Ecology North American Biomes
by
Debra P.C. Peters, Stacey L.P. Scroggs, Jin Yao
  • LAST REVIEWED: 11 July 2019
  • LAST MODIFIED: 30 July 2014
  • DOI: 10.1093/obo/9780199830060-0099

Introduction

Biomes of North America are contained within the land area of Canada, the United States, Mexico, and countries in Central America. The area is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the west and south by the Pacific Ocean, and to the southeast by South America and the Caribbean Sea. This large area (over 24 x 106 km2) is characterized by a broad range of temperature and precipitation that result in nine biomes ranging from tropical rainforests and seasonal deciduous forests in the south near the equator to boreal forests and tundra at high latitudes near the North Pole. Temperate forests (deciduous, coniferous), grasslands, deserts, and Chaparral woodlands occur at mid-latitudes. Landscape-scale patterns in contemporary ecosystems within each biome reflect variability in climate and soil parent material combined with human activities that have increased in extent and intensity over the past several centuries. These patterns are often influenced by the redistribution of organisms, water and sediment, fire, and air chemistry. Connections with biome types on other continents on Earth can lead to invasion by exotic species including pests and pathogens, large climatic events such as hurricanes and drought, and changes in air quality through dust storms and volcanic eruptions. These tele-connections often occur infrequently, yet with large and surprising effects on ecosystem properties and dynamics. Directional changes in climate are expected to influence biome distributions and composition in novel ways. Increasing awareness of these broad-scale dynamics that connect biomes globally is leading to new avenues of research that intersect ecology with other disciplines.

General Overviews

There are several synthetic works that describe and compare biomes within the North American continent. Many, such as Bolen 1998, Molles 2005, and Whittaker 1975, are written for undergraduate courses and are not limited to biomes of North America but rather provide information on biome types located globally. Shelford 1913 and Shelford 1963 are classics that provide a compendium of information known at that time about what we now call biomes within the North American continent. The material in these books was recently updated by Bolen 1998. Vegetation types of North America are described in Barbour and Billings 2000 that go beyond the definition of biomes to include locally important vegetation types.

  • Barbour, Michael G., and William Dwight Billings. 2000. North American terrestrial vegetation. Cambridge, UK: Cambridge Univ. Press.

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    Descriptions of vegetation types within North America, including the vegetation composition, properties of the abiotic environment, conservation issues, management problems, and areas for future research. Unique resource for students and researchers. Also see Defining and Describing North American Biomes and Biogeography of Biomes.

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  • Bolen, Eric G. 1998. Ecology of North America. New York: Wiley.

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    An updated and expanded version of Shelford 1963, this book by Bolen is a very readable introduction to biomes that is designed to acquaint undergraduate students and others interested in understanding and protecting the biomes of North America. See also Defining and Describing North American Biomes.

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  • Molles, Manuel C. 2005. Ecology: Concepts and applications. New York: Sinauer Associates.

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    This is an excellent textbook that is widely used for undergraduate courses. Ten biomes are described with their global distribution.

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  • Shelford, Victor E. 1913. Animal communities in temperate America. Chicago: Univ. of Chicago Press.

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    Although this book focuses on animal communities within 160 km of Chicago, it is notable for the biogeographic approach that was adapted by others for plants. The focus is a study of the ecology of animals within communities that lead to changes in species responses and community dynamics as the habitat changes.

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  • Shelford, Victor E. 1963. The ecology of North America. Urbana: Univ. of Illinois Press.

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    This classic treatise describes important features and successional dynamics of the major biomes of North America north of 21oN latitude. Discussion of dominant animals within each biome updates and expands Shelford 1913. The book is notable for including both plant and animal dynamics across large areas. Also see Defining and Describing North American Biomes.

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  • Whittaker, Robert H. 1975. Communities and ecosystems. New York: Macmillan.

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    An introduction to population, community, and ecosystem dynamics with descriptions and comparisons of the major biomes of the world. Undergraduate and graduate students will find this book to be excellent background to more recent comparative analyses of biomes.

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Journals

Because of the large spatial extent and large number of biomes, research on North American biomes appears in all ecological journals. The most relevant journals for patterns in environmental drivers and ecological responses across large parts of the continent are journals that are not specific to North America. Three sister journals provide comprehensive coverage of topics relevant to the study of biomes: Global Ecology and Biogeography contains papers that address general ecological hypotheses that are explored and tested using data of broad geographic, taxonomic, or temporal scope; Journal of Biogeography focuses on broad-scale patterns in species, communities, and ecosystems and how those patterns change through time; Diversity and Distributions publishes papers on the application of biogeographic principles to the conservation of biodiversity. The journal Ecography focuses on papers dealing with population and community ecology and macroecology. Ecosphere, a new online only, open-access journal published by the Ecological Society of America, encourages a broad suite of papers relevant to dynamics of biomes, although most papers focus on specific aspects of ecosystems.

Online Databases

There are a number of important online databases that provide information about North American biomes. These databases typically include either environmental driver data on climate or soils, such as the Canadian Department of Agriculture and Agri-Food, and Canada Department of the Environment, the National Atmospheric Deposition Program, US National Oceanic and Atmospheric Administration, and PRISM or they provide information on ecological patterns, such as the US Census Bureau and US Department of Agriculture soils maps and plants database. Some online databases such as EcoTrends provide both environmental driver and ecological data to promote synthesis using sites located across the North American continent, with a focus on the United States.

Historical Perspectives on Conceptualizing Biomes

The term “biome” was originally coined in 1916 by Frederick Clements at a meeting of the Ecological Society of America based on a review by J. Richard Carpenter 1939. The original definition referred only to the biotic community (plants and animals) found within a particular habitat. Similar ideas were proposed by Du Rietz 1929 in Europe. Clements and Shelford 1939 provided an expanded definition where biome was viewed as a social, complex, or super organism. Within the same time frame, Tansley 1935 developed the concept of an ecosystem that includes the physical environment as well as the biota at any spatial extent. Golly 1993 describes the International Biological Program in the 1960s as bringing together these terms where biomes were differentiated based on vegetation formations within which intensive, site-based ecosystem studies were conducted. The current view of a biome has been maintained where a complex of organisms (plants and animals) inhabit a given region defined by the abiotic environment, but the developmental connotation associated with a super organism has been dropped. Biome, ecosystem, and vegetation type are often considered synonymous at the regional scale where the recognizable feature of the biome is the vegetation or formation under current climatic conditions. Much of this work led to the field of biogeography with Dansereau 1957 providing one of the earliest syntheses of how fine-scale processes can scale up to regional biomes.

  • Carpenter, J. Richard. 1939. The biome. American Midland Naturalist 21:75–91.

    DOI: 10.2307/2420376Save Citation »Export Citation »

    Excellent historical account of the events in ecology in the United States in the early 1900s. Describes development of plant and animal ecology as well as the concept of the biome. Discussion notes at the end present question and answer session among most notable ecologists in the late 1920s and provide insight into their thinking at that time.

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  • Clements, Frederic E., and V. E. Shelford. 1939. Bio-ecology. New York: Wiley.

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    The last major work by Clements before his death in 1945. Provides the most complete description of the biome concept as a synonym of formation and climax at large spatial extents.

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  • Dansereau, Pierre. 1957. Biogeography: An ecological perspective. New York: Ronald.

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    Originally written as an undergraduate textbook, this book is one of the first to synthesize social sciences, anthropology, genetics, and biogeography into an ecological perspective. Excellent reading for graduate students as a general introduction to how ecological ideas have developed over the past fifty years.

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  • Du Rietz, Gustaf. 1929. The fundamental units of vegetation. Proceedings of the International Congress of Plant Sciences 1:623–627.

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    Although published in English, this small document is only available in a few libraries. The importance of Du Rietz to European vegetation classification is described in Lawesson, et al. 2000.

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  • Golly, Frank B. 1993. A history of the ecosystem concept in ecology: More than the sum of the parts. New Haven, CT: Yale Univ. Press.

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    Excellent historical accounts of the development of the term ecosystem, with a section on the International Biological Program and the relationships between biomes and ecosystems.

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  • Lawesson, Jonas E., ed. 2000. A concept for vegetation studies and monitoring in the Nordic countries. Copenhagen: Nordic Council of Ministers.

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    This book provides an excellent historical account of the development of vegetation classification at small scales for Nordic countries with linkages to American authors and contributions (e.g., Whittaker).

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  • Tansley, Arthur G. 1935. The use and abuse of vegetational concepts and terms. Ecology 16:284–307.

    DOI: 10.2307/1930070Save Citation »Export Citation »

    Classic paper that presents the term “ecosystem” as consisting of both the biotic and abiotic components, in contrast to biome that was defined by the plants and animals.

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  • Whittaker, R. H. 1975. Communities and ecosystems. New York: Macmillan.

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    Essential background reading for descriptions of communities and ecosystems.

