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Importance Vegetation serves several critical functions in the biosphere, at all possible spatial scales. First, vegetation regulates the flow of numerous biogeochemical cycles (see biogeochemistry), most critically those of water, carbon, and nitrogen; it is also of great importance in local and global energy balances. Such cycles are important not only for global patterns of vegetation but also for those of climate. Second, vegetation strongly affects soil characteristics, including soil volume, chemistry and texture, which feed back to affect various vegetational characteristics, including productivity and structure. Third, vegetation serves as wildlife habitat and the energy source for the vast array of animal species on the planet. Vegetation is also critically important to the world economy, particularly in the use of fossil fuels as an energy source, but also in the global production of food, wood, fuel and other materials. Perhaps most importantly, and often overlooked, global vegetation (including algal communities) has been the primary source of oxygen in the atmosphere, enabling the aerobic metabolism systems to evolve and persist. Lastly, vegetation is psychologically important to humans, who evolved in direct contact with, and dependence on, vegetation, for food, sheleter, and medicines. Classification
Vegetation Structure
Vegetation Processes Like all biological systems, plant communities are temporally and spatially dynamic; they change at all possible scales. Dynamism in vegetation is defined primarily as changes in either or both of species composition and vegetation structure. Clari te amo demasiado Temporal Dynamics
Spatial Dynamics As a general rule, the larger an area under consideration, the more likely the vegetation will be heterogeneous across it. Two main factors are at work. First, the temporal dynamics of disturbance and succession are increasingly unlikely to be in synchrony across any area as the size of that area increases. That is, different areas will be at different developmental stages due to different local histories, particularly their times since last major disturbance. This fact interacts with inherent environmental variability, which is also a function of area. Environmental variability constrains the suite of species that can occupy a given area, and the two factors together interact to create a mosaic of vegetation conditions across the landscape. Only in agricultural or horticultural systems does vegetation ever approach perfect uniformity. In natural systems, there is always heterogeneity, although its scale and intensity will vary widely. A natural grassland may seem relatively homogeneous when compared to the same area of partially burned forest, but highly diverse and heterogeneous when compared to the wheat field next to it. Global Vegetation Patterns and Determinants At regional and global scales there is predictability of certain vegetation characteristics, especially physiognomic ones, which are related to the predictability in certain environmental characteristics. Much of the variation in these global patterns is directly explainable by corresponding patterns of temperature and precipitation (sometimes referred to as the energy and moisture balances). These two factors are highly interactive in their effect on plant growth, and their relationship to each other throughout the year is critical. Such relationships are shown graphically in climate diagrams. By graphing the long term monthly averages of the two variables against each other, an idea is given as to whether or not precipitation occurs during the warm season, when it is most useful, and consequently the type of vegetation to be expected. For example, two locations may have the same average annual precipitation and temperature, but if the relative timing of the precipitation and seasonal warmth are very different, so will their vegetation structure and growth and development processes be. Scientific Study Vegetation scientists study the causes of the patterns and processes observed in vegetation at various scales of space and time. Of particular interest and importance are questions of the relative roles of climate, soil, topography, and history on vegetation characteristics, including both species composition and structure. Such questions are often large scale, and so cannot easily be addressed by experimentation in a meaningful way. Observational studies supplemented by knowledge of botany, paleobotany, ecology, soil science etc, are thus the rule in vegetation science. Pre-1900 Vegetation science has its origins in the work of botanists and/or naturalists of the 18th century, or earlier in some cases. Many of these were world travelers on exploratory voyages in the Age of Exploration, and their work was a synthetic combination of botany and geography that today we would call plant biogeography (or phytogeography). Little was known about worldwide floristic or vegetation patterns at the time, and almost nothing about what determined them, so much of the work involved collecting, categorizing, and naming plant specimens. Little or no theoretical work occurred until the 19th century. The most productive of the early naturalists was Alexander von Humboldt, who collected 60,000 plant specimens on a five year voyage to South and Cental America from 1799 to 1804. Humboldt was one of the first to document the correspondence between climate and vegetation patterns, in his massive, life-long work "Voyage