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Environmental Factors and Ecological Processes in Boreal Forests

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Environmental Factors and Ecological Processes in Boreal Forests Annu. Rev. Ecol. Syst. 1989. 20:1-28 Copyright © 1989 by Annual Reviews Inc. All rights reserved ENVIRONMENTAL FACTORS AND ECOLOGICAL PROCESSES IN BOREAL FORESTS Gordon B. Bonan Earth Resources Branch/Code -623, Laboratory for Terrestrial Physics, NASAl Goddard Space Flight Center, Greenbelt, Maryland 20771 Herman H. Shugart Department of Environmental Sciences, University of Virginia, Charlottesville, Virgi­ nia 22903 INTRODUCTION The boreal forest is a broad, circumpolar mixture of cool coniferous and deciduous tree species which covers over 14.7 million km2, or 11%, of the earth's terrestrial surface (Figure 1). At these latitudes, a strong correlation exists between the seasonal dynamics of atmospheric carbon dioxide and the seasonal dynamics of the "greenness" (49) of the earth (145)- A possible causal relation, in which the dynamics of the forests at these latitudes reg­ ulates the atmospheric carbon concentrations, appears to be consistent with the present-day understanding of ecological processes in these ecosystems Annu. Rev. Ecol. Syst. 1989.20:1-28. Downloaded from www.annualreviews.org by Swedish University of Agricultural Sciences on 02/05/14. For personal use only. (31, 46). Along with its familiar role in plant photosynthesis, carbon dioxide is a "greenhouse" gas that markedly affects the heat budget of the earth (23). Thus, the possibility that boreal forests may actively participate in the dynam­ ics of atmospheric carbon dioxide is of considerable significance, especially since the climatic response to elevated atmospheric carbon dioxide con­ centrations seems to be strongly directed to the boreal forests of the world (32, 128). These large-scale forest/environment interactions provide motivation to understand better the environmental factors controlling the structure and 0066-4162/89/1120-0001$02.00 2 HONAN & SHUGART 1100 1000 900E � Continuous Permafrost ELZI Discontinuous Permafrost [ill] Boreal Forest 1300 1100 Annu. Rev. Ecol. Syst. 1989.20:1-28. Downloaded from www.annualreviews.org by Swedish University of Agricultural Sciences on 02/05/14. For personal use only. Figure 1 Circumpolar distribution of the boreal forest. From Van Cleve & Dymess (151)_ function of boreal forest ecosystems. Numerous researchers have examined specific aspects of boreal forests, but no one has formulated a unifying model of the boreal forest, a paradigm to link the pattern of forest vegetation with causal environmental factors. In this review, we develop a qualitative con­ ceptual model of environmental factors and vegetation patterns in the cir­ cumpolar boreal forest. Our model represents a complex interrelationship among climate, solar radiation, soil moisture, soil temperature and per­ mafrost, the forest floor organic layer, nutrient availability, forest fires, insect outbreaks, and forest structure (Figure 2). FOREST STRUCTURE -- - -------i CLOUDINESS ,I - ---) -- - - -- - NUTRIENT _____ �I IWAILABILITY r---------, r----SOLAR iSLOPEI, ASPECT L--:" RADIATION ,�' ===--4=':=-"':"::4��"':":::':':"::+t--.!:=======t========:;-, �-----------; :..------------�----, ---- ttl 1------ - , ,"========:j' SOIL PHYSICAL: r PROPERTIES , __________-, I 1 ____________ L.. • � , l ·l���:��R�����_�: :::::=t-.._-.__r-..J �==== ru-- l -- U; 25 ELEVATION : en§ -- J >-i L u ur u .f��E-C�P�T-A-T���':... -' � , , ________ I ____________ , L.. J 5 Figure 2 Hypothesized environmental processes controlling forest structure and vegetation patterns in boreal forests. Dashed boxes represent climatic and Annu. Rev. Ecol. Syst. 1989.20:1-28. Downloaded from www.annualreviews.org edaphic input parameters. Arrows indicate interacting processes. � by Swedish University of Agricultural Sciences on 02/05/14. For personal use only. w 4 BONAN & SHUGART CLIMATE Geographic Patterns The climate of the boreal forest is characterized by strong seasonal variation with short, moderately warm, and moist summers and long, extremely cold, and dry winters (88, 122). This seasonal variation is most pronounced in the continental climates of interior Alaska and eastern Siberia, where seasonal fluctuation of mean monthly temperature is on the order of 44°C and more than 56°C, respectively (122). These harsh, continental regions are also characterized by a range of up to a 100°C in seasonal temperature extremes (122). Pronounced interannual variation in air temperatures is coupled with this strong seasonal variation. At Fairbanks, Alaska, mean monthly January air temperature between 1971 and 1986 ranged from -35 AOC to -7. 