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Relationships between Biome Classification and Other Systems

Large areas of homogeneous vegetation have been classified in a number of ways, depending on the goals and interests of the author. Climatologists and physical scientists typically assume that vegetation can be mapped and classified if the climate is known. A well-known and widely used approach is the Köppen-Geiger system originally published in 1884. A recent update by Peel, et al. 2007 used monthly precipitation and temperature to show that all five major climate types of the world occur in North America. A total of twenty-four classes were found that roughly correspond to major biomes. The Holdridge life zone system is a global scheme to classify land area based on climate. The underlying assumption is that soil and vegetation can be mapped and classified once climate (precipitation, temperature, potential evapotranspiration) is known. This system was originally published in 1947, updated in 1967, and applied to the United States by Lugo, et al. 1999. Other approaches use a hierarchical approach that depend on ecoregions—the term was first proposed by Orie Loucks in 1962 to refer to ecosystems (abiotic and biotic) at various spatial scales. One well-known ecoregion classification was developed by Bailey 2005 and others that delineates ecoregions at three hierarchical levels: domain, division, and province. The spatial distribution of Bailey’s divisions roughly correlates with biomes. For North America, the Bailey system defines four domains and fifteen divisions. Another well-known ecoregion classification system was developed for the World Wildlife Fund by Olson, et al. 2001. This system offers features for conservation planning at global and regional scales, and was applied to Latin America by Dinerstein, et al. 1995. Other efforts have focused on classifications aimed at response to disturbance by Omernik 1987. Efforts have compared and contrasted classification systems, in particular across US federal agencies by McMahon, et al. 2001.

  • Bailey, Robert G. 2005. Identifying ecoregion boundaries. Environmental Management 34:S14–S26.

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    A general description of the spatial locations of biomes in North America mapped as ecoregions defined as geographically distinct assemblages of species, natural communities, and environmental conditions. Describes assumptions and limitations as well as justification of the approach.

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  • Dinerstein, Eric, David M. Olson, D. J. Graham, et al. 1995. A conservation assessment of the terrestrial ecoregions of Latin America and the Caribbean. Washington, DC: World Bank.

    DOI: 10.1596/0-8213-3295-3Save Citation »Export Citation »

    A hierarchy of 191 ecoregions is proposed for Latin America. The conservation status of each eco-region is determined and combined with its biological distinctiveness to suggest geographic priorities for conservation.

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  • Lugo, Ariel E., S. L. Brown, R. Dodson, T. S. Smith, and H. H. Shugart. 1999. The Holdridge life zones of the conterminous United States in relation to ecosystem mapping. Journal of Biogeography 26:1025–1038.

    DOI: 10.1046/j.1365-2699.1999.00329.xSave Citation »Export Citation »

    Excellent example of the use of the Holdridge system for mapping life zones of the United States. The Holdridge system was applied using data at 4-km resolution, which led to thirty-eight life zones in the United States. They also compare their map with other global vegetation classification systems.

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  • McMahon, Gerald, Steven W. Gregonis, Sharon W. Waltman, et al. 2001. Developing a spatial framework of common ecological regions for the conterminous United States. Environmental Management 28:293–316.

    DOI: 10.1007/s0026702429Save Citation »Export Citation »

    This paper describes the origin, capabilities, and limitations of three major federal agency frameworks for defining ecoregions and suggests why a common framework to describe and map ecoregions for the conterminous United States is desirable.

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  • Olson, David M., E. Dinerstein, Eric D. Wikramanayake, et al. 2001. Terrestrial ecoregions of the world: A new map of life on Earth. BioScience 51:933–938.

    DOI: 10.1641/0006-3568(2001)051[0933:TEOTWA]2.0.CO;2Save Citation »Export Citation »

    Essential reading for users interested in using biomes and finer scales for conservation planning.

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  • Omernik, James M. 1987. Ecoregions of the conterminous United States. Map (scale 1:7,500,000). Annals of the Association of American Geographers 77:118–125.

    DOI: 10.1111/j.1467-8306.1987.tb00149.xSave Citation »Export Citation »

    This work, conducted in collaboration with the US Environmental Protection Agency regional offices, other federal agencies, state resource management agencies, and groups from neighboring North American countries, refines and subdivides ecoregions stratifed by response to disturbance.

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  • Peel, M. C., B. L. Finlayson, and T. A. McMahon. 2007. Updated world map of the Köppen-Geiger climate classification. Hydrology and Earth System Sciences 11:1633–1644.

    DOI: 10.5194/hess-11-1633-2007Save Citation »Export Citation »

    Provides good description of the climate classification system originally developed, and the update based on thousands of weather stations located globally. The maps are available online as an additional resource.

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Defining and Describing North American Biomes

A number of efforts have been conducted to map and delineate the biomes of North America. The classic text on the ecology of the biomes by Shelford 1963 and mapping of potential natural vegetation in the conterminous United States by Küchler 1964 continue to be widely cited. The classic map of biomes in Mexico is provided by Leopold 1950. Locations of paleo-biomes have been mapped by Williams, et al. 2004 and Dyke 2005 using pollen and tree-ring data.

  • Dyke, Arthur S. 2005. Late Quaternary vegetation history of Northern North America based on pollen, macrofossil, and faunal remains. Géographie physique et Quaternaire 59:211–262.

    DOI: 10.7202/014755arSave Citation »Export Citation »

    Biome maps spanning the interval from the last glacial maximum to modern times are presented for northern North America.

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  • Küchler, A. W. 1964. Potential natural vegetation of the conterminous United States. Special Publication 36. New York: American Geographic Society.

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    Considered by many to be the authority on the geographic distribution of potential natural vegetation in the United States (pre-European settlement).

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  • Leopold, Aldo S. 1950. Vegetation zones of Mexico. Ecology 31:507–518.

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    The classification and map of Mexican vegetation was compiled as part of a survey of the wildlife resources of Mexico.

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  • Williams, John W., Bryan N. Shuman, Thompson Webb III, Patrick J. Bartlein, and Phillip L. Leduc. 2004. Late-quaternary vegetation dynamics in North America: Scaling from taxa to biomes. Ecological Monographs 74:309–334.

    DOI: 10.1890/02-4045Save Citation »Export Citation »

    Using a fossil-pollen data set of over seven hundred sites, late-Quaternary vegetation history in northern and eastern North America across levels of ecological organization is used to show that biomes are dynamic entities that have changed in distribution, composition, and structure over time.

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Modeling Potential Vegetation and Plant Species Distributions

Responses of biomes and, in some cases, ecosystems at smaller spatial extents to environmental drivers have been simulated using models, such as Neilson 1995 in response to climate and Bachelet, et al. 2003 in response to changing fire regimes and biogeochemistry. Geographical shifts in distributions of species in response to changing climate, such as trees by McKenney, et al. 2007, have also been conducted.

  • Bachelet, Dominique, Ronald P. Neilson, Thomas Hickler, et al. 2003. Simulating past and future dynamics of natural ecosystems in the United States. Global Biogeochemical Cycles 17:1045.

    DOI: 10.1029/2001GB001508Save Citation »Export Citation »

    Simulations of potential vegetation distribution, natural fire frequency, carbon pools, and fluxes are presented for two Dynamic Global Vegetation Models. Good example of linking vegetation dynamics to biogeochemical cycling for the conterminous United States.

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  • McKenney, Daniel W., John H. Pedlar, Kevin Lawrence, Kathy Campbell, and Michael F. Hutchinson. 2007. Potential impacts of climate change on the distribution of North American trees. BioScience 57:939–948.

    DOI: 10.1641/B571106Save Citation »Export Citation »

    Present-day climatic niches for 130 North American tree species are determined and then future distributions under climate change are predicted. Provides excellent example of potential shifts in geographic distributions of individual tree species with climate change.

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  • Neilson, Ronald P. 1995. A model for predicting continental-scale vegetation distribution and water balance. Ecological Applications 5:362–385.

    DOI: 10.2307/1942028Save Citation »Export Citation »

    Describes a Mapped Atmosphere-Plant-Soil System (MAPSS) for simulating the potential biosphere impacts and biosphere-atmosphere feedbacks from climatic change with a focus on North America. One of the first biogeographic models of biomes developed for North America. Essential reading for large-scale simulation modeling.

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Mapping Land Use and Land Cover Classes

Maps of current vegetation are often developed using imagery from remote sensing products to show patterns in land use and land cover. Anderson, et al. 1976 provides a classification system for the United States, and Di Gregorio and Jansen 2005 provides a general classification system and software for use anywhere on Earth. Fry, et al. 2011 provides updated North American map, and Muchoney, et al. 2000 provides a similar map for Central America.

  • Anderson, James. R., Ernest E. Hardy, John T. Roach, and Richard E. Witmer, 1976. A land use and land cover classification system for use with remote sensor data. Geological Survey Professional Paper 964. Washington, DC: US Government Printing Office.

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    The framework of a national land use and land cover classification system is presented for use with remote sensing data. Three hierarchical levels of classifying land cover and land use are developed with level three representing over one hundred land-use types. This was an early attempt at classifying biomes using satellite-based imagery and can be used to compare with more recent approaches that use imagery at finer spatial resolutions.

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  • Di Gregorio, A., and L. J. Jansen. 2005. Land cover classification system, classification concepts and user manual software version 2. Environmental and Natural Resources Series 8. Rome: Food and Agriculture Organization, United Nations.

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    The Land Cover Classification System is a comprehensive, standardized a priori classification system designed to meet specific user requirements and created for mapping exercises, independent of the scale or means used to map. It enables a comparison of land cover classes regardless of data source, thematic discipline or country. Although developed for biomes globally, the North American maps are also useful.

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  • Fry, Joyce A., George Xian, Suming Jin, et al. 2011. Completion of the 2006 national land cover database for the conterminous United States. Photogrammetric Engineering and Remote Sensing 77:858–864.

