity hotspots that include African mountain systems are the Mediterranean Basin (Atlas), Cape Floristic region (Great
Escarpment), Eastern Afromontane (e.g., Ethiopian, Kenyan, and Tanzanian Highlands), Guinean Forests of West Africa (e.g.,
Cameroon and Nimba Highlands, Bioko Montane Forests ecoregion), and Maputaland-Pondoland-Albany (mountains east of the
Great Escarpment) (see Online Databases and Web Pages). An overview of African mountain systems give Messerli and Winiger
1992 and Bussmann 2006. Burgess, et al. 2007 assesses the importance of biodiversity in east African mountains, and chapters 4
and 17 in Vuilleumier and Monasterio 1986 (cited under Tropical Alpine Environments) discuss the ecology and origins of the flora
of these mountains. Further information on East African (afroalpine) plant diversity can be found in publications by Hedberg (see
Vegetation Zones and Mountain Biodiversity). Clark, et al. 2011 gives an overview of biodiversity in the Great Escarpment mountain
system of southern Africa.
Burgess, N., T. Butynski, N. Cordeiro, et al. 2007. The biological importance of the Eastern Arc Mountains of Tanzania and
Kenya. Biological Conservation 134:209–231.
Gives an overview of biodiversity patterns, both animals and plants, in the Eastern Arc Mountains of Tanzania and Kenya, and it
also discusses conservation issues. A web page online is cited in the article that gives further information on the region.
Bussmann, R. W. 2006. Vegetation zonation and nomenclature of African mountains: An overview. Lyonia 11:41–66.
Gives an overview of the vegetation zonation of a large part of Africa’s mountains. The focus is on floristic patterns.
Clark, V. R., N. P. Barker, and L. Mucina. 2011. The Great Escarpment of southern Africa: A new frontier for biodiversity
exploration. Biodiversity and Conservation 20:2543–2561.
The southern African Great Escarpment is a 5,000-kilometers-long, semi-continuous mountain range system stretching from
northwestern Angola through South Africa into eastern Zimbabwe and Mozambique. The article gives a country-based assessment
of biodiversity, endemism, research needs, and conservation priorities.
Messerli, B., and M. Winiger. 1992. Climate, environmental change, and resources of the African mountains from the
Mediterranean to the equator. Mountain Research and Development 12:315–336.
The synthesis article gives an overview of Africa’s mountain systems, vegetation zonation in these mountains, and the role of the
climatic history of the last 20,000 years for these biotas.
ISLANDS
Many oceanic island archipelagos include high-elevation islands. Most oceanic island archipelagos are included in a biodiversity
hotspot region (see Online Databases and Web Pages). Leuschner 1996 compares high elevation island vegetation worldwide, and
Whittaker, et al. 2008 emphasizes the importance of island ontogeny at geological timescales for island biodiversity.
Leuschner, C. 1996. Timberline and alpine vegetation on the tropical and warm-temperate oceanic islands of the world:
Elevation, structure and floristics. Vegetatio 123:193–206.
Compares treeline and alpine vegetation across high-elevation, mid-latitude islands worldwide.
Whittaker, R. J., K. A. Triantis, and R. J. Ladle. 2008. A general dynamic theory of oceanic island biogeography. Journal of
Biogeography 35:977–994.
Emphasizes that island biodiversity and its key biological processes (migration, speciation, extinction) must be understood with the
dynamics of island ontogeny (formation, changes in altitude and topographic complexity, and submergence) in mind.
Climate, Geomorphology, Hydrology
Physical aspects of mountains differ between regions (see Mountain Regions of the World) and vegetation zones (see Vegetation
Zones). There are, however, also physical characteristics that many mountains share. Elevational gradients and orographic effects
(climate patterns induced by local topography; in particular strong contrasts in rainfall patterns between leeward and windward
sides of mountains) generate high climatic variability. Volcanic and tectonic processes shape mountains, and steep slopes
(combined with climatic extremes) result in highly dynamic geomorphology and frequent natural disturbances (e.g., landslides,
erosion, avalanches, glaciers). Mountain processes (especially in hydrology) affect ecosystems and people at the foot of
mountains. High elevations are characterized by similar conditions such as an islandlike topography, high UV radiation, low
atmospheric pressure, and often harsh climates (cold temperature, wind exposure, and dryness). Several chapters in Price, et al.
2013 (cited under General Overviews) give excellent overviews of physical aspects of mountains: mountain formation (chapter 2),
climate (chapter 3), snow, ice, avalanches, and glaciers (chapter 4), geomorphology (chapter 5), and soils (chapter 6). More
detailed information can be found in general textbooks on these topics. Here only a few mountain-specific textbooks are listed:
Barry 2013 on mountain climate, Lu and Godt 2013 on geomorphology and hydrology in steep terrain, and Wohl 2010 on mountain
rivers and hydrology.
Barry, R. G. 2013. Mountain weather and climate. 3d ed. Cambridge, UK: Cambridge Univ. Press.
