Beyond the forest edge: Ecology, diversity and conservation of the grassy biomes

The objectives of this synthesis are (1) to review the factors that influence the ecological, geographical, and palaeoecological distributions of plants possessing C4 photosynthesis and (2) to propose a hypothesis/model to explain both the distribution of C4 plants with respect to temperature and CO2 and why C4 photosynthesis is relatively uncommon in dicotyledonous plants (hereafter dicots), especially in comparison with its widespread distribution in monocotyledonous species (hereafter monocots). Our goal is to stimulate discussion of the factors controlling distributions of C4 plants today, historically, and under future elevated CO2 environments. Understanding the distributions of C3/C4 plants impacts not only primary productivity, but also the distribution, evolution, and migration of both invertebrates and vertebrates that graze on these plants. Sixteen separate studies all indicate that the current distributions of C4 monocots are tightly correlated with temperature: elevated temperatures during the growing season favor C4 monocots. In contrast, the seven studies on C4 dicot distributions suggest that a different environmental parameter, such as aridity (combination of temperature and evaporative potential), more closely describes their distributions. Differences in the temperature dependence of the quantum yield for CO2 uptake (light-use efficiency) of C3 and C4 species relate well to observed plant distributions and light-use efficiency is the only mechanism that has been proposed to explain distributional differences in C3/C4 monocots. Modeling of C3 and C4 light-use efficiencies under different combinations of atmospheric CO2 and temperature predicts that C4-dominated ecosystems should not have expanded until atmospheric CO2 concentrations reached the lower levels that are thought to have existed beginning near the end of the Miocene. At that time, palaeocarbonate and fossil data indicate a simultaneous, global expansion of C4-dominated grasslands. The C4 monocots generally have a higher quantum yield than C4 dicots and it is proposed that leaf venation patterns play a role in increasing the light-use efficiency of most C4 monocots. The reduced quantum yield of most C4 dicots is consistent with their rarity, and it is suggested that C4 dicots may not have been selected until CO2 concentrations reached their lowest levels during glacial maxima in the Quaternary. Given the intrinsic light-use efficiency advantage of C4 monocots, C4 dicots may have been limited in their distributions to the warmest ecosystems, saline ecosystems, and/or to highly disturbed ecosystems. All C4 plants have a significant advantage over C3 plants under low atmospheric CO2 conditions and are predicted to have expanded significantly on a global scale during full-glacial periods, especially in tropical regions. Bog and lake sediment cores as well as pedogenic carbonates support the hypothesis that C4 ecosystems were more extensive during the last glacial maximum and then decreased in abundance following deglaciation as atmospheric CO2 levels increased.

C4 photosynthesis is a series of anatomical and biochemical modifications that concentrate CO2 around the carboxylating enzyme Rubisco, thereby increasing photosynthetic efficiency in conditions promoting high rates of photorespiration. The C4 pathway independently evolved over 45 times in 19 families of angiosperms, and thus represents one of the most convergent of evolutionary phenomena. Most origins of C4 photosynthesis occurred in the dicots, with at least 30 lineages. C4 photosynthesis first arose in grasses, probably during the Oligocene epoch (24-35 million yr ago). The earliest C4 dicots are likely members of the Chenopodiaceae dating back 15-21 million yr; however, most C4 dicot lineages are estimated to have appeared relatively recently, perhaps less than 5 million yr ago. C4 photosynthesis in the dicots originated in arid regions of low latitude, implicating combined effects of heat, drought and/or salinity as important conditions promoting C4 evolution. Low atmospheric CO2 is a significant contributing factor, because it is required for high rates of photorespiration. Consistently, the appearance of C4 plants in the evolutionary record coincides with periods of increasing global aridification and declining atmospheric CO2 . Gene duplication followed by neo- and nonfunctionalization are the leading mechanisms for creating C4 genomes, with selection for carbon conservation traits under conditions promoting high photorespiration being the ultimate factor behind the origin of C4 photosynthesis. Contents Summary 341 I. Introduction 342 II. What is C4 photosynthesis? 343 III. Why did C4 photosynthesis evolve? 347 IV. Evolutionary lineages of C4 photosynthesis 348 V. Where did C4 photosynthesis evolve? 350 VI. How did C4 photosynthesis evolve? 352 VII. Molecular evolution of C4 photosynthesis 361 VIII. When did C4 photosynthesis evolve 362 IX. The rise of C4 photosynthesis in relation to climate and CO2 363 X. Final thoughts: the future evolution of C4 photosynthesis 365 Acknowledgements 365 References 365.

The incidence and importance of fire in the Amazon have increased substantially during the past decade, but the effects of this disturbance force are still poorly understood. The forest fire dynamics in two regions of the eastern Amazon were studied. Accidental fires have affected nearly 50 percent of the remaining forests and have caused more deforestation than has intentional clearing in recent years. Forest fires create positive feedbacks in future fire susceptibility, fuel loading, and fire intensity. Unless current land use and fire use practices are changed, fire has the potential to transform large areas of tropical forest into scrub or savanna.