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    This product provides not only the land cover characterization for 2006 but also land cover change from 2001 to 2006 for every 30 m grid cell in the United States. This database is particularly useful for examining changes in land use within biomes through time.

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  • Muchoney D., J. Borak, H. Chi, et al. 2000. Application of the MODIS global supervised classification model to vegetation and land cover mapping of Central America. International Journal of Remote Sensing 21:1115–1138.

    DOI: 10.1080/014311600210100Save Citation »Export Citation »

    The product is a map of land cover characterization in Central America at the spatial resolution of 1 km.

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Animal Species Distributions

Although biome studies have traditionally focused on plants and vegetation, the distribution of animal species have been quantified and mapped within North American biomes. In Central America, Almendra and Rogers 2012 provides maps of mammals, and Howell and Webb 1995 presents distributions of birds. Venter, et al. 2006 discusses threats to endangered species in Canada. Fleming 1973 presents distributions of mammals in North and Central forests, and Laliberte and Ripple 2004 compares historic and current ranges of carnivores and ungulates in North America.

  • Almendra, A. L., and D. S. Rogers. 2012. Biogeography of Central America mammals: Patterns and processes. In Bones, clones, and biomes: The history and geography of recent Neotropical mammals. Edited by B. D. Patterson and L. P. Costa. Chicago: Univ. of Chicago Press.

    DOI: 10.7208/chicago/9780226649214.001.0001Save Citation »Export Citation »

    Description of Central American mammal biodiversity within a biogeographic context.

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  • Fleming, Theodore H. 1973. Numbers of mammal species in North and Central American forest communities. Ecology 54.3: 555–563.

    DOI: 10.2307/1935340Save Citation »Export Citation »

    Latitudinal trends in the species diversity of North and Central America mammalian communities are documented for habitats ranging from the tundra of Alaska to the moist tropical forests of Panama. Although this paper focuses on mammals in forested biomes, the trends in diversity are useful for comparing across biomes.

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  • Howell, Steve N. G., and Sophie Webb. 1995. A guide to the birds of Mexico and northern Central America. New York: Oxford Univ. Press.

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    This book describes the spatial distribution of approximately 1,070 species of birds in Mexico and northern Central America.

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  • Laliberte, Andrea S., and William J. Ripple. 2004. Range contractions of North American carnivores and ungulates. BioScience 54:123–138.

    DOI: 10.1641/0006-3568(2004)054[0123:RCONAC]2.0.CO;2Save Citation »Export Citation »

    This paper compares the historic and current geographical ranges of forty-three North American carnivores and ungulates. The approach is particularly useful in identifying large-scale patterns in range contractions and expansions that can be compared among biomes.

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  • Venter, Oscar, Nathalie N. Brodeur, Leah Nemiroff, Brenna Belland, Ivan J. Dolinsek, and James W. A. Grant. 2006. Threats to endangered species in Canada. BioScience 56:903–910.

    DOI: 10.1641/0006-3568(2006)56[903:TTESIC]2.0.CO;2Save Citation »Export Citation »

    Threats facing 488 species in Canada were quantified and categorized as extinct, extirpated, endangered, threatened, or of special concern.

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Patterns in Climate, Atmospheric Chemistry, and Soils

Environmental drivers vary spatially across the North American continent. Patterns in climate are provided by Bonsal, et al. 2001 for Canada, by Bullock 1986 for southern Mexico, and by Daly, et al. 2008 for the United States. Effects of human activities on a climate signal are discussed in Karoly, et al. 2003. Patterns in nitrogen emissions and deposition for the United States are shown in Fenn, et al. 2003, and Foth and Schafer 1980 and Liu, et al. 2013 provide a unified soil map for North America.

  • Bonsal, B. R., X. Zhang, L. A. Vincent, and W. D. Hogg. 2001. Characteristics of daily and extreme temperatures over Canada. Journal of Climate 14:1959–1976.

    DOI: 10.1175/1520-0442(2001)014%3C1959:CODAET%3E2.0.CO;2Save Citation »Export Citation »

    This paper examines trends and variability in daily minimum and maximum temperature with particular emphasis on extremes for the periods of 1900–1998 (southern Canada), and 1950–1998 (the entire country). These trends in temperature can easily be compared across biomes within Canada.

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  • Bullock, S. H. 1986. Climate of Chamela, Jalisco, and trends in the south coastal region of Mexico. Archives for Meteorology, Geophysics, and Bioclimatology, Series B 36:297–316.

    DOI: 10.1007/BF02263135Save Citation »Export Citation »

    This paper describes the warm, seasonally dry climate of Mexico’s south coast using data from the biological station at Chamela, Jalisco, and comparing it with twenty-six other localities in the region. This data set is regionally important for the southern coast of Mexico.

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  • Daly, Christopher, Michael Halbleib, Joseph I. Smith, et al. 2008. Physiographically-sensitive mapping of temperature and precipitation across the conterminous United States. International Journal of Climatology 28:2031–2064.

    DOI: 10.1002/joc.1688Save Citation »Export Citation »

    The PRISM modeling system is described where 1971–2000 long-term averages were simulated, and an uncertainty analysis was conducted for the United States. PRISM model results for monthly temperature and precipitation are also available online and are used by many ecologists to characterize the climatic regime of specific locations or to compare across large extents.

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  • Fenn, Mark E., Richard Haeuber, Gail S. Tonnesen, et al. 2003. Nitrogen emissions, deposition, and monitoring in the western United States. BioScience 53:391–403.

    DOI: 10.1641/0006-3568(2003)053[0391:NEDAMI]2.0.CO;2Save Citation »Export Citation »

    Nitrogen deposition is compared for sites distributed across the western United States using data from long-term monitoring sites. These data from the National Atmospheric Deposition Program are a valuable resource for patterns in atmospheric chemistry and trends through time.

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  • Foth, Henry D., and John W. Schafer. 1980. Soil geography and land use. New York: Wiley.

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    Distribution of major soil orders are presented for North America.

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  • Karoly, David. J., Karl Braganza, Peter A. Stott, et al. 2003. Detection of a human influence on North American climate. Science 302:1200–1203.

    DOI: 10.1126/science.1089159Save Citation »Export Citation »

    Variation in surface temperature data over the 20th century are examined as an indicator of climate change.

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  • Liu, S., Y. Wei, W. M. Post, R. B. Cook, K. Schaefer, and M. M. Thornton. 2013. The Unified North American Soil Map and its implication on the soil organic carbon stock in North America. Biogeosciences 10:2915–2930.

    DOI: 10.5194/bg-10-2915-2013Save Citation »Export Citation »

    The Unified North American Soil Map was developed by combining information from US STATSGO2 and Soil Landscape of Canada databases and filling gaps with the Harmonized World Soil Database version 1.21. Patterns in soils show variation within biomes that can be sued to help explain fine-scale patterns in vegetation and plant and animal species distributions.

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Patterns in Human Activities

Human activities vary spatially across the biome with consequences for vegetation dynamics within biomes. A number of texts have presented information on the paleo-history of these activities, including Piperno 2006 for Central America. Current patterns in human activities, including urbanization and pollution, are provided in Grimm, et al. 2008. Studies of coupled human-natural systems are prevalent, including comparisons of biomes in the book edited by Redman and Foster 2008, and the use of long-term data from multiple locations to address biome-level questions, as presented by Robertson, et al. 2012.

  • Grimm, Nancy B., David Foster, Peter Groffman, et al. 2008. The changing landscape: Ecosystem responses to urbanization and pollution across climatic and societal gradients. Frontiers in Ecology and the Environment 6:264–272.

    DOI: 10.1890/070147Save Citation »Export Citation »

    Shows continental-scale patterns in urbanization and pollution driven by land-use change across the biome. Essential reading to understand broad-scale patterns in human activities across biomes.

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  • Piperno, Dolores R. 2006. Quaternary environmental history and agricultural impact on vegetation in Central America. Annals of the Missouri Botanical Garden 93:274–296.

    DOI: 10.3417/0026-6493(2006)93[274:QEHAAI]2.0.CO;2Save Citation »Export Citation »

    Historical data from lake sediments are used to relate climate, vegetation, and human land use of the lowland Central American tropical forest between ca. 20,000 BP and the time of European contact. Excellent historical account of these drivers of patterns in vegetation.

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  • Redman, Charles L., and David R. Foster, eds. 2008. Agrarian landscapes in transition. Oxford: Oxford Univ. Press.

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    Unique resource for comparing coupled human-natural ecosystems in six biomes based on site-specific studies. Although each has its own unique agricultural history, the sites characterizing biomes show similar patterns in human impacts.

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  • Robertson, G. Philip, Scott L. Collins, David R. Foster, et al. 2012. Long-term ecological research in a human-dominated world. BioScience 62:342–353.

    DOI: 10.1525/bio.2012.62.4.6Save Citation »Export Citation »

    Conceptual paper that describes how long-term research in multiple biomes can be used to address emergent continent-scale questions that couple human and natural ecosystems.