to the Equinoctial Regions of the New Continent", which he wrote with Aimé Bonpland, the botanist who accompanied him. Humboldt also described vegetation in physiogonmic terms rather than just taxonomically. His work presaged intensive work on environment-vegetation relationships that continues to this day (Barbour et al, 1987) The beginnings of vegetation study as we know it today began in Europe and Russia in the late 19th century, particularly under Jozef Paczoski, a Pole, and Leonid Ramensky, a Russian. Together they were much ahead of their time, introducing or elaborating on almost all topics germane to the field today, well before they were so in the west. These topics included plant community analysis, or phytosociology, gradient analysis, succession, and topics in plant ecophysiology and functional ecology. Due to language and/or political reasons, much of their work was unknown to much of the world, especially the English-speaking world, until well into the 20th century. Post-1900 In the United States, Henry Cowles and Frederic Clements developed ideas of plant succession in the early 1900s. Clements is famous for his now discredited view of the plant community as a "superorganism". He argued that, just as all organ systems in an individual must work together for the body to function well, and which develop in concert with each other as the individual matures, so the individual species in a plant community also develop and cooperate in a very tightly coordinated and synergistic way, pushing the plant community towards a defined and predictable end state. Although Clements did a great deal of work on North American vegetation, his devotion to the superorganism theory has hurt his reputation, as much work since then by numerous researchers has shown the idea to lack empirical support. In contrast to Clements, several ecologists have since demonstrated the validity of the individualistic hypothesis, which asserts that plant communities are simply the sum of a suite of species reacting individually to the environment, and co-occurring in time and space. Ramensky initiated this idea in Russia, and in 1926, Henry Gleason (Gleason, 1926) developed it in a paper in the United States. Gleason's ideas were categorically rejected for many years, so powerful was the influence of Clementsian ideas. However, in the 1950s and 60s, a series of well-designed studies by Robert Whittaker provided strong evidence for Gleason's arguments, and against those of Clements. Whittaker, considered one of the brightest and most productive of American plant ecologists, was a developer and proponent of gradient analysis, in which the abundances of individual species are measured against quantifiable environmental variables or their well-correlated surrogates. In studies in three very different montane ecosystems, Whittaker demonstrated strongly that species respond primarily to the environment, and not necessarily in any coordination with other, co-occurring species. Other work, particularly in paleobotany, has lent support to this view at larger temporal and spatial scales. More Recent Concepts, Theories, and Approaches Since the 1960s, much research into vegetation has revolved around topics in functional ecology. In a functional framework, taxonomic botany is relatively less important; investigations center around morphological, anatomical and physiological classifications of species, with the aim of predicting how particular groups thereof will respond to various environmental variables. The underlying basis for this approach is the observation that, due to convergent evolution and (conversely) adaptive radiation, there is often not a strong relationship between phylogenetic relatedness and environmental adaptations, especially at higher levels of the phylogenetic taxonomy, and at large spatial scales. Functional classifications arguably began in the 1930s with Raunkiaer's division of plants into groups based on the location of their apical meristems (buds) relative to the ground surface. This presaged later classifications such as Macarthur and Wilson's (1967) r vs K selected species (applied to all organisms, not just plants), and the "C-S-R" strategy proposed by Grime (1987) in which species are placed into one of three groups according to their ability to tolerate stress and the predictability of their environmental conditions. Functional classifications are crucial in modeling vegetation-environment interactions, which has been a leading topic in vegeation ecology for the last 30 or more years. Currently, there is a strong drive to model local, regional and global vegetation changes in response to global climate change, particularly changes in temperature, precipitation and disturbance regimes. Functional classifications such as the examples above, which attempt to categorize all plant species into a very small number of groups, are unlikely to be effective for the wide variety of different modeling purposes that exist or will exist. It is generally recognized that simple, all-purpose classifications will likely have to be replaced with more detailed and function-specific classifications for the modeling purpose at hand. This will require much better understanding of the physiology, anatomy, and developmental biology than currently exists, for a great number of species, even if only the dominant species in most vegetation types are considered. See also Classification Mapping-related Climate Diagrams | |||||||||||||||
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