A textbook on mountain climatology. The book is available in electronic form.
Lu, N., and J. W. Godt. 2013. Hillslope hydrology and stability. Cambridge, UK: Cambridge Univ. Press.
A textbook on hillslope stability and rainfall-induced landslides. The bulk of the book introduces quantitative approaches for
advanced students in geology or geo-engineering, but the introductory chapters are easily accessible for a broad audience. The
book is available in electronic form.
book is available in electronic form.
Wohl, E. 2010. Mountain rivers revisited. Washington, DC: American Geophysical Union.
A textbook on mountain river geomorphology, hydrology, chemistry, and biology. The book is available in electronic form.
Vegetation Zones
A key feature of mountains is that they encompass different types of ecosystems (vegetation zones) along their elevational
gradients. While characteristics of the highest elevational zones (alpine and nival) tend to converge, at the foot of mountains all
climates of the world are represented—from cold to hot and from dry to humid. Consequently, the range of climates represented
along elevational gradients varies widely between mountains. In a particular mountain system often a wide range of ecosystem
types is represented along the elevational gradient (e.g., from tropical humid to alpine). Therefore there is no such thing as a single
mountain biome; there are different types of ecosystems in different elevational belts. Textbooks on vegetation science (including
the classical work of Heinrich Walther) generally include an introduction to vegetation zonation in mountains, where such zonal
biomes are sometimes called orobiomes. In French, Ozenda 2002 introduces vegetation zonation in mountains of the temperate
zone. Historically an important vegetation classification system based on climate variables is the life zone concept of Holdridge
1967. A general introduction of vegetation zonation in mountains is found in chapter 7 of Price, et al. 2013 and Körner 2004 (both
cited under General Overviews), a formal definition of different zones based on bioclimatic variables is given in Körner, et al. 2011
(cited under Defining the Mountain Biome). Alexander von Humboldt’s work (in Buttimer 2012), Merriam and Steineger 1890,
Schroeter 1926, Daubenmire 1943, and Whittaker 1956 illustrate historical work on the topic (all cited under Early Plant Ecology
and Biogeology). More detailed information for different world regions can be found in regional accounts (see Mountain Regions of
the World). In the following, the most relevant vegetation zones from mountaintops to mid-elevation are covered: the nival, alpine,
treeline ecotone, and montane forest zones, and inselbergs as a special mountain type.
Holdridge, L. R. 1967. Life Zone Ecology. San Jose, Costa Rica: Tropical Science Center San Jose.
In the 1940s Holdridge developed a “life zones” classification scheme based on bioclimatic variables and built on earlier work by
Merriam (see Merriam and Steineger 1890, cited under Early Plant Ecology and Biogeology). It equals altitudinal belts to latitudinal
regions; an analogy that has limitations (see chapter 7 of Price, et al. 2013, cited under General Overviews). The Holdridge biome
classification is still important.
Ozenda, P. 2002. Perspectives pour une géobiologie des montagnes Lausanne, Switzerland: Presses Polytechniques et
Universitaires Romandes.
Provides an overview of vegetation zonation and ecological features of mountain systems in the temperate zone of the Northern
Hemisphere. In French.
NIVAL ZONE (INCLUDING SNOWBED VEGETATION)
The nival zone is characterized by snow cover throughout most of the year and consequently a short growing season. In shaded
micro-topographical locations such as in depressions, snowbeds can form that are also characterized by a long duration of snow
cover and a specially adapted biota. Laybourn-Parry, et al. 2012 discusses microbial life in ice and snow. More information on the
nival zone and life in snow and ice environments can be found in the literature listed under Alpine Vegetation, and in the entries on
Polar Regions and African Biomes of Oxford Bibliographies in Ecology.
Laybourn-Parry, J., M. Tranter, and A. J. Hodson. 2012. The ecology of snow and ice environments. Oxford: Oxford Univ.
Press.
An easily accessible textbook on the biology (mostly microorganisms) of ice and snow habitats in polar and alpine environments.
ALPINE VEGETATION
Alpine vegetation is found below the nival zone and above the treeline. In the alpine zone typically grasslands (montane tundra),
mires, and low-statured heathlands are found. Alpine and arctic environments are often treated together, but there are important
differences that are discussed in chapter 7 of Price, et al. 2013 (cited under General Overviews). There are several comprehensive
treatments of alpine ecosystems and ecological adaptation to such environments available including the classical synthesis by
Chapin and Körner 1996 and Wielgolaski 1997, the comprehensive recent textbooks by Körner 2003 and Nagy and Grabherr 2009,
and Lütz 2012 with a focus on the ecophysiology of alpine plants. Cushion plants that are one important life form of alpine
environments are comprehensively covered in Aubert, et al. 2014. Although the books listed in this section focus on the alpine
zone, they give information of more general relevance for mountain ecology; in particular on important microhabitats such as rock
vegetation and disturbance-influenced habitat (e.g., succession on glacier forefields, fell-fields).