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Biogeography of Biomes

Biomes have been classified in numerous ways by different authors. Gurevitch, et al. 2006 is one example of this adoption with some modifications. Olson, et al. 2001 classification of major habitat types is similar to biome types, but with more classes based on conservation issues. Barbour and Billings 2000 classes are based on vegetation type that includes locally important vegetation at a finer resolution than biomes considered by other authors. In the sections that follow, nine major biomes are described in more detail that generally follow the Whittaker 1975 and Gurevitch, et al. 2006 classifications based on temperature and moisture but with fewer classes based on physiognomy of the plants. Along a temperature gradient, the hottest forests are either wet year-round (tropical evergreen, broad-leaved rainforest) or seasonally dry (tropical broad-leaved deciduous forest). As annual average temperature decreases, cool, wet locations are dominated by temperate coniferous forests, and seasonally cold, moderately wet locations are dominated by temperate deciduous broad-leaved forests. As average temperature continues to decrease, boreal evergreen forests are found in cold, moderately wet locations and the coldest, moderately wet or dry locations are dominated by tundra (grasses, sedges, shrublands). Hot or mild locations that are dry are dominated by deserts (shrubs, succulents, grasses), and seasonally warm, moderately wet locations are dominated by temperate grasslands. Mild locations that are seasonally wet or dry are dominated by Mediterranean woodlands (Chaparral) with evergreen sclerophylous leaves.

Tropical Rainforest

Tropical rainforests are among the most diverse and productive biomes on Earth. These forests are found within a band extending from 10oN to 10oS latitude at elevations below 1 km. Rainfall occurs year-round with amounts totaling more than 250 cm each year. In North America, tropical rainforests mainly occur in Central America and around the Caribbean. The first general account of this forest type was Richards 1952. Hypotheses to explain high species richness are explored in Denslow 1987. Deforestation and the associated loss of biodiversity are an important conservation concern described in Achard, et al. 2002; Geist and Lambin 2002; and Myers, et al. 2000. However, Parés-Ramos, et al. 2008 and Wright and Samaniego 2008 show that recent demographic and economic development has reduced rates of deforestation or led to forest expansion in some areas in the Latin America.

Tropical Deciduous Forest

Tropical deciduous forests, also known as tropical dry forests, are described by Murphy and Lugo 1986 as defined by four factors: (i) occur in frost-free zones; (ii) the mean annual temperature must be higher than 17°C; (iii) the mean annual precipitation is between 250 and 2000 mm; and (iv) the annual ratio of potential evapotranspiration to precipitation exceeds 1.0. The impacts of major disturbances, such as hurricanes and drought, on forest ecosystem structure and dynamics are documented in Dirzo, et al. 2011 and Brokaw, et al. 2012. Studies of biodiversity are common, such as Bullock and Arturo Solis-Magallanes 1990. Due to their low rainfall and high humidity, humans have developed large population centers in tropical dry forests, as described in Tosi and Voertman 1964.

  • Brokaw, Nicholas, Todd A. Crowl, Ariel E. Lugo, William H. McDowell, and Frederick N. Scatena, eds. 2012. A Caribbean forest tapestry: The multidimensional nature of disturbance and response. Oxford: Oxford Univ. Press.

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    Excellent background on tropical ecosystem dynamics over multiple scales of time and space with a focus on long-term research.

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  • Bullock, Stephen H., and J. Arturo Solis-Magallanes. 1990. Phenology of canopy trees of a tropical deciduous forest in Mexico. Biotropica 22:22–35.

    DOI: 10.2307/2388716Save Citation »Export Citation »

    Example of biodiversity study using phenology of 108 species (1,094 trees) monitored for forty-two months in a tropical deciduous forest in the Pacific lowlands of Mexico.

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  • Dirzo, R., H. S. Young, H. A. Mooney, and G. Ceballos, eds. 2011. Seasonally dry tropical forests, ecology and conservation. Washington, DC: Island.

    DOI: 10.5822/978-1-61091-021-7Save Citation »Export Citation »

    Essential reading on a broad range of subjects including distribution of tropical dry forests, biodiversity of flora and fauna (from community to genetic level), physiological ecology, plant animal interactions, ecosystem processes, and conservation biology.

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  • Murphy, Peter G., and Ariel E. Lugo. 1986. Ecology of tropical dry forest. Annual Review of Ecology and Systematics 17:67–88.

    DOI: 10.1146/annurev.es.17.110186.000435Save Citation »Export Citation »

    Essential reading as general overview of tropical dry forests, their definition, climate, vegetation and human interactions.

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  • Tosi, Joseph A., and Robert F. Voertman. 1964. Some environmental factors in the economic development of the tropics. Economic Geography 40:189–205.

    DOI: 10.2307/142421Save Citation »Export Citation »

    Good discussion of environmental factors that dictated human settlement and development in dry forests of the tropics.

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Temperate Deciduous Forest

Temperate deciduous forests described by Braun 1950 make up a large portion of the eastern United States and parts of Canada between latitudes 23.5°N and 66.5°N. The vegetation is primarily deciduous hardwood species, with conifers dominating the colder climates and/or areas with less fertile soils. The use of these forests by humans and influence by natural disturbance over the past one thousand years are described using long-term data in the edited volume by Foster and Aber 2006. Comparisons with other forest biomes in North America are provided by Kuennecke 2008. Whittaker 1957 was instrumental in ecology by showing that temperate deciduous forests follow the Gleasonian individualistic concept rather than a Clementsian superorganism paradigm. The global overview of these forests is provided by Gilliam 2012.

  • Braun, E. Lucy. 1950. Deciduous forests of eastern North America. Philadelphia: Blakiston.

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    Classic monograph that set the standard for vegetation science in North America and elsewhere. Essential background reading.

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  • Foster, David R., and John D. Aber, eds. 2006. Forests in time: The environmental consequences of 1,000 years of change in New England. New Haven, CT: Yale Univ. Press.

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    This seminal book, based on innovative research at Harvard Forest, describes the dramatic natural and human-induced changes in the land and environment of New England over the past one thousand years.

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  • Gilliam, Frank S. 2012. “Temperate deciduous forests.” In Oxford Bibliographies: Ecology.

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    Excellent overview of the temperate deciduous forest biome globally.

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  • Kuennecke, Bernd. 2008. Temperate forest biomes. Westport, CT: Greenwood.

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    This volume compares temperate deciduous forests with boreal forests and Mediterranean woodland and scrub in North America. Useful for general overview as well as details on each biome.

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  • Whittaker, Robert H. 1957. Recent evolution of ecological concepts in relation to the eastern forests of North America. American Journal of Botany 44:197–206.

    DOI: 10.2307/2438311Save Citation »Export Citation »

    Classic paper that draws upon the author’s original work in the southern Appalachian Mountains to support Gleason’s individualistic concept of communities rather than the Clementsian paradigm of a super-organism.

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Temperate Coniferous Forest

Temperate coniferous forests of North America, as described by Waring and Franklin 1979, are dominated by coniferous trees, usually evergreens. The climate is characterized by warm summers and cool winters. Here, temperate coniferous forest includes the coniferous forests in the temperate North America: (i) along the Pacific northwestern coast of the continent, (ii) in the Sierra Nevada of California, (iii) in the mountainous areas between the Sierra Nevada and the Rocky Mountains, and (iv) in the southeastern United States. In the Pacific Northwest, these forests can sustain among the highest levels of biomass of any terrestrial biome and are notable for massive trees; biodiversity is described by Franklin 1988. In addition, old growth forests within the biome are particular important, as described by Spies and Duncan 2009. Fire has been an important factor in shaping the structure and function of these forest ecosystems based on Agee 1996. Land management practices (e.g., fire suppression, selective logging, and livestock grazing) and changes in climate have contributed to increased wildfire activities in recent decades, as discussed by Belsky and Blumenthal 1997, and Westerling, et al. 2006. Proper management of these fire-prone forests for log productivity and conservation purpose has been frequently studied, including Fontaine and Kennedy 2012 and Noss, et al. 2006.

  • Agee, J. K. 1996. Fire ecology of Pacific Northwest forests. Washington, DC: Island.

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    Provides good overviews of wildland fire history and the effects of fire on maintaining forests of the Pacific Northwest. The knowledge in fire ecology is applicable to other forests.

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  • Belsky, A. J., and D. M. Blumenthal. 1997. Effects of livestock grazing on stand dynamics and soils in upland forests of the Interior West. Conservation Biology 11:315–327.

    DOI: 10.1046/j.1523-1739.1997.95405.xSave Citation »Export Citation »

    Excellent review of the dynamics and factors driving the dynamics of coniferous forests before and after Euro-American settlement, with a focus on effects of livestock grazing.

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  • Fontaine, Joseph B., and Patricia L. Kennedy. 2012. Meta-analysis of avian and small-mammal response to fire severity and fire surrogate treatments in US fire-prone forests. Ecological Applications 22:1547–1561.

    DOI: 10.1890/12-0009.1Save Citation »Export Citation »

    Quantitatively synthesizes bird and small animal demographic responses to burn severity, fire surrogates (forest thinning), and combined treatments in fire-prone forests of the United States.

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  • Franklin, Jerry F. 1988. Structural and functional diversity in temperate forests. Biodiversity. Washington, DC: National Academy Press.

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    Chapter on the diversity of temperate forests (pp. 166–175) is good background.

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  • Noss, R. F., J. F. Franklin, W. L. Baker, T. Schoennagel, and P. B. Boyle. 2006. Managing fire-prone forests in the western United States. Frontiers in Ecology and the Environment 4:418–487.

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    Reviews the ecological science relevant to developing and implementing fire and fuel management policies for forests before, during, and after wildfires. Although not limited to coniferous forests, the management principles developed apply to this forest biome.

