Preface to the Princeton Landmarks in Biology Edition vii Preface xi Symbols Used xiii 1. The Importance of Islands 3 2. Area and Number of Speicies 8 3. Further Explanations of the Area-Diversity Pattern 19 4. The Strategy of Colonization 68 5. Invasibility and the Variable Niche 94 6. Stepping Stones and Biotic Exchange 123 7. Evolutionary Changes Following Colonization 145 8. Prospect 181 Glossary 185 References 193 Index 201

The Theory of Island Biogeography

Nitrogen is a key element controlling the species composition, diversity, dynamics, and functioning of many terrestrial, freshwater, and marine ecosystems. Many of the original plant species living in these ecosystems are adapted to, and function optimally in, soils and solutions with low levels of available nitrogen. The growth and dynamics of herbivore populations, and ultimately those of their predators, also are affected by N. Agriculture, combustion of fossil fuels, and other human activities have altered the global cycle of N substantially, generally increasing both the availability and the mobility of N over large regions of Earth. The mobility of N means that while most deliberate applications of N occur locally, their influence spreads regionally and even globally. Moreover, many of the mobile forms of N themselves have environmental consequences. Although most nitrogen inputs serve human needs such as agricultural production, their environmental conse- quences are serious and long term. Based on our review of available scientific evidence, we are certain that human alterations of the nitrogen cycle have: 1) approximately doubled the rate of nitrogen input into the terrestrial nitrogen cycle, with these rates still increasing; 2) increased concentrations of the potent greenhouse gas N 2O globally, and increased concentrations of other oxides of nitrogen that drive the formation of photochemical smog over large regions of Earth; 3) caused losses of soil nutrients, such as calcium and potassium, that are essential for the long-term maintenance of soil fertility; 4) contributed substantially to the acidification of soils, streams, and lakes in several regions; and 5) greatly increased the transfer of nitrogen through rivers to estuaries and coastal oceans. In addition, based on our review of available scientific evidence we are confident that human alterations of the nitrogen cycle have: 6) increased the quantity of organic carbon stored within terrestrial ecosystems; 7) accelerated losses of biological diversity, especially losses of plants adapted to efficient use of nitrogen, and losses of the animals and microorganisms that depend on them; and 8) caused changes in the composition and functioning of estuarine and nearshore ecosystems, and contributed to long-term declines in coastal marine fisheries.

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Human alteration of the global nitrogen cycle: sources and consequences

Ecological extinction caused by overfishing precedes all other pervasive human disturbance to coastal ecosystems, including pollution, degradation of water quality, and anthropogenic climate change. Historical abundances of large consumer species were fantastically large in comparison with recent observations. Paleoecological, archaeological, and historical data show that time lags of decades to centuries occurred between the onset of overfishing and consequent changes in ecological communities, because unfished species of similar trophic level assumed the ecological roles of overfished species until they too were overfished or died of epidemic diseases related to overcrowding. Retrospective data not only help to clarify underlying causes and rates of ecological change, but they also demonstrate achievable goals for restoration and management of coastal ecosystems that could not even be contemplated based on the limited perspective of recent observations alone.

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Historical overfishing and the recent collapse of coastal ecosystems.

Dead zones in the coastal oceans have spread exponentially since the 1960s and have serious consequences for ecosystem functioning. The formation of dead zones has been exacerbated by the increase in primary production and consequent worldwide coastal eutrophication fueled by riverine runoff of fertilizers and the burning of fossil fuels. Enhanced primary production results in an accumulation of particulate organic matter, which encourages microbial activity and the consumption of dissolved oxygen in bottom waters. Dead zones have now been reported from more than 400 systems, affecting a total area of more than 245,000 square kilometers, and are probably a key stressor on marine ecosystems.

Supporting Online Material for Spreading Dead Zones and Consequences for Marine Ecosystems

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Spreading Dead Zones and Consequences for Marine Ecosystems

The second largest zone of coastal hypoxia (oxygen-depleted waters) in the world is found on the northern Gulf of Mexico continental shelf adjacent to the outflows of the Mississippi and Atchafalaya Rivers. The combination of high freshwater discharge, wind mixing, regional circulation, and summer warming controls the strength of stratification that goes through a well-defined seasonal cycle. The physical structure of the water column and high nutrient loads that enhance primary production lead to an annual formation of the hypoxic water mass that is dominant from spring through late summer. Paleoindicators in dated sediment cores indicate that hypoxic conditions likely began to appear around the turn of the last century and became more severe since the 1950s as the nitrate flux from the Mississippi River to the Gulf of Mexico tripled. Whereas increased nutrients enhance the production of some organisms, others are eliminated from water masses (they either emigrate from the area or die) where the...

