Atmosphere and Climate Regulation Life on earth plays a critical role in regulating the earth's physical, chemical, and geological properties, from influencing the chemical composition of the atmosphere to modifying climate. About 3.5 billion years ago, early life forms (principally cyanobacteria) helped create an oxygenated atmosphere through photosynthesis, taking up carbon dioxide from the atmosphere and releasing oxygen (Schopf 1983; Van Valen 1971). Over time, these organisms altered the composition of the atmosphere, increasing oxygen levels, and paved the way for organisms that use oxygen as an energy source (aerobic respiration), forming an atmosphere similar to that existing today. Carbon cycles on the planet between the land, atmosphere, and oceans through a combination of physical, chemical, geological, and biological processes (IPCC 2001). One key way biodiversity influences the composition of the earth's atmosphere is through its role in carbon cycling in the oceans, the largest reservoir for carbon on the planet (Gruber and Sarmiento, in press). In turn, the atmospheric composition of carbon influences climate. Phytoplankton (or microscopic marine plants) play a central role in regulating atmospheric chemistry by transforming carbon dioxide into organic matter during photosynthesis. This carbon-laden organic matter settles either directly or indirectly (after it has been consumed) in the deep ocean, where it stays for centuries, or even thousands of years, acting as the major reservoir for carbon on the planet. In addition, carbon also reaches the deep ocean through another biological process -- the formation of calcium carbonate, the primary component of the shells in two groups of marine organisms coccolithophorids (a phytoplankton) and foraminifera (a single celled, shelled organism that is abundant in many marine environments). When these organisms die, their shells sink to the bottom or dissolve in the water column. This movement of carbon through the oceans removes excess carbon from the atmosphere and regulates the earth's climate. Over the last century, humans have changed the atmosphere's composition by releasing large amounts of carbon dioxide. This excess carbon dioxide, along with other 'greenhouse' gases, is believed to be heating up our atmosphere and changing the world's climate, leading to 'global warming'. There has been much debate about how natural processes, such as the cycling of carbon through phytoplankton in the oceans, will respond to these changes. Will phytoplankton productivity increase and thereby absorb the extra carbon from the atmosphere? Recent studies suggest that natural processes may slow the rate of increase of carbon dioxide in the atmosphere, but it is doubtful that either the earth's oceans or its forests can absorb the entirety of the extra carbon released by human activity (Falkowski et al. 2000).

Land Use Change and Climate Regulation The energy source that ultimately drives the earth's climate is the sun. The amount of solar radiation absorbed by the earth depends primarily on the characteristics of the surface. Although the link between solar absorption, thermodynamics, and ultimately climate is very complex, newer studies indicate that vegetation cover and seasonal variation in vegetation cover affects climate on both global and local scales. New generations of atmospheric circulation models are increasingly able to incorporate more complex data related to these parameters (Sellers et al. 1997). Besides regulating the atmosphere's composition, the extent and distribution of different types of vegetation over the globe modifies climate in three main ways: affecting the reflectance of sunlight (radiation balance); regulating the release of water vapor (evapotranspiration); and changing wind patterns and moisture loss (surface roughness). The amount of solar radiation reflected by a surface is known as its albedo; surfaces with low albedo reflect a small amount of sunlight, those with high albedo reflect a large amount. Different types of vegetation have different albedos; forests typically have low albedo, whereas deserts have high albedo. Deciduous forests are a good example of the seasonal relationship between vegetation and radiation balance. In the summer, the leaves in deciduous forests absorb solar radiation through photosynthesis; in winter, after their leaves have fallen, deciduous forests tend to reflect more radiation. These seasonal changes in vegetation modify climate in complex ways, by changing evapotranspiration rates and albedo (IPCC 2001). Vegetation absorbs water from the soil and releases it back into the atmosphere through evapotranspiration, which is the major pathway by which water moves from the soil to the atmosphere. This release of water from vegetation cools the air temperature. In the Amazon region, vegetation and climate is tightly coupled; evapotranspiration of plants is believed to contribute an estimated fifty percent of the annual rainfall (Salati 1987). Deforestation in this region leads to a complex feedback mechanism, reducing evapotranspiration rates, which leads to decreased rainfall and increased vulnerability to fire (Laurance and Williamson 2001). Deforestation also influences the climate of cloud forests in the mountains of Costa Rica. The Monteverde Cloud Forest harbors a rich diversity of organisms, many of which are found nowhere else in the world. However, deforestation in lower-lying lands, even regions over 50 kilometers way, is changing the local climate, leaving the "cloud" forest cloudless (Lawton et al. 2001). As winds pass over deforested lowlands, clouds are lifted higher, often above the mountaintops, reducing the ability for cloud forests to form. Removing the clouds from a cloud forest dries the forest, so it can no longer support the same vegetation or provide appropriate habitat for many of the species originally found there. Similar patterns may be occurring in other, less studied montane cloud forests around the world. Different vegetation types and topographies have varying surface roughness, which change the flow of winds in the lower atmosphere and in turn influences climate. Lower surface roughness also tends to reduce surface moisture and increase evaporation. Farmers apply this knowledge when they plant trees to create windbreaks (Johnson et al. 2003). Windbreaks reduce wind speed and change the microclimate, increase surface roughness, reduce soil erosion, and modify temperature and humidity. For many field crops, windbreaks increase yields and production efficiency. They also minimize stress on livestock from cold winds.