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  • Spies, T. A., and S. L. Duncan, eds. 2009. Old growth in a new world: A Pacific Northwest icon reexamined. Washington, DC: Island.

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    Show the importance of old growth forests as a valuable system to be protected and restored.

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  • Waring, R. H., and J. F. Franklin. 1979. Evergreen coniferous forests of the Pacific Northwest. Science 204:1380–1386.

    DOI: 10.1126/science.204.4400.1380Save Citation »Export Citation »

    Describe the evergreen coniferous forests of the Pacific Northwest in detail. Essential background reading.

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  • Westerling, A. L., H. G. Hidalgo, D. R. Cayan, and T. W. Swetnam. 2006. Warming and earlier spring increase Western U.S. Forest wildfire activity. Science 313:940–943.

    DOI: 10.1126/science.1128834Save Citation »Export Citation »

    A review paper shows that recent climatic trends are strongly associated with increased wildfire activity in western US forests. Although not limited to temperate coniferous forests, this is an excellent review of fire activity as climate changes.

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Boreal Forest

Boreal forests, also known as “taiga,” as described by Larson 1980, are dominated by coniferous trees and found at high latitudes of Alaska and Canada with short cool summers and long cold winters. Boreal forests dynamics, as described by Chapin, et al. 2006, are driven by wildfire and other disturbances interacting with global warming. High-intensity fires are described by Bond-Lamberty, et al. 2004 as common occurrences with effects on biomass accumulation and primary production. An important ecosystem service provided by boreal forests, as described by Kasischke, et al. 1995, is carbon storage, although climate change threatens this storage as temperature increases with global warming. Snyder, et al. 2004 provides evidence that boreal forests can feed back to influence air temperature. An overview of boreal forests globally is provided by Frelich 2013.

Tundra

The tundra biome is located in North America in the arctic at high latitudes (in the northern parts of Alaska and Canada) and in the alpine at high elevations in the Rocky Mountains. The alpine tundra is located above tree line where the short statured vegetation has adapted to harsh climatic conditions (cold temperature, high winds, frozen soils). The tundra biome is described in general by Billings and Mooney 1968 and updated by Quinn 2008. A more detailed description of the arctic tundra is provided by Bliss, et al. 1973, with updates by Hobbie and King 2014, and of alpine systems in the Rocky Mountains based on long-term data in the edited volume Bowman and Seastedt 2001. The arctic tundra is expected to be sensitive to global change based on research by Chapin, et al. 1995 and others. Human activities that threaten the Rocky Mountain tundra are described by Willard and Marr 1970.

  • Billings, W. D., and H. A. Mooney. 1968. The ecology of arctic and alpine plants. Biological Reviews 43:481–529.

    DOI: 10.1111/j.1469-185X.1968.tb00968.xSave Citation »Export Citation »

    Geographic extent and general characteristics of arctic and alpine vegetation with a focus on adaptations of plants and their primary productivity.

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  • Bliss, L. C., G. M. Courtin, D. L. Pattie, R. R. Riewe, D. W. A. Whitfield, and P. Widden. 1973. Arctic tundra ecosystems. Annual Review of Ecology and Systematics 4:359–399.

    DOI: 10.1146/annurev.es.04.110173.002043Save Citation »Export Citation »

    Review paper that describes role of biotic and abiotic factors affecting arctic tundra.

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  • Bowman, William D., and Timothy R. Seastedt, eds. 2001. Structure and function of an alpine ecosystem: Niwot Ridge, Colorado. Oxford: Oxford Univ. Press.

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    Essential reading for overview of alpine dynamics based on long-term research conducted at the Niwot Ridge Long Term Ecological Research site.

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  • Chapin, F. Stuart, III, Gaius R. Shaver, Anne E. Giblin, Knute J. Nadelhoffer, and James A. Laundre. 1995. Responses of arctic tundra to experimental and observed changes in climate. Ecology 76:694–711.

    DOI: 10.2307/1939337Save Citation »Export Citation »

    One paper of many describing an experimental study to determine how changes in light, temperature, and nutrients can affect community and ecosystem changes in arctic tundra.

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  • Hobbie, John E., and George W. King. 2014. Alaska’s changing Arctic: Ecological consequences for tundra, streams, and lakes. Oxford: Oxford Univ. Press.

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    Essential reading for background on arctic tundra processes and dynamics through the use of long-term data at the Toolik Lake Long Term Ecological Research site. The book includes linkages among the tundra and major aquatic systems (lakes, streams).

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  • Quinn, Joyce A. 2008. Arctic and alpine biomes. Westport, CT: Greenwood.

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    This volume covers vegetation, geographic distribution, and soil properties of tundra, and includes challenges posed by the environment, adaptations of plants and animals, and conservation efforts.

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  • Willard, Beatrice E., and John W. Marr. 1970. Effects of human activities on alpine tundra ecosystems in Rocky Mountain National Park, Colorado. Biological Conservation 2:257–265.

    DOI: 10.1016/0006-3207(70)90008-XSave Citation »Export Citation »

    Discusses effects of human activities on the alpine tundra.

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Temperate Grassland

Grasslands cover more than 12 percent of North America and occur in regions that receive too little rain to support trees and too much rain to be deserts. Grasslands are fertile lands that are biologically diverse and are heavily used for agriculture (farming and livestock). The grassland biome is defined by the presence or dominance by native grasses. Grasslands of North America are described as individual chapters in Coupland 1979 and updated in more recent volumes for the shortgrass steppe by Lauenroth and Burke 2008, for tallgrass prairie by Knapp, et al. 1998, and summarized by Gibson 2012. Many of the classic ecological studies of the early 20th century were conducted in grasslands, including the work by Clements 1916 and the author’s students and colleagues, for example, Weaver 1954.

  • Clements, Frederic E. 1916. Plant succession: An analysis of the development of vegetation. Washington, DC: Carnegie Institution of Washington.

    DOI: 10.5962/bhl.title.56234Save Citation »Export Citation »

    Classic book that describes Clementsian view of succession. Good historic account of ideas in grasslands at turn of century.

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  • Coupland, Robert T., ed. 1979. Grassland ecosystems of the world: Analysis of grasslands and their uses. IBP Synthesis Series 18. Cambridge, UK: Cambridge Univ. Press.

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    An edited book of thirty-three contributions summarizing grassland ecosystem research conducted as part of the International Biology Program. Includes chapters on each grassland type in North America.

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  • Gibson, David J. 2012. “Grassland Biome.” In Oxford Bibliographies: Ecology.

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    Excellent annotated bibliography of the grassland biome. Essential reading for characteristics of grasslands and current issues.

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  • Knapp, Alan K., John M. Briggs, David C. Hartnett, and Scott L. Collins, eds. 1998. Grassland dynamics: Long-term ecological research in tallgrass prairie. New York: Oxford Univ. Press.

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    This volume provides a comprehensive perspective on the tallgrass prairie of North America based on long-term research from the Konza Long Term Ecological Research Program.

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  • Lauenroth, William K., and Ingrid C. Burke, eds. 2008. Ecology of the shortgrass steppe: A long-term perspective. New York: Oxford Univ. Press.

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    A summary of more than fifty years of research on shortgrass steppe in central North America.

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  • Weaver, John E. 1954. North American prairie. Lincoln, NE: Johnsen.

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    Comprehensive book that summarizes much of Weaver’s work in the central Great Plains.

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Desert

Deserts can be either hot or cold, but all deserts receive very little precipitation (less than 250 mm of precipitation per year). McClaran and Van Devender 1995 provides excellent overview of deserts, and Whitford 2002 provides an updated, ecosystem perspective. The southwestern deserts of North America (Chihuahuan, Sonoran, Mojave) are hot and the intermountain desert (Great Basin) is cold. Rain falls in seasons specific to each desert that determine population, community and ecosystem dynamics, as summarized in Havstad, et al. 2006 for the Chihuahuan Desert; Rundel and Gibson 1996 for the Mojave Desert; and Phillips and Comus 1999 for the Sonoran Desert. Schlesinger, et al. 1990 describes deserts as highly dynamic in response to livestock overgrazing and drought that can shift grasslands to desertified shrublands or savannas, described by McPherson 1997. Peters, et al. 2006 provides a more recent perspective that includes changes in patterns and processes as the spatial extent of the desertified area increases. Ward 2012 provides global overview of the desert biome.

Mediterranean Woodlands “Chaparral”

The Mediterranean woodlands of North America are known as the “chaparral,” due to its sclerophyllous shrub vegetation. The North American Chaparral is concentrated in the western United States as well as parts of Mexico. The Chaparral is characterized by a Mediterranean-like climate defined by cool, wet winters and hot, dry summers. The Chaparral, similar to other Mediterranean woodlands globally, has high plant diversity, as described by Cowling, et al. 1996. Fire is a commonly occurring disturbance that has been well-studied in California by Keeley, et al. 2005; Keeley, et al. 2006; and others.

  • Cowling, Richard M., Philip W. Rundel, Byron B. Lamont, Mary Kalin Arroyo, and Margarita Arianoutsou. 1996. Plant diversity in Mediterranean-climate regions. Trends in Ecology and Evolution 11:362–366.

    DOI: 10.1016/0169-5347(96)10044-6Save Citation »Export Citation »

    Essential reading for review of patterns and determinants of local and regional plant diversity in Mediterranean climate regions.

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  • Keeley, Jon E., Melanie Baer-Keeley, and C. J. Fotheringham. 2005. Alien plant dynamics following fire in Mediterranean-climate California shrublands. Ecological Applications 15:2109–2125.