Aubert, S., F. Boucher, S. Lavergne, J. Renaud, and P. Choler. 2014. 1914–2014: A revised worldwide catalogue of cushion
plants 100 years after Hauri and Schröter. Alpine Botany 124:59–70.
A typology of cushion plants and a comprehensive global synthesis of their taxonomic and geographic distribution. A web catalogue
gives access to the full data online.
Chapin, F. S., and C. Körner. 1996. Arctic and alpine biodiversity: Its patterns, causes and ecosystem consequences. In
Functional roles of biodiversity: A global perspective. Scope 55. Edited by H. A. Mooney, J. H. Cushman, E. Medina, O. E.
Sala, and E. D. Schulze, 7–32. Chichester, UK: Wiley.
This synthesis chapter gives a good overview of the state of knowledge at the time about the role of biodiversity in arctic and alpine
ecosystems, although the main focus is on arctic ecosystems and on plants. It builds on a book edited by the same authors that
was published by Springer in 1995. The article is available online.
Körner, C. 2003. Alpine plant life: Functional plant ecology of high mountain ecosystems. 2d ed. Berlin: Springer.
A classical textbook on plant adaptation to alpine environments: cold temperature adaptation, plant nutrition, water relations, carbon
budgets and growth, phenology, and plant reproduction. Includes also a chapter on treelines and some information on elevational
gradients in mountains.
Lütz, C., ed. 2012. Plants in alpine regions: Cell physiology of adaption and survival strategies. Berlin: Springer.
On the ecophysiology of plants in alpine and nival environments with an emphasis on cell physiology.
Nagy, L., and G. Grabherr. 2009. The biology of alpine habitats. Oxford: Oxford Univ. Press.
A textbook on alpine habitats with a focus on biogeography, ecosystem ecology (including the role of animals), and global change
impacts.
Wielgolaski, F. E., ed. 1997. Polar and alpine tundra. Amsterdam: Elsevier.
Includes regional accounts of alpine vegetation in the Alps, the Carpathians, the former USSR, central Himalaya, tropical Africa,
southern Africa, northern America, southern American Paramo, and New Zealand. Although the focus is on high-altitude
ecosystems, some of the chapters also give more general information on geography, biogeography, climate, and human use of
these mountains systems.
these mountains systems.
Tropical Alpine Environments
Vuilleumier and Monasterio 1986 and Rundel, et al. 1994 cover biodiversity patterns and plant adaptation, respectively, in tropical
alpine environments. A concise account of tropical alpine environments and plant adaptation gives Lüttge 2008. Hedberg and
Hedberg 1979 is a classical treatise of plant life forms in this habitat, and Anthelme and Dangles 2012 reviews recent literature on
plant-plant interactions.
Anthelme, F., and O. Dangles. 2012. Plant-plant interactions in tropical alpine environments. Perspectives in Plant
Ecology, Evolution and Systematics 14:363–372.
A literature review on plant-plant interactions in tropical alpine environments.
Hedberg, I., and O. Hedberg. 1979. Tropical-alpine life-forms of vascular plants. Oikos 33:297–307.
A classical paper on plant life forms in tropical alpine environments. A more comprehensive but less accessible excellent work by
Olov Hedberg is Features of Afroalpine Plant Ecology (Acta Phytogeographica Suecica 49, 1964).
Lüttge, U. 2008. Páramos. In Physiological ecology of tropical plants. 2d ed. Edited by U. Lüttge, 419–441. Berlin: Springer.
A concise introduction to different plant life forms (including giant rosette plants, tussock grasses and sedges, cushion plants, and
sclerophyllous shrubs) in tropical alpine environments. The book is available in electronic form.
Rundel, P. W., A. P. Smith, and F. C. Meinzer. 1994. Tropical alpine environments: Plant form and function. Cambridge,
UK: Cambridge Univ. Press.
A classical textbook on the functional ecology of plants in tropical alpine environments. The book is available in electronic form.
Vuilleumier, F., and M. Monasterio, eds. 1986. High altitude tropical biogeography. Oxford: Oxford Univ. Press.
A classical reference book on the evolution and biogeography of alpine biotas (plant, invertebrates, vertebrates) in the Andes and
East Africa.
TREELINES
Treelines separate the montane from the alpine zone. At treelines the elevational distribution of a life form—tall trees of at least a
few meters—stops. Above the treeline only small statured woody species and herbaceous species are present. The treeline
ecotone is often a transition zone and not a strict line. Sometimes a distinction is made between the upper limit of closed montane
forest (timberline) and the beginning of the treeless alpine zone where the occurrence of individual and low-statured (krummholz)
trees stops (treeline). Treelines have long fascinated ecologists as exemplified in Troll 1973 (cited under Rise of a Global Mountain
Research Community). Körner 2012, Holtmeier 2009, and Schickhoff 2005 (cited under Hindu Kush-Himalaya-Tibet) give an
overview of historical research on treelines. Several books summarize the state of knowledge about treelines in mountains
including Körner 2012 and Holtmeier 2009. Information on regional treeline patterns is found in regional accounts Schickhoff 2005
for the Himalayas and Leuschner 1996 for islands (cited under Mountain Regions of the World).