Soil and Water Conservation Biodiversity is also important for global soil and water protection. Terrestrial vegetation in forests and other upland habitats maintain water quality and quantity, and controls soil erosion. In watersheds where vegetation has been removed, flooding prevails in the wet season and drought in the dry season. Soil erosion is also more intense and rapid, causing a double effect: removing nutrient-rich topsoil and leading to siltation in downstream riverine and ultimately oceanic environments. This siltation harms riverine and coastal fisheries as well as damaging coral reefs (Turner and Rabalais 1994; van Katwijk et al. 1993). One of the most productive ecosystems on earth, wetlands have water present at or near the surface of the soil or within the root zone, all year or for a period of time during the year, and the vegetation there is adapted to these conditions. Wetlands are instrumental for the maintenance of clean water and erosion control. Microbes and plants in wetlands absorb nutrients and in the process filter and purify water of pollutants before they can enter coastal or other aquatic ecosystems. Wetlands also reduce flood, wave, and wind damage. They retard the flow of floodwaters and accumulate sediments that would otherwise be carried downstream or into coastal areas. Wetlands also serve as breeding grounds and nurseries for fish and support thousands of bird and other animal species.

Nutrient Cycling Nutrient cycling is yet another critical service provided by biodiversity -- particularly by microorganisms. Fungi and other microorganisms in soil help break down dead plants and animals, eventually converting this organic matter into nutrients that enrich the soil (Pimentel et al. 1995). Nitrogen is essential for plant growth, and an insufficient quantity of it limits plant production in both natural and agricultural ecosystems. While nitrogen is abundant in the atmosphere, only a few organisms (commonly known as nitrogen-fixing bacteria) can use it in this form. Nitrogen-fixing bacteria extract nitrogen from the air, and transform it into ammonia, then other bacteria further break down this ammonia into nitrogenous compounds that can be absorbed and used by most plants. In addition to their role in decomposition and hence nutrient cycling, microorganisms also help detoxify waste, changing waste products into forms less harmful to humans.

Pollination and Seed Dispersal An estimated 90 percent of flowering plants depend on pollinators such as wasps, birds, bats, and bees, to reproduce. Plants and their pollinators are increasingly threatened around the world (Buchmann and Nabhan 1995; Kremen and Ricketts 2000). Pollination is critical to most major crops and virtually impossible to replace. For instance, imagine how costly fruit would be (and how little would be available) if its natural pollinators no longer existed and each developing flower had to be fertilized by hand. Many animal species are important dispersers of plant seeds. It has been hypothesized that the loss of a seed disperser could cause a plant to become extinct. At present, there is no example where this has occurred. A famous example that has often been cited previously is the case of the dodo (Raphus cucullatus) and the tambalacoque (Sideroxylon grandiflorum). The dodo, a large flightless bird that inhabited the island of Mauritius in the Indian Ocean, became extinct due to overhunting in the late seventeenth century. It was once thought that the tambalacoque, a now endangered tree, depended upon the dodo to germinate its hard-cased seeds (Temple 1977). In the 1970s, only 13 trees remained and it was thought the tree had not reproduced for 300 years. The seeds of the tree have a very hard coat, as an experiment they were fed to a turkey; after passing through its gizzard the seeds were viable and germinated. This experiment led scientists to believe that the extinction of the dodo was coupled to the tambalacoque's inability to reproduce. However, this hypothesis has not stood up to further scrutiny, as there were several other species (including three now extinct species, a large-billed parrot, a giant tortoise, and a giant lizard) that were also capable of cracking the seed (Witmar and Cheke 1991; Catling 2001). Thus many factors, including the loss of the dodo, could have contributed to the decline of the tambalacoque. (For further details of causes of extinction see Historical Perspectives on Extinction and the Current Biodiversity Crisis). Unfortunately, declines and/or extinctions of species are often unobserved and thus it is difficult to tease out the cause of the end result, as multiple factors are often operating simultaneously. Similar problems exist today in understanding current population declines. For example, in a given species, population declines may be caused by loss of habitat, loss in prey species or loss of predators, a combination of these factors, or possibly some other yet unidentified cause, such as disease. In the pine forests of western North America, corvids (including jays, magpies, and crows), squirrels, and bears play a role in seed dispersal. The Clark's nutcracker (Nucifraga columbiana) is particularly well adapted to dispersal of whitebark pine (Pinus albicaulis) seeds (Lanner 1996). The nutcracker removes the wingless seeds from the cones, which otherwise would not open on their own. Nutcrackers hide the seeds in clumps. When the uneaten seeds eventually grow, they are clustered, accounting for the typical distribution pattern of whitebark pine in the forest. In tropical areas, large mammals and frugivorous birds play a key role in dispersing the seeds of trees and maintaining tree diversity over large areas. For example, three-wattled bellbirds (Procnias tricarunculata) are important dispersers of tree seeds of members of the Lauraceae family in Costa Rica. Because bellbirds return again and again to one or more favorite perches, they take the fruit and its seeds away from the parent tree, spreading Lauraceae trees throughout the forest (Wenny and Levy 1998).