    DOI: 10.1890/04-1222Save Citation »Export Citation »

    This study monitored alien plants the first five years after fire in southern California chaparral.

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  • Keeley, Jon E., C. J. Fotheringham, and Melanie Baer-Keeley. 2006. Demographic patterns of postfire regeneration in Mediterranean-climate shrublands of California. Ecological Monographs 76:235–255.

    DOI: 10.1890/0012-9615(2006)076[0235:DPOPRI]2.0.CO;2Save Citation »Export Citation »

    Following massive wildfires in southern California, this paper reports on detailed demographic data obtained from sampling chaparral and sage scrub vegetation from ninety sites for five postfire years.

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Connections Among Biomes within North America

Biomes within North America can be connected by a number of vectors of transport, including water, wind, animals (including humans), and pests and pathogens, which are described in Peters, et al. 2008. These vectors are influenced by atmospheric (e.g., climate, air chemistry) and physical drivers, such as wildfire, topography, and soils, that interact with each other and with the vegetation to determine the rate, direction, and magnitude of movement of materials and organisms that affect the dynamics of each biome. There are a number of books and papers that provide overviews of the climate system that connects biomes, including the El Niño Southern Oscillation (ENSO) globally, including Allan, et al. 1996.

  • Allan R., J. Lindesay, and D. Parker. 1996. El Niño: Southern oscillation and climate variability. Collingwood, Australia: CSIRO.

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    Good general description of the climate system and the ENSO based on information known in 1996.

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  • Peters, Debra P. C., Peter M. Groffman, Knute J. Nadelhoffer, et al. 2008. Living in an increasingly connected world: A framework for continental-scale environmental science. Frontiers in Ecology and the Environment 5:229–237.

    DOI: 10.1890/070098Save Citation »Export Citation »

    Essential reading for overview of continental-scale patterns in climate change drivers and ecological responses.

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Climatic Drivers as Vectors of Transport that Connect Biomes

Spatial variation in sea surface temperatures at multiple temporal scales can synchronize climatic patterns in drought or wet periods across multiple biomes, as described by Trenberth and Guillemot 1996, or can lead to spatial variation in climate based on research by Hu and Feng 2012. Veblen, et al. 2003 shows that continental-scale patterns in drought and modes of sea surface temperatures can generate variation among biomes in fire regimes and fire impacts. The relationship between climate and streamflow for multiple biomes is explored in Jones, et al. 2012, and climate with stream chemistry is discussed for nitrogen by Preston, et al. 2011 and for phosphorous by Alexander, et al. 2004. Barry 1998 shows how major floods are another driver that connect large areas and multiple biomes, and Knox 1993 provides evidence that changes in climate are expected to lead to changes in flood regimes in the future.

  • Alexander, Richard B., Richard A. Smith, and Gregory E. Schwarz. 2004. Estimates of diffuse phosphorus sources in surface waters of the United States using a spatially referenced watershed model. Water Science Technology 49:1–10.

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    The watershed model SPARROW (SPAtially Referenced Regression On Watershed attributes) provides an excellent example of an approach to estimate the sources and transport of total phosphorus in surface waters across the United States.

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  • Barry, John M. 1998. Rising tide: The Great Flood of 1927 and how it changed America. New York: Simon & Schuster.

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    Describes Mississippi River flood of 1927 and shows how the river connected different locations and their biomes across the United States.

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  • Hu, Qi, and Song Feng. 2012. AMO- and ENSO-driven summertime circulation and precipitation variations in North America. Journal of Climate 25:6477–6495.

    DOI: 10.1175/JCLI-D-11-00520.1Save Citation »Export Citation »

    Description of the relationship between the Atlantic multi-decadal oscillation and ENSO, including their resulting inter-annual to multi-decadal time-scale variations, in affecting summertime precipitation for biomes in North America.

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  • Jones, Julia A., Irena F. Creed, Kendra L. Hatcher, et al. 2012. Ecosystem processes and human influences regulate streamflow response to climate change at long-term ecological research sites. BioScience 62:390–404.

    DOI: 10.1525/bio.2012.62.4.10Save Citation »Export Citation »

    Compares long-term records at thirty-five headwater basins in the United States and Canada from a number of biomes. Because these sampled streams are reference basins in natural areas, the effects of connectivity with nearby or upstream ecosystems is indicated.

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  • Knox, James C. 1993. Large increases in flood magnitude in response to modest changes in climate. Nature 361:430–432.

    DOI: 10.1038/361430a0Save Citation »Export Citation »

    A seven thousand-year geological record of overbank floods for upper Mississippi river tributaries is used to show that small, rare floods occurred during warmer, drier climate whereas frequent, larger floods occurred in cooler and wetter climate. Can be used to estimate potential changes in this river in the future as climate changes.

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  • Preston, Stephen D., Richard B. Alexander, Gregory E. Schwarz, and Charles G. Crawford. 2011. Factors affecting stream nutrient loads: A synthesis of regional SPARROW model results for the continental United States. Journal of American Water Resources Association 47:891–915.

    DOI: 10.1111/j.1752-1688.2011.00577.xSave Citation »Export Citation »

    Twelve regional SPARROW models covering most of the continental United States were used to evaluate regional differences in factors affecting stream nutrient loads. Urban and agricultural sources influenced stream nutrient loads nationally and regionally. Because these regions are in different biomes, the biomes can be compared for differences in stream nutrient loads.

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  • Trenberth, Kevin E., and Christian J. Guillemot. 1996. Physical processes involved in the 1988 drought and 1993 floods in North America. Journal of Climate 9:1288–1298.

    DOI: 10.1175/1520-0442(1996)009%3C1288:PPIITD%3E2.0.CO;2Save Citation »Export Citation »

    An analysis of the spring–summer 1988 drought and 1993 floods over North America as related to strong La Niña conditions in 1988 and a mature El Niño in 1993. Provides background to drivers of seasonal precipitation in different biomes.

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  • Veblen, Thomas T., William L. Baker, Gloria Montenegro, and Thomas W. Swetnam, eds. 2003. Fire and climatic change in temperate ecosystems of the western Americas. New York: Springer-Verlag.

    DOI: 10.1007/b97443Save Citation »Export Citation »

    Fifteen chapters describe fire regimes and climate change primarily for woody plant-dominated biomes in North and South America. Good within biome information about fire and comparisons among biomes are possible using information from multiple chapters.

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Dust and Aerosols as Vectors of Transport that Connect Biomes

Dust deposition driven by wind can link alpine lakes to low elevation, overgrazed rangelands, described by Neff, et al. 2008, with consequences for duration of snow cover and earlier spring runoff in rivers described by Painter, et al. 2010. Fenn, et al. 2003 shows patterns in atmospheric chemistry with a focus on nitrogen have the potential to link biomes or landscapes within a biome. Driscoll, et al. 2001 shows how patterns in acid rain can vary across the biome as a result of the source of the chemicals, and Likens 2010 provides background to acid rain regulation by the US federal government.

  • Driscoll, Charles T., Gregory B. Lawrence, Arthur J. Bulger, et al. 2001. Acidic deposition in the northeastern United States: Sources and inputs, ecosystem effects, and management strategies. BioScience 51:180–198.

    DOI: 10.1641/0006-3568(2001)051[0180:ADITNU]2.0.CO;2Save Citation »Export Citation »

    Good general overview of acid rain with examples from northeastern United States.

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  • Fenn, Mark E., Jill S. Baron, Edith B. Allen, et al. 2003. Ecological effects of nitrogen deposition in the western United States. BioScience 53.4: 404–420.

    DOI: 10.1641/0006-3568(2003)053[0404:EEONDI]2.0.CO;2Save Citation »Export Citation »

    Good general description of nitrogen in the atmosphere, soils, and vegetation. Greatest acidification occurs in high alpine lakes during spring when sulfate and nitrates are released from melting snow pack. Air quality from lower elevations connects these high elevation streams and lakes with cities at lower elevations. The result is nitrogen saturation of plants and soils with potential consequences for ecosystem dynamics.

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  • Likens, Gene E. 2010. The role of science in decision-making: Does evidence-based science drive environmental policy? Frontiers in Ecology and the Environment 8:e1–e9.

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    Good historical review of acid rain science and policy in the United States beginning in the early 1960s to 2010 with a focus on recommendations for facilitating communication between scientists, policymakers, and the general public.

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  • Neff, J. C., A. P. Ballantyne, G. L. Farmer, N. M. Mahowald, et al. 2008. Increasing eolian dust deposition in the western US linked to human activity. Nature Geoscience 1:189–195.

    DOI: 10.1038/ngeo133Save Citation »Export Citation »

    Excellent background paper for understanding relationships between human activities and dust deposition. Alpine lake sediments in western United States in 19th century showed 500 percent increase in dust load levels above the late Holocene average following settlement by Europeans and expansion of livestock grazing in low elevation sites.

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  • Painter, Thomas H., Jeffrey S. Deems, Jayne Belnap, Alan F. Hamlet, Christopher C. Landry, and Bradley Udall. 2010. Response of Colorado River runoff to dust radiative forcing in snow. Proceedings of the National Academy of Sciences 107:17125–17130.