Harsch, M. A., P. E. Hulme, M. S. McGlone, and R. P. Duncan. 2009. A global meta-analysis of treeline response to climate
warming. Ecology Letters 12:1040–1049.
warming. Ecology Letters 12:1040–1049.
A meta-analysis of treeline changes in response to climate change at 166 sites (both at high latitude and elevation).
Holtmeier, F.-K. 2009. Mountain timberlines: Ecology, patchiness, and dynamics. Berlin: Springer.
A comprehensive synthesis of research on treelines in mountains that covers both historical and recent work. The book is available
in electronic form.
Körner, C. 2012. Alpine treelines: Functional ecology of the global high elevation tree limits. Berlin: Springer.
A concise textbook on the functional ecology of plant life at treelines in mountains. The book is available in electronic form.
Körner, C., and J. Paulsen. 2004. A world-wide study of high altitude treeline temperatures. Journal of Biogeography
31:713–732.
Based on field measurements at forty-six treeline sites the authors conclude that high altitude treelines are globally associated with
a common thermal threshold (seasonal mean ground temperature of 5–8 °C).
MONTANE FORESTS (INCLUDING MONTANE CLOUD FORESTS)
Below the timberline is where the montane forest zone starts. Montane forests are important to mountain people as they protect
water reservoirs and against natural hazards; they are also a source of wood and other products, and they are an important habitat
of diverse mountain wildlife. In northern temperate regions montane forests are generally dominated by conifers, while in the
Southern Hemisphere broad-leaved tree species dominate (e.g., Eucalyptus in Australian mountains, Nothofagus and/or Araucaria
in New Zealand, Argentina, and Chile). In the tropics, montane cloud forests form that are often immersed in the cloud layer and
are characterized by stunted trees, abundant mosses, epiphytes and ferns, and a thick humus layer in the topsoil. Montane cloud
forests are considered cradles and refugia of biodiversity and a priority habitat for conservation. During dry seasons they depend in
particular on clouds as a water source, and there is concern that an elevational shift of the cloud layer due to climate change could
threaten them in many world regions. General information about montane forests can be found in chapter 7 in Price, et al. 2013
(cited under General Overviews) and literature about the treeline ecotone (see Treelines). More specific literature can be found in
regional accounts (see Mountain Regions of the World): see, for instance, several chapters in Barbour and Billings 2000 (cited
under American Cordilleras). Price and Butt 2000 is a comprehensive assessment of the ecology and conservation of mountain
forests, and Price, et al. 2011 is a policy-oriented summary. Bruijnzeel, et al. 2010 is a comprehensive overview about tropical
montane cloud forests. Eller, et al. 2013 demonstrates a unique adaptation of montane cloud forest plants.
Bruijnzeel, L. A., F. N. Scatena, and L. Hamilton. 2010. Tropical montane cloud forests: Science for conservation and
management. Cambridge, UK: Cambridge Univ. Press.
A comprehensive overview of current knowledge about tropical montane cloud forests. It comprises seventy-two chapters divided
into seven sections covering a wide spectrum of topics including cloud forest distribution, climate, soils, biodiversity, hydrological
processes, hydrochemistry and water quality, climate change impacts, and cloud forest conservation, management, and
restoration. Available in electronic form.
Eller, C., A. Lima, and R. Oliveira. 2013. Foliar uptake of fog water and transport belowground alleviates drought effects in
cloud forest tree species, Drimys brasilensis (Winteraceae). New Phytologist 199:151–162.
There is a growing body of research demonstrating that trees covered in clouds in a montane cloud forest can absorb water
through their leaves and move it downward into the soil.
Price, M. F., and N. Butt. 2000. Forests in sustainable mountain development: A state of knowledge report for 2000. New
York: CABI.
A comprehensive assessment of the ecology and conservation of mountain forests that includes both thematic synthesis chapters
and regional case studies. The book focuses on ecosystem services including cultural, institutional, and political aspects. Freely
available online.
Price, M. F., G. Gratzer, L. A. Duguma, T. Kohler, D. Maselli, and R. Romeo. 2011. Mountain forests in a changing world:
Realizing values, addressing challenges. Rome: FAO.
An easily accessible and colorful brochure on ecosystem services provided by mountain forests and on conservation issues.
Includes both short synthetic overviews of different topics and case studies from different world regions. Freely available online.
INSELBERGS
Inselbergs are isolated and steep-sided rock outcrops that rise above plains and are characterized by harsh environmental
conditions and a rocky underground with shallow soils. Porembski and Barthlott 2000 is a comprehensive treatment of inselberg
ecology, and Lüttge 2008 is a concise overview.
