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Principles of terrestrial ecosystem ecology.pdf

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284 13. Temporal Dynamics<br />

Small rodents (index)<br />

50<br />

25<br />

0<br />

Flowers<br />

1972 1975 1978 1981<br />

Year<br />

Figure 13.2. Interannual variation in flowering<br />

density <strong>of</strong> an understory shrub (Vaccinium myrtillus)<br />

and <strong>of</strong> small rodents in northern Finland (Laine and<br />

<strong>of</strong> years. During the Tertiary they occupied a<br />

warm moist environment throughout much <strong>of</strong><br />

western North America. Their range is now<br />

restricted to valleys where coastal fog minimizes<br />

summer drought stress. Aspen clones in<br />

the Rocky Mountains range in age from a few<br />

years to as much as 10,000 years (Kemperman<br />

and Barnes 1976, Tuskan et al. 1996). The<br />

current distribution <strong>of</strong> redwoods and aspens<br />

is therefore a product <strong>of</strong> past population and<br />

community processes and cannot be fully<br />

understood with reference only to the present<br />

environment. Many species are still migrating<br />

poleward in response to the disappearance <strong>of</strong><br />

continental ice sheets 10,000 years ago. The<br />

soils beneath these recent arrivals may still<br />

reflect the properties <strong>of</strong> earlier communities<br />

rather than being completely a function <strong>of</strong><br />

current vegetation.<br />

The current functioning <strong>of</strong> the East Siberian<br />

coastal plain also reflects processes that occurred<br />

thousands <strong>of</strong> years ago. During the<br />

Pleistocene, this region was a steppe grassland<br />

that accumulated highly organic loess soils and<br />

ice lenses that now occupy 50 to 70% <strong>of</strong> the soil<br />

volume (Zimov et al. 1995). As the ice melted<br />

in the warmer Holocene climate, the soils subsided,<br />

forming lakes with organic-rich sediments.<br />

Pleistocene-age organic matter is now<br />

the major carbon source for methane production<br />

by these lakes (Zimov et al. 1997). In other<br />

words, the current processing <strong>of</strong> carbon in these<br />

lakes is strongly influenced by processes that<br />

occurred 10,000 to 100,000 years ago.<br />

Current <strong>ecosystem</strong> processes also respond<br />

to changes that have occurred more recently.<br />

Small<br />

rodents<br />

160<br />

Flowers (number m -2 )<br />

Henttonen 1983). These herbivores and their food<br />

plants show approximately 4-year cycles <strong>of</strong> abundance.<br />

Large areas <strong>of</strong> Europe and northeastern North<br />

America were deforested for agriculture in<br />

recent centuries and have more recently<br />

reverted to forests. Even forests older than 200<br />

years still exhibit composition and dynamics<br />

that reflect their earlier history (Foster and<br />

Motzkin 1998) (Fig. 13.3). A plow layer is still<br />

evident in these forests, for example, resulting<br />

in a sharp vertical discontinuity in soil processes<br />

and nutrient supply. Net primary production<br />

(NPP) substantially exceeds the rates<br />

<strong>of</strong> heterotrophic respiration in these <strong>ecosystem</strong>s,<br />

so vegetation and soils are actively accumulating<br />

carbon (Goulden et al. 1996). Any<br />

study <strong>of</strong> carbon dynamics that ignored these<br />

historical legacies would seriously misinterpret<br />

the relationship between carbon balance <strong>of</strong><br />

Landscape composition (% <strong>of</strong> land)<br />

100<br />

80<br />

60<br />

40<br />

20<br />

Forest<br />

Pasture<br />

80<br />

0<br />

0<br />

1650 1750 1850 1950<br />

Year<br />

0<br />

Cultivated<br />

100<br />

Figure 13.3. Changes in land use and population<br />

density in central Massachusetts (northeastern<br />

United States) since European colonization. Most<br />

forests in this region were previously croplands<br />

or pastures. (Redrawn with permission from<br />

Ecosystems; Foster and Motzkin 1998.)<br />

80<br />

60<br />

40<br />

20<br />

Population density (people km -2 )

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