    DOI: 10.1073/pnas.0913139107Save Citation »Export Citation »

    Builds on work by Neff, et al. 2008 to show that peak runoff from Colorado River has occurred on average three weeks earlier under heavier dust loading and that evapotranspiration increases from earlier exposure of vegetation and soils decreases annual runoff.

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Animals as Vectors that Connect Biomes

Comparing prehistoric animal distributions from Lyons, et al. 2010 with predicted future distributions from Lawler, et al. 2009 and Lawler, et al. 2010 suggest that future faunal distributions may be quite different from present. Large-scale migrations of insects, including the Monarch butterfly based on information in Urquhart and Urquhart 1978, the gypsy moth from information in Haynes, et al. 2013, and birds from information in Robbins, et al. 1989 can connect different locations within one biome or different biomes to lead to increased susceptibility of animals to human activities year-round in different parts of their ranges. Krauss, et al. 2004 and Reisen 2013 show that birds are also effective transport vectors of viruses that can affect humans.

  • Haynes, Kyle J., Ottar N. Bjørnstad, Andrew J. Allstadt, and Andrew M. Liebhold. 2013. Geographical variation in the spatial synchrony of a forest-defoliating insect: Isolation of environmental and spatial drivers. Proceedings of the Royal Society B: Biological Sciences 280.1753: 2012–2373.

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    The importance of synchronous precipitation to spatial synchronization over long distances (approx. 900 km) of the outbreak dynamics of the gypsy moth is shown, an invasive species in North America. Excellent example of how precipitation synchronicity can influence invasive species dynamics to connect biomes.

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  • Krauss, Scott, David Walker, S. Paul Pryor, et al. 2004. Influenza A viruses of migrating wild aquatic birds in North America. Vector-Borne and Zoonotic Diseases 4:177–189.

    DOI: 10.1089/vbz.2004.4.177Save Citation »Export Citation »

    Surveillance of North America’s wild ducks and shorebirds for twenty-six and sixteen years, respectively, revealed differences in the prevalence of orthomyxoviruses between these hosts that migrate across large distances to connect biomes.

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  • Lawler, Joshua J., Sarah L. Shafer, Betsy A. Bancroft, and Andrew R. Blaustein. 2010. Projected climate impacts for the amphibians of the Western Hemisphere. Conservation Biology 24:38–50.

    DOI: 10.1111/j.1523-1739.2009.01403.xSave Citation »Export Citation »

    Three approaches used to map areas in the western hemisphere where amphibians are particularly likely to be affected by climate change show differential sensitivity of biomes to climate.

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  • Lawler, Joshua J., Sarah L. Shafer, Denis White, et al. 2009. Projected climate-induced faunal change in the Western Hemisphere. Ecology 90:588–597.

    DOI: 10.1890/08-0823.1Save Citation »Export Citation »

    Study showed the potential effects of thirty coupled atmosphere-ocean general circulation model future-climate simulations on the geographic ranges of 2,954 species of birds, mammals, and amphibians in the Western Hemisphere. Useful for demonstrating effects of climate change on linkages by birds among biomes.

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  • Lyons, S. Kathleen, Peter J. Wagner, and Katherine Dzikiewicz. 2010. Ecological correlates of range shifts of Late Pleistocene mammals. Philosophical Transactions Royal Society B 365:3681–3693.

    DOI: 10.1098/rstb.2010.0263Save Citation »Export Citation »

    The study examines whether Pleistocene mammal species’ range shifts differed because of taxonomic affinity, life-history traits, body size or topographic heterogeneity, and whether the species survived the megafaunal extinction. Useful approach for examining drivers of future changes in distributions.

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  • Reisen, William K. 2013. Ecology of West Nile virus in North America. Viruses 4.5: 2079–2105.

    DOI: 10.3390/v5092079Save Citation »Export Citation »

    Excellent review of the introduction, dispersal and establishment of West Nile virus in North America with a focus on factors that may have enhanced receptivity and enabled the invasion process.

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  • Robbins, Chandler S., John R. Sauer, Russell S. Greenberg, and Sam Droege. 1989. Population declines in North American birds that migrate to the Neotropics. Proceedings of the National Academy of Sciences 86:7658–7662.

    DOI: 10.1073/pnas.86.19.7658Save Citation »Export Citation »

    Illustrates how data from the North American Breeding Bird Survey can be used to examine linkages among biomes. Most neotropical migrant bird species that breed in forests of the eastern United States and Canada (1978–1987) declined in abundance after a period of stable or increasing populations.

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  • Urquhart, F. A., and N. R. Urquhart. 1978. Autumnal migration routes of the eastern population of the monarch butterfly (Danaus p. plexippus L.; Danaidae; Lepidoptera) in North America to the overwintering site in the Neovlcanic Plateau of Mexico. Canadian Journal of Zoology 56:1759–1764.

    DOI: 10.1139/z78-240Save Citation »Export Citation »

    One of the first studies to document the large-scale migration routes of Monarch butterflies and to show the connection between North and Central America.

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Connectivity with Other Biome Types Globally

North American biomes are connected with biomes on other continents and with oceans by the same transport vectors that connect biomes described above but across much longer distances. Dust storms in Asia and Africa can lead to dust deposition in North America with effects on local precipitation patterns and amounts, described by Creamean, et al. 2013. Hurricanes connect northern Africa with the Atlantic Coast and Gulf Coast of North America, described by Gray and Landsea 1992, Elsner and Birol Kara 1999, and Pielke and Landsea 1999. Warming of polar regions is increasing glacial melt with increases in sea level at lower latitudes in North America, described by Hopkinson, et al. 2008. Human activities, including transportation and land-use change, and climate change are leading to the spread of exotic species, pests, and pathogens across biomes globally, including North America, described by Crowl, et al. 2008 and Perrings, et al. 2010, with feedbacks to disease in humans described by Runstadler, et al. 2013.

  • Creamean, Jessie M., Kaitlyn J. Suski, Daniel Rosenfeld, et al. 2013. Dust and biological aerosols from the Sahara and Asia influence precipitation in the western US. Science 339:1572–1578.

    DOI: 10.1126/science.1227279Save Citation »Export Citation »

    This study presents the first direct cloud and precipitation measurements showing that Saharan and Asian dust and biological aerosols probably play an important role in precipitation processes over the western United States.

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  • Crowl, Todd A., Thomas O. Crist, Robert R. Parmenter, Gary Belovsky, and Ariel E. Lugo. 2008. The spread of invasive species and infectious disease as drivers of ecosystem change in an increasingly connected world. Frontiers in Ecology and the Environment 6:238–246.

    DOI: 10.1890/070151Save Citation »Export Citation »

    Excellent synthesis of the importance of long-distance movement of invasive plants and animals that is often facilitated by human activities.

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  • Elsner, James B., and A. Birol Kara. 1999. Hurricanes of the North Atlantic: Climate and society. Oxford: Oxford Univ. Press.

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    Reference guide for users interested in information on hurricanes in terms of physical processes and economic impacts. Emphasis is on physical models to explain statistical relationships between hurricane activity and weather or climate events.

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  • Gray, William M., and Christopher W. Landsea. 1992. African rainfall as a precursor of hurricane-related destruction on the U.S. East Coast. Bulletin American Meteorological Society 73:1352–1364.

    DOI: 10.1175/1520-0477(1992)073%3C1352:ARAAPO%3E2.0.CO;2Save Citation »Export Citation »

    This paper describes a predictive relationship between West African rainfall and hurricane destruction along the US East Coast based on information for the forty-two year period, 1949–1990. Similar hurricane damage along the Gulf Coast shows only a negligible relationship with African rainfall. One of the first papers to document tele-connections between North American biomes and rainfall in Africa.

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  • Hopkinson, Charles S., Ariel E. Lugo, Meryl A. Alber, Alan P. Covich, and Skip J. Van Bloem. 2008. Forecasting effects of sea-level rise and windstorms on coastal and inland ecosystems. Frontiers in Ecology and the Environment 6:255–263.

    DOI: 10.1890/070153Save Citation »Export Citation »

    Coastal wetland sites in North America are experiencing increases in sea level as glaciers and the polar ice cap melt with global warming. This paper illustrates spatial variation in these drivers using data from sites located in many different biomes.

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  • Perrings, Charles, Harold Mooney, and Mark Williamson. 2010. Bioinvasions and globalization: Ecology, economics, management, and policy. Oxford: Oxford Univ. Press.

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    Forward-looking book that links bioinvasions as one manifestation of globalization through the movement of humans. Improved transportation is leading to spread of pests and pathogens with ecological consequences for agricultural crops.

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  • Pielke, Roger A., Jr., and Christopher N. Landsea. 1999. La Niña, El Niño, and Atlantic hurricane damage in the United States. Bulletin of the American Meteorological Society 80:2027–2033.

    DOI: 10.1175/1520-0477(1999)080%3C2027:LNAENO%3E2.0.CO;2Save Citation »Export Citation »

    This paper compares the historical record of La Niña and El Niño events defined by eastern Pacific sea surface temperature with a dataset of hurricane losses normalized to 1997 values. The significant relationship found between the ENSO cycle and US hurricane losses, with La Niña years exhibiting much more damage, can be used in disaster prevention.

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  • Runstadler, Jonathan, Nichola Hill, Islam T. M. Hussein, Wendy Puryear, and Mandy Keogh. 2013. Connecting the study of wild influenza with the potential for pandemic disease. Infection, Genetics and Evolution 17:162–187.