Lüttge, U. 2008. Inselbergs. In Physiological ecology of tropical plants. 2d ed. Edited by U. Lüttge, 379–418. Berlin:
Springer.
A concise introduction to plant life on inselbergs. The book is available in electronic form.
Porembski, S., and W. Barthlott, eds. 2000. Inselbergs: Biotic diversity of isolated rock outcrops in tropical and temperate
regions. Berlin: Springer.
This edited book gives a comprehensive overview of the geomorphology, environmental conditions, plant ecophysiology, and
algae, fungi, lichens, bryophytes and vascular plant and faunistic diversity on inselbergs. Regional accounts comprehensive treat
inselbergs in different regions in Africa, North and South America, and on islands.
Mountain Biodiversity
Mountains are global biodiversity hotspots with unique floras and faunas. General overviews of mountain biodiversity can be found
in chapters 7 (vegetation) and 8 (wildlife) in Price, et al. 2013; and Körner, et al. 2005; Körner 2004; and Körner and Spehn 2002
(cited under General Overviews). Information on biodiversity in particular mountain ecosystems are accessible through the WWF
Ecoregions and Biodiversity Hotspots web pages (see Online Databases and Web Pages), regional accounts (see Mountain
Regions of the World) and publications on different vegetation zones (see Vegetation Zones). In this section literature is included
that highlight aspects of mountain biodiversity and biogeography that represent general characteristics of mountains rather than of
particular vegetation zones including: large-scale biogeographic patterns (Hughes and Eastwood 2006, Rickart 2001), vicariant
species (Hedberg 1969), and glacial refugia (Birks 2008, Holderegger and Thiel-Egenter 2009, Dobrowski 2011). Particularly
important features of mountains are elevational gradients that are treated in a separate section.
Birks, H. John. 2008. The late-Quaternary history of arctic and alpine plants. Plant Ecology and Diversity 1:135–146.
This easily accessible short review of the paleohistory of arctic and alpine plants during recent glacial periods is rich in detailed
knowledge and broad in coverage. It explains the empirical evidence that comes from plant fossils, reconstructs responses of arctic
and alpine plants to different phases of Quaternary climate changes, and gives also a short historic account of research on the
and alpine plants to different phases of Quaternary climate changes, and gives also a short historic account of research on the
subject.
Dobrowski, S. Z. 2011. A climatic basis for microrefugia: The influence of terrain on climate. Global Change Biology
17:1022–1035.
The role and definition of glacial microrefugia at a fine spatial resolution has attracted recent research interest especially because it
is expected that it will help to better understand the potential of mountain species to persist in mountain microhabitats despite
climate change. See also Scherrer and Körner 2011, cited under Mountain Ecosystems.
Hedberg, O. 1969. Evolution and speciation in a tropical high mountain flora. Biological Journal of the Linnean Society
1:135–148.
Vicarious species form through the division of an original population into two or more populations by a geographic barrier. Vicariant
speciation is common in mountains thanks to their topographic heterogeneity and frequent major disturbances (e.g., volcanoes).
The article gives an overview of vicariant species in the afroalpine flora of eastern Africa.
Holderegger, R., and C. Thiel-Egenter. 2009. A discussion of different types of glacial refugia used in mountain
biogeography and phylogeography. Journal of Biogeography 36:476–480.
An important aspect of the biogeography of temperate mountain biotas is where species survived ice ages. The article discusses
three main types of glacial refugia of mountain species: nunatak, peripheral, and lowland refugia.
Hughes, C., and R. Eastwood. 2006. Island radiation on a continental scale: Exceptional rates of plant diversification after
uplift of the Andes. Proceedings of the National Academy of Sciences 103:10334–10339.
Many different north temperate plant genera arrived in the Andes from North America after uplift of the northern Andes where they
then speciated in high elevation habitat. This paper analyzes such a plant radiation in the Lupinus genus. The analysis illustrates
the interplay of three factors for the formation of mountain biotas: mountain chains as corridors, mountaintops and mountain
habitats as islands, and mountain dynamics in geological times.
Rickart, E. A. 2001. Elevational diversity gradients, biogeography and the structure of montane mammal communities in
the intermountain region of North America. Global Ecology and Biogeography 10:77–100.
The author analyzes distribution patterns of nonvolant small mammals along elevational gradients in mountain ranges in the
western United States. The analysis illustrates how the interplay of paleohistory, elevational gradients, and the islandlike
topography of mountains can shape the biogeography of mountain biotas.
ELEVATIONAL GRADIENTS
Elevational gradients are a defining feature of mountains and are therefore of fundamental importance to the understanding of
mountain biodiversity. McCain and Grytnes 2010 presents a synthesis of the current understanding of species richness patterns
along elevational gradients, and Körner 2007 is a critical assessment of the use of elevational gradients in ecological research
(compare also Nunez-Farfan and Schlichting 2001, cited under Early Plant Ecology and Biogeography). Janzen 1967 highlights an
important difference between elevational gradients in temperate regions and the tropics. Changes in the elevational distribution of
species as a result of ongoing climate change and its consequences for mountain biodiversity has attracted much recent research
interest (Pauli, et al. 2012; Crimmins, et al. 2011; Chen, et al. 2009; Laurance, et al. 2011, and Harsch, et al. 2009 [cited under
Treelines] on treeline changes in Vegetation Zones).