7°C, and averaged -22.1°C with a standard deviation of 7.1 (14). Together, these large seasonal and annual temperature fluctuations form a distinct feature of the continental climate of the boreal forest (88, 122). Where a more oceanic climate prevails, such as in eastern Canada and Scandinavia, these tempera­ ture fluctuations are not as extreme. Throughout the boreal forest, annual rainfall is relatively light. In North America, winters are characteristically dry, and more than half of the annual precipitation falls in the summer (22). In the northwest, where high moun­ tain ranges restrict the inland penetration of moisture-laden air, annual rainfall is less than 38 cm and, in Fort Yukon, Alaska, is as low as 18 cm (122). East of the Rocky Mountains, annual rainfall increases to 38-51 cm in central Canada and to 51-89 em in eastern Canada (122). In contrast to northwestern North America, the northern European boreal forest region west of the Ural Mountains lacks high mountain barriers. Here, under the moderating influence of maritime air from the north Atlan­ tic, the mean annual temperature range is relatively low (122). Though sum­ mers are cooler than in areas east of the Ural Mountains, winters are less Annu. Rev. Ecol. Syst. 1989.20:1-28. Downloaded from www.annualreviews.org severe (122). East of the Ural Mountains, the continentality of the climate by Swedish University of Agricultural Sciences on 02/05/14. For personal use only. increases. Here, seasonal weather patterns resemble those of northern North America, but the large seasonal contrast between warm, moist summers and cold, dry winters is exaggerated by the larger land area, which insulates most of the interior of the Soviet Union from moderating oceanic influences (122). In Eurasia, as in North America, most of the annual rainfall occurs in the summer (122). Annual rainfall is greater than 51 cm west of the Ural mountains, but less than 51 cm east of the Urals (122). In northeastern Siberia, precipitation averages less than 25 em and is as low as 13 em in BOREAL FOREST ECOLOGY 5 Verkhoyansk, which is well removed from the Arctic Ocean and isolated from the Pacific Ocean by mountains to the south and east (122). Climate-Vegetation Interactions Long-term average values of many climatic parameters, especially measures of summer temperatures, are coincident with the northern and southern boundaries of the boreal forest (56, 88, 107, 146, 170). Certain air mass characteristics also coincide with these boundaries. Throughout Canada, the taiga-tundra boundary is marked by the modal July position of the front that separates continental Arctic and maritime Pacific air masses (21), though this relationship is not as clear in the Labrador-Quebec peninsula (82). The Arctic air mass is associated with cool summers, long, cold winters, and low annual precipitation. Pacific air is typically warmer, moister, and more unstable than continental Arctic air. To the south, the boreal forest is bounded by the winter position of the Arctic front (21). Similar relationships have been found in the boreal forests of Eurasia (82). Climatic gradients usually are perpendicular to frontal positions, and if climate controls vegetation patterns, one would expect corresponding vegeta­ tion gradients normal to the mean Arctic frontal position. Indeed, radial growth of black spruce, Picea mariana, and white spruce, P. glauca, in central Canada declines sharply as the northern tree line (and hence Arctic front) is approached; isopleths of similar growth rates parallel the location of the Arctic front (101). In central Canada, the similarity of Picea mari­ ana communities also parallels the northern tree line (86). In addition, the occurrence of Picea mariana communities is positively correlated with the frequency of Arctic air mass and negatively correlated with the frequency of Pacific air mass; Picea glauca communities show the converse relation­ ship (87). However, the net energy received at the earth's surface, rather than air temperature or air mass characteristics, may be the critical factor for vegeta­ tion (50, 88). Both total stand biomass and net annual production increase Annu. Rev. Ecol. Syst. 1989.20:1-28. Downloaded from www.annualreviews.org with increases in annual net radiation (50). Hare & Ritchie (50) used this by Swedish University of Agricultural Sciences on 02/05/14. For personal use only. criterion to propose a zonal division of the boreal forest, and they speculated that the relationship Bryson (21) found between vegetation and air mass characteristics is manifested through the effect of air masses on the radiative balance. Larsen (88) also downplayed the importance of frontal activity, suggesting that it is significant only to the degree that air mass characteristics determine the characteristics of the local energy budget during the growing season. In high-latitude forests, where the growing season is restricted by cold temperatures, plant processes may be more limited by extreme, or anomalous,
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