    DOI: 10.1016/j.meegid.2013.02.020Save Citation »Export Citation »

    Reviews the current gaps in wild animal and environmental surveillance of avian influenza virus and the current understanding of genetic signatures in potentially pandemic strains.

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Climate Change and the Dynamics of Biomes

Current geographic distribution of biomes based on long-term climate and soil properties may not be the same as future distributions under a changing climate. Understanding biome responses to multiple types of drivers, such as those described for forests by Dale, et al. 2001, for the North America continent by Marshall, et al. 2008, and globally by Peters 2011, can be useful in projecting future biome distributions. Historic shifts in the geographic distribution of tree species with changes in climate, described by Davis and Shaw 2001, suggest that species composition of biomes may also change with climate. Understanding the mechanisms responsible for these shifts, similar to Gray, et al. 2006 which inferred mechanisms using pollen from packrat middens combined with tree ring analyses, will be instrumental to accurate predictions. Approaches to predicting future shifts in geographic distributions, such as native and invasive species by Peterson, et al. 2003, include habitat or niche models described by Phillips and Dudki 2008 and process-based simulation models that require more species and environmental parameters, described by Lenihan, et al. 2008.

  • Dale, Virginia H., Linda A. Joyce, Steve McNulty, et al. 2001. Climate change and forest disturbances. BioScience 51.9: 723–734.

    DOI: 10.1641/0006-3568(2001)051[0723:CCAFD]2.0.CO;2Save Citation »Export Citation »

    Classic paper showing that climate change can affect forests by altering the frequency, intensity, duration, and timing of fire, drought, introduced species, insect and pathogen outbreaks, hurricanes, windstorms, ice storms, or landslides. Essential reading about the importance of multiple drivers to biome responses.

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  • Davis, Margaret B., and Ruth G. Shaw. 2001. Range shifts and adaptive responses to Quaternary climate change. Science 292:673–679.

    DOI: 10.1126/science.292.5517.673Save Citation »Export Citation »

    One of first papers to show that tree species in North America shifted latitude or elevation range in response to changes in Quaternary climate. A classic, must read for anyone interested in climate change impacts.

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  • Gray, Stephen T., Julio L. Betancourt, Stephen T. Jackson, and Robert G. Eddy. 2006. Role of multidecadal climate variability in a range extension of Pinyon Pine. Ecology 87:1124–1130.

    DOI: 10.1890/0012-9658(2006)87[1124:ROMCVI]2.0.CO;2Save Citation »Export Citation »

    Used evidence from woodrat middens and tree rings at in northeastern Utah to reveal spatiotemporal patterns of pinyon pine (Pinus edulis Engelm.) colonization and expansion in the past millennium. Novel methods provided interesting results.

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  • Lenihan, James M., Dominique Bachelet, Ronald P. Neilson, and Raymond Drapek. 2008. Simulated response of conterminous United States ecosystems to climate change at different levels of fire suppression, CO2 emission rate, and growth response to CO2. Global and Planetary Change 64:16–25.

    DOI: 10.1016/j.gloplacha.2008.01.006Save Citation »Export Citation »

    A modeling experiment was used to investigate the impact of fire management, CO2 emission rate, and the growth response to CO2 on the response of ecosystems to climate scenarios produced by three different General Circulation Models. Interesting example of biome model used to examine finer-scale dynamics.

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  • Marshall, John D., John M. Blair, Debra P. C. Peters, Greg S. Okin, Albert Rango, and Mark Williams. 2008. Predicting and understanding ecosystem responses to climate change at continental scales. Frontiers in Ecology and the Environment 6:273–280.

    DOI: 10.1890/070165Save Citation »Export Citation »

    Excellent summary of regional- to continental-scale patterns in four drivers (drought, warming, snowpack loss, and wildfire) and their ecological responses.

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  • Peters, Debra P. C. 2011. Globalization: Ecological consequences of global-scale connectivity in people, resources, and information. In The systemic dimension of globalization. Edited by P. Pachura Rijeka, Croatia: InTech.

    DOI: 10.5772/19793Save Citation »Export Citation »

    Provides an overview of how various drivers, including drought and nitrogen deposition, can influence and interact with heterogeneity in land surface properties of vegetation and soils to synchronize, attenuate or amplify impacts of the drivers on ecological systems.

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  • Peterson, A. Townsend, Monica Papes, and Daniel A. Kluza. 2003. Predicting the potential invasive distributions of four alien plant species in North America. Weed Science 51:863–868.

    DOI: 10.1614/P2002-081Save Citation »Export Citation »

    Ecological niche modeling, one approach to investigating species responses to climate, was tested based on four invasive plant species (garlic mustard, sericea lespedeza, Russian olive, and hydrilla) in North America.

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  • Phillips, Steven J., and Miroslav Dudki. 2008. Modeling of species distributions with Maxent: New extensions and a comprehensive evaluation. Ecography 31:161–175.

    DOI: 10.1111/j.0906-7590.2008.5203.xSave Citation »Export Citation »

    This paper evaluates the species habitat model approach (Maxent: Maximum Entropy Modeling) to modeling species geographic ranges for 226 species from six regions.

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New Research Avenues and Contemporary Views

Recent studies of North American biomes focus on comparing their properties and dynamics based on data collected at research sites organized as networks, such as those described in Foster 2012 for the US Long Term Ecological Research Network. Hargrove, et al. 2003 and Parton, et al. 2007 provide examples of a bottom–up approach where comparable data collected by individual researchers or sites are analyzed to address specific questions. Keller, et al. 2008 describes a new top–down approach in ecology where the same data are collected by a team from a set of sites located strategically to represent the biomes of the United States. The data collected from these ecological observatories, such as the National Ecological Observatory Network, are sufficiently general, yet with the same methods from all sites, to be used for many questions in the future. Combining the data from bottom–up and top–down approaches provides new challenges described in Peters, et al. 2014 that builds on a macroecology perspective from Kerr, et al 2007 and new understanding of the role of climate in biome-level dynamics from Jackson, et al. 2007.

  • Foster, David R. 2012. Expanding the integration and application of long-term ecological research. BioScience 62:323.

    DOI: 10.1525/bio.2012.62.4.1Save Citation »Export Citation »

    Good description of the twenty-six Long Term Ecological Research sites that encompass diverse ecosystems in the continental United States, Alaska, Antarctica and islands in the Caribbean and the Pacific—including deserts, estuaries, lakes, oceans, coral reefs, prairies, forests, alpine and Arctic tundra, urban areas, and production agriculture. For more than three decades, the Network has generated rigorous, site-based scientific research that has led to important findings on regional and continental scales. This paper is an introduction to more detailed papers in the same issue.

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  • Hargrove, William W., Forest M. Hoffman, and Beverly E. Law. 2003. New analysis reveals representativeness of the AmeriFlux network. Eos Transactions, American Geophysical Union 84:529–535.

    DOI: 10.1029/2003EO480001Save Citation »Export Citation »

    Describes the AmeriFlux network that contains the longest-running sites in the world for ecosystem CO2, water, and energy fluxes. It includes 168 sites in North and South America: Brazil, Canada, Costa Rica, Mexico, Panama, and the United States (148 sites).

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  • Jackson, Stephen T., Julio L. Betancourt, Robert K. Booth, and Stephen T. Gray. 2007. Ecology and the ratchet of events: Climate variability, niche dimensions, and species distributions. Proceedings National Academy of Sciences 106:19685–19692.

    DOI: 10.1073/pnas.0901644106Save Citation »Export Citation »

    Draws on North American examples to develop approach to understanding and predicting broad-scale ecological responses to climate change that integrates correlational niche modeling with mechanistic niche modeling, dynamic ecological modeling, targeted experiments, and systematic observations of past and present patterns and dynamics at broad scales.

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  • Keller, Michael, David S. Schimel, William W. Hargrove, and Forrest M. Hoffman. 2008. A continental strategy for the National Ecological Observatory Network. Frontiers in Ecology and the Environment 6:282–284.

    DOI: 10.1890/1540-9295(2008)6[282:ACSFTN]2.0.CO;2Save Citation »Export Citation »

    Describes the National Ecological Observatory Network, a continental-scale observatory that measures the causes and effects of climate change, land-use change and invasive species on US ecosystems within biomes of North America using standard methods and instrumentation.

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  • Kerr, Jeremy T., Heather M. Kharouba, and David J. Currie. 2007. The macroecological contribution to global change solutions. Science 316:1581–1584.

    DOI: 10.1126/science.1133267Save Citation »Export Citation »

    Describes macroecology as approach to describe broad-scale predictions of species’ distributions and abundances to complement fine-scale, local studies that dominate global change biology.

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  • Parton, William, Whendee L. Silver, Ingrid C. Burke, et al. 2007. Global-scale similarities in nitrogen release patterns during long-term decomposition. Science 315:361–364.

    DOI: 10.1126/science.1134853Save Citation »Export Citation »

    Described a ten-year, twenty-eight-site experiment designed to test the effect of substrate quality and macroclimate on long-term decomposition and nutrient dynamics across multiple biomes in the United States.

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  • Peters, Debra P. C., Henry W. Loescher, Michael D. SanClements, and Kris M. Havstad. 2014. Taking the pulse of a continent: Expanding site-based research infrastructure for regional—to continental-scale ecology. Ecosphere 5.3: art. 29.

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    Describes limitations of current site-based approaches, and provides new framework to link different types of networks as way to address continental-scale, biome-level questions.

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