Chen, I.-C., H.-J. Shiu, S. Benedick, et al. 2009. Elevation increases in moth assemblages over 42 years on a tropical
mountain. Proceedings of the National Academy of Sciences 106:1479–1483.
A non-plant and tropical example of elevational species shifts due to climate change.
Crimmins, S. M., S. Z. Dobrowski, J. A. Greenberg, J. T. Abatzoglou, and A. R. Mynsberge. 2011. Changes in climatic water
balance drive downhill shifts in plant species’ optimum elevations. Science 331:324–327.
The analysis highlights that climate change will also affect water availability and that both uphill and downhill shifts of species
distributions can result.
Janzen, D. H. 1967. Why mountain passes are higher in the tropics. American Naturalist 101:233–249.
A classic paper that highlights an important difference between mountains of the temperate and tropical zones. Due to seasonal
climate fluctuations and longer-term glacial and interglacial phases temperate zone species tend to have broader climate niches,
and low and high elevation climates are similar during summer months. In contrast, high elevation climates in the tropics strongly
contrast with the surrounding landscape year round.
Körner, C. 2007. The use of “altitude” in ecological research. Trends in Ecology and Evolution 22:569–574.
The author emphasizes that multiple environmental factors vary along elevational gradients (including atmospheric pressure,
temperature, moisture, season length, geology and human land use) and that their interplay must be taken into account in
elevational gradient studies.
Laurance, W. F., D. Carolina Useche, L. P. Shoo, et al. 2011. Global warming, elevational ranges and the vulnerability of
tropical biota. Biological Conservation 144:548–557.
The authors argue based on elevational distribution data that tropical species tend to be thermally specialized and therefore
particularly vulnerable to global warming. Compare Janzen 1967 above.
McCain, Christy M., and John-Arvid Grytnes. 2010. Elevational gradients in species richness. In Encyclopedia of life
sciences (ELS). Chichester, UK: John Wiley.
A comprehensible and easily accessible synthesis of the current understanding of the mechanisms that form species richness
patterns along elevational gradients.
Pauli, H., M. Gottfried, S. Dullinger, et al. 2012. Recent plant diversity changes on Europe’s mountain summits. Science
336:353–355.
This vegetation monitoring study presents recent (2001 to 2008) changes in vascular plant species richness across Europe’s major
mountain ranges. Species have moved upslope on average. These shifts had a positive effect on the summit floras’ species
richness in boreal-temperate mountains and a negative one on Mediterranean mountains, probably because recent climatic trends
have decreased the availability of water in the European south.
Mountain Ecosystems
Many aspects of ecosystem ecology in mountains are specific to particular vegetation zones (see Vegetation Zones). In this section
Many aspects of ecosystem ecology in mountains are specific to particular vegetation zones (see Vegetation Zones). In this section
the literature presented treats selected ecosystem processes that are of more general importance to mountains. At a local scale
there is ecology of high-elevation ecosystems (Bowman and Seastedt 2001), major disturbances (Dale, et al. 2005), soil stability
(Pohl, et al. 2009), and microtopography (Scherrer and Körner 2011). At a landscape-scale there is large wildlife (White, et al.
2013), insect outbreaks (Büntgen, et al. 2009), redistribution of chemical elements (Seastedt, et al. 2004), and linkages between
mountains and surrounding landscapes (Gomi, et al. 2002, Green 2008).
Bowman, W. D., and T. R. Seastedt. 2001. Structure and function of an alpine ecosystem: Niwot Ridge, Colorado. Oxford:
Oxford Univ. Press.
The long-term ecological research (LTER) site at Niwot Ridge in Colorado is the source of some of the most comprehensive and
long-term research on a high elevation ecosystem (at above 3000 m asl). For updates see related information online.
Büntgen, U., D. Frank, A. Liebhold, et al. 2009. Three centuries of insect outbreaks across the European Alps. New
Phytologist 182:929–941.
Based on dendrochronological data, the authors demonstrate that cyclical larch budmoth outbreaks persisted in the European Alps
for centuries.
Dale, V. H., F. J. Swanson, and C. M. Crisafulli. 2005. Disturbance, survival, and succession: Understanding ecological
responses to the 1980 eruption of Mount St. Helens. Berlin: Springer.
In 1980 a major eruption destroyed large tracts of vegetation at Mount St. Helens in the northwest United States. The volcano has
become an outdoors laboratory for studying how mountain ecosystems response to major disturbances and how the barren
landscape is recolonized. The book brings together the work of plant, animal, and ecosystem ecologists on secondary successions
at the site. The book is available in electronic form.
