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Principles of terrestrial ecosystem ecology.pdf

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228 10. Aquatic Carbon and Nutrient Cycling<br />

shallow-water <strong>ecosystem</strong>s, including coral reefs,<br />

seagrass beds, and kelp forests. In <strong>ecosystem</strong>s<br />

where physical attachment is possible, phytoplankton,<br />

benthic algal mats, multicellular algae<br />

such as sea lettuce (e.g., Ulva) and kelp (e.g.,<br />

Laminaria), and vascular plants like eelgrass<br />

(Zostera) occur in various combinations. The<br />

relative abundance <strong>of</strong> different types <strong>of</strong><br />

primary producers depends on many factors,<br />

including nutrient availability, water depth, the<br />

stability <strong>of</strong> substrates, and disturbance regime.<br />

Nutrient addition, for example, favors rapidly<br />

growing phytoplankton, which reduce the light<br />

available to multicellular algae at depth. Intermediate<br />

levels <strong>of</strong> disturbance and presence <strong>of</strong><br />

stable substrates favor multicellular primary<br />

producers (Sousa 1985).<br />

Oceans<br />

The large area and low productivity per unit<br />

area <strong>of</strong> ocean cancel out, so the ocean contributes<br />

nearly half (about 40%) <strong>of</strong> global NPP.<br />

MARINE PRODUCERS<br />

Corals<br />

Kelp and rockweeds<br />

Salt marsh grasses<br />

Sea grasses<br />

Mangroves<br />

Benthic microalgae<br />

Coastal phytoplankton<br />

Open sea phytoplankton<br />

FRESH-WATER PRODUCERS<br />

Macrophytes<br />

Phytoplankton (nutrient rich)<br />

Phytoplankton (nutrient poor)<br />

TERRESTRIAL PRODUCERS<br />

Tropical wet forests<br />

Temperate forests<br />

Grasslands<br />

Deserts and tundra<br />

0 1 2 3 4<br />

Production (kg C m -2 yr -1 )<br />

Although oceans cover 70% <strong>of</strong> Earth’s surface,<br />

the average NPP per unit area is only 20% <strong>of</strong><br />

that on land (Table 10.2). Aquatic productivity,<br />

however, is highly variable, just as on land.<br />

The most productive aquatic <strong>ecosystem</strong>s, such<br />

as coral reefs, kelp forests, and eutrophic lakes,<br />

can be at least as productive as the most productive<br />

<strong>terrestrial</strong> <strong>ecosystem</strong>s (Fig. 10.3). NPP<br />

in the open ocean, which accounts for 90% <strong>of</strong><br />

the ocean area, however, is similar to that <strong>of</strong><br />

<strong>terrestrial</strong> deserts and tundra. Because <strong>of</strong> its<br />

large area, the open ocean accounts for 60% <strong>of</strong><br />

marine production, with picoplankton accounting<br />

for about 90% <strong>of</strong> this production (Valiela<br />

1995).<br />

Carbon and Light Availability<br />

Photosynthesis is seldom carbon limited in<br />

the ocean. In marine pelagic <strong>ecosystem</strong>s, for<br />

example, only 1% <strong>of</strong> the carbon in a given<br />

water volume is involved in primary production,<br />

whereas the nitrogen in this water may<br />

cycle through primary production 10 times a<br />

Figure 10.3. NPP <strong>of</strong> selected marine,<br />

fresh-water, and <strong>terrestrial</strong> <strong>ecosystem</strong>s.<br />

Marine and fresh-water <strong>ecosystem</strong>s exhibit<br />

the same range <strong>of</strong> NPP that occurs on land,<br />

but unproductive marine <strong>ecosystem</strong>s (the<br />

open ocean) are much more widespread.<br />

(Redrawn with permission from Springer-<br />

Verlag; Valiela 1995.)

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