Gomi, T., R. C. Sidle, and J. S. Richardson 2002. Understanding processes and downstream linkages of headwater
systems. Bioscience 52:905–916.
An exemplary article that discusses how the ecology of headwaters in mountains influences the ecology of ecosystems
downstream.
Green, K. 2008. Migratory bogong moths (Agrotis infusa) transport arsenic and concentrate it to lethal effect by estivating
gregariously in alpine regions of the Snowy Mountains of Australia. Arctic, Antarctic, and Alpine Research 40:74–80.
The bogong moth migrates annually to the Australian Alps in spring, before returning to the surrounding plains in autumn to breed.
In the alpine zone they occur at high density and are an important food source for mountain wildlife. This study indicates that
through this migration, high levels of arsenic are brought to an alpine ecosystem from the surrounding lowlands.
Pohl, M., D. Alig, C. Körner, and C. Rixen. 2009. Higher plant diversity enhances soil stability in disturbed alpine
ecosystems. Plant and Soil 324:91–102.
An exemplary article that demonstrates the importance of plant diversity for soil stability in mountain ecosystems.
Scherrer, D., and C. Körner. 2011. Topographically controlled thermal-habitat differentiation buffers alpine plant diversity
against climate warming. Journal of Biogeography 38:406–416.
Based on high-resolution infra-red thermometry substantial temperature variability at a microscale due to microtopography is
demonstrated for three alpine slopes of contrasting exposure. The authors argue that such climatic variability will help mountain
species to persist despite climate change. See also Dobrowski 2011, cited under Mountain Biodiversity.
Seastedt, T. R., W. D. Bowman, T. N. Caine, D. McKnight, A. Townsend, and M. W. Williams. 2004. The landscape
continuum: A model for high-elevation ecosystems. Bioscience 54:111–121.
The authors present a conceptual model of the landscape-scale redistribution of chemical elements in mountains that links
terrestrial and aquatic ecosystems.
White, P. J., R. A. Garrott, G. E. Plumb, eds. 2013. Yellowstone’s wildlife in transition. Cambridge, MA: Harvard Univ.
Press.
In contrast to many plants and smaller animals, large predators and herbivores in mountains are often generalists with broad
elevational activity ranges. The book presents a synthesis of ecosystem research at Yellowstone National Park (United States) with
a focus on the role of large wildlife as keystone species and associated human conflicts (e.g., wolves, overabundance of deer).
Land Use, Global Change Impacts, and Sustainable Development
Mountains are of great importance to humanity especially as water reservoirs and tourism destinations. The services provided by
mountain ecosystems are essential for soil stability, protection against natural hazards, climate regulation, agriculture, forestry, and
products for local livelihoods. Many mountains have special cultural or spiritual significance (e.g., Mount Fuji in Japan or Mount
Etna in Italy). Many mountain systems experienced a long history of human land use, and more recently often abrupt land-use
changes both due to intensification and abandonment. Mountain ecosystems are increasingly exposed to human pressures
including land-use changes (e.g., agriculture, tourism, urbanization, or abandonment), pollution, and climate change (e.g., loss of
glaciers, melting of permafrost, desertification, increased frequency of natural hazards such as floods, landslides and avalanches,
and spread of diseases and invasive species). The human dimension of mountains is extensively covered in literature listed under
other headings, especially: chapters 9–12 in Price, et al. 2013 and Körner, et al. 2005 (cited under General Overviews). Historical
Accounts and Foundational Works presents milestones in the formation of a global environmental awareness and sustainable
development research in mountains. Information on threats affecting particular mountain ecosystems are accessible through the
WWF Ecoregions and Biodiversity Hotspots web pages and other online sources (see Online Databases and Web Pages), regional
accounts (see Mountain Regions of the World) and publications on different vegetation zones (see Vegetation Zones). White, et al.
2013 (cited under Mountain Ecosystems) discusses wildlife management issues (large predators and overabundance of large
herbivores). There are a number of comprehensive publications about global change and conservation issues in mountains that are
not necessarily all up-to-date and/or are compilations of case studies, but they give a good overview of the relevant issues and
provide a good starting point for searching more recent and specific literature: global change (Huber, et al. 2005), land-use impacts
(Spehn, et al. 2006), ecological connectivity (Worboys, et al. 2010), invasive species (Kueffer, et al. 2013), ecosystem services
(Grêt-Regamey, et al. 2012), and sustainable development (Price, et al. 2004). In mountains, climate change impacts are of
particular importance, and they are therefore treated in a separate section.
Grêt-Regamey, A., S. H. Brunner, and F. Kienast. 2012. Mountain ecosystem services: Who cares? Mountain Research and
Development 32:3–34.
This short open-access article reviews the literature on ecosystem services in mountains.
Huber, U. M., H. K. Bugmann, and M. A. Reasoner. 2005. Global change and mountain regions: An overview of current
knowledge. Berlin: Springer.
A compilation on global change in mountains, including sections on paleoenvironmental, cryospheric, hydrological and ecological
changes, and human dimensions. The book is available in electronic form.
Kueffer, C., K. McDougall, J. Alexander, et al. 2013. Plant invasions into mountain protected areas: Assessment,
prevention and control at multiple spatial scales. In Plant invasions in protected areas: Patterns, problems and
challenges. Edited by F. L. C. P. Py!ek, D. M. Richardson, and P. Genovesi, 89–113. Berlin: Springer.
Synthesizes current knowledge on plant invasions in mountains. It builds in particular on the work of the Mountain Invasion
Research Network that can be found online. The book is available in electronic form.
Price, M. F., L. F. Jansky, and A. A. Iatsenia. 2004. Key issues for mountain areas. New York: United Nations Univ. Press.
On socioeconomic dimensions of sustainable development in mountains. The book is available in electronic form.
Spehn, E. M., M. Libermann, and C. Körner, eds. 2006. Land use change and mountain biodiversity. Andover, UK: CRC.
An edited volume that includes mostly local case studies on the impact of fire and grazing on mountain biodiversity.
Worboys, G. L., W. L. Francis, and M. J. Lockwood. 2010. Connectivity conservation management: A global guide.
Washington, DC: Earthscan.
Establishing large-scale ecological corridors at a transnational scale through connecting protected areas is a current conservation
priority. Many of these projects are established in mountain areas, as this book demonstrates. Mountain areas are, in many world
regions, the only remaining relatively undisturbed places that stretch across large distances.
CLIMATE CHANGE
Due to their often extreme climates (e.g., cold, dry, or cloud-covered), exposure to natural hazards (e.g., avalanches, landslides),
vulnerability of abiotic and biotic factors to climate variation (e.g., short growing season, snow cover), often specialized biota with
small and fragmented populations, and the islandlike topography, mountain floras, faunas and ecosystems might be particularly
vulnerable to climate change. Although species at lower elevations in mountains might also profit from short distances along steep
climate gradients for tracking a changing climate. Such short distances might, however, also lead to particularly fast spread of pests
and diseases into high elevation ecosystems once climate change has removed climate barriers (see Kueffer, et al. 2013, cited
under Land Use, Global Change Impacts, and Sustainable Development). Literature covering climate change is mentioned and
listed in the general section on Land Use, Global Change Impacts, and Sustainable Development, but there are also syntheses
focusing exclusively on climate change impacts, including chapter 7 in Barry 2013 (cited under Climate, Geomorphology,
Hydrology), Theurillat and Guisan 2001, and Grabherr, et al. 2010. Many references listed in the sections General Overviews,
Mountain Regions of the World and Vegetation Zones address climate change; for instance Harsch, et al. 2009, cited under
Treelines on shifting treelines or Bruijnzeel, et al. 2010, cited under Montane Forests (Including Montane Cloud Forests) on the
impacts on montane cloud forests. The section on Mountain Biodiversity includes literature on changing elevational species ranges
due to climate change. Gottfried, et al. 2012 documents a shift from cold-adapted to warm-adapted species on European
mountaintops through vegetation monitoring, and the modeling study Engler, et al. 2011 predicted a negative but region-specific
climate change impact on European mountain floras. Updates on climate change impacts can be found in the reports of the
Intergovernmental Panel on Climate Change (IPCC) (e.g., in chapter 4 of the 2014 report on impacts, adaptation, and vulnerability).
Engler, R., C. F. Randin, W. Thuiller, et al. 2011. Twenty-first century climate change threatens mountain flora unequally
across Europe. Global Change Biology 17:2330–2341.
In this modeling study climate change impacts on 2,632 plant species across all major European mountain ranges were assessed.
Projected habitat loss was greatest for species distributed at higher elevations and for floras from regions projected to undergo
increased warming accompanied by decreased precipitation, such as the Pyrenees and the Eastern Austrian Alps.
Gottfried, M., H. Pauli, A. Futschik, et al. 2012. Continent-wide response of mountain vegetation to climate change. Nature
Climate Change 2:111–115.
Data from sixty summits in Europe collected through GLORIA (cited under Online Databases and Web Pages) indicates that coldadapted species decline and the more warm-adapted species increase due to recent climate change.
Grabherr, G., M. Gottfried, and H. Pauli. 2010. Climate change impacts in alpine environments. Geography Compass
4:1133–1153.
A comprehensive and easily accessible literature review of climate change impacts in alpine environments.
Theurillat, J. P., and A. Guisan. 2001. Potential impact of climate change on vegetation in the European Alps: A review.
Climatic Change 50:77–109.
Although not most up-to-date, it gives a good overview of ecological processes of relevance to climate change impacts on
vegetation, including changing species distributions, vegetation shifts, phenology, productivity, and landscape-scale changes.
LAST MODIFIED: 10/28/2014
DOI: 10.1093/OBO/9780199830060-0119
BACK TO TOP
Copyright © 2014. All rights reserved.