Gulf of Mexico Hypoxia, A.K.A. “The Dead Zone”

Rabalais, N. N., Turner, R. E., Diaz, R. J., and Justic, D. 2009. Global change and eutrophication of coastal waters. - ICES Journal of Marine Science, 66: 1528-1537.The cumulative effects of global change, including climate change, increased population, and more intense industrialization and agribusiness, will likely continue and intensify the course of eutrophication in estuarine and coastal waters. As a result, the symptoms of eutrophication, such as noxious and harmful algal blooms, reduced water quality, loss of habitat and natural resources, and severity of hypoxia (oxygen depletion) and its extent in estuaries and coastal waters will increase. Global climate changes will likely result in higher water temperatures, stronger stratification, and increased inflows of freshwater and nutrients to coastal waters in many areas of the globe. Both past experience and model forecasts suggest that these changes will result in enhanced primary production, higher phytoplankton and macroalgal standing stocks, and more frequent or severe hypoxia. The negative consequences of increased nutrient loading and stratification may be partly, but only temporarily, compensated by stronger or more frequent tropical storm activity in low and mid-latitudes. In anticipation of the negative effects of global change, nutrient loadings to coastal waters need to be reduced now, so that further water quality degradation is prevented.

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Global change and eutrophication of coastal waters

The Mississippi River system ranks among the world's top 10 rivers in freshwater and sediment inputs to the coastal ocean. The river contributes 90% of the freshwater loading to the Gulf of Mexico, and terminates amidst one of the United States' most productive fisheries regions and the location of the largest zone of hypoxia, in the western Atlantic Ocean. Significant increases in riverine nutrient concentrations and loadings of nitrate and phosphorus and decreases in silicate have occurred this century, and have accelerated since 1950. Consequently, major alterations have occurred in the probable nutrient limitation and overall stoichiometric nutrient balance in the adjacent continental shelf system. Changes in the nutrient balances and reduction in riverine silica loading to, the continental shelf appear to have led to phytoplankton species shifts offshore and to an increase in primary production. The phytoplankton community response, as indicated by long-term changes in biological uptake of silicate and accumulation of biologically bound silica in sediments, has shown how the system has responded to changes in riverine nutrient loadings. Indeed, the accumulation of biologically bound silica in sediments beneath the Mississippi River plume increased during the past two decades, presumably in response to, increased nitrogen loading. The duration, size, and severity of hypoxia has probably increased as a consequence of the increased primary production. Management alternatives directed at water pollution issues within the Mississippi River watershed may have unintended and contrasting impacts on the coastal waters of the northern Gulf of Mexico.

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Nutrient changes in the Mississippi River and system responses on the adjacent continental shelf

CHANGES in delivery of river-borne nutrients such as dissolved phosphate, nitrate and silicate, owing to land-use changes and anthropogenic emissions, are known to result in eutrophication1— enhanced phytoplankton blooms—and more severe hypoxic events2–1 in many enclosed bays and seas. Although similar ecological effects might be expected on continental shelves, the occurrence of such eutrophication has remained unresolved5. Here we present evidence of eutrophication of the continental shelf near the outflow of the Mississippi river, obtained by quantifying biologically bound silica (BSi) in diatom remnants within dated sediment cores. BSi accumulation rates are greatest in water depths of 20 to 50 m within 100 km of the river mouth, and have increased by as much as 100% this century. The increases were substantial by 1980, by which time riverine nitrogen loading had doubled relative to the beginning of the century, even though the silica loading had declined by 50% over the same period. Thus changes in river-borne nutrient loadings can modify coastal food webs and affect the amount and distribution of oxygen in bottom waters on the scale of continental shelves.

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Coastal eutrophication near the Mississippi river delta

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R. Eugene Turner

Papers

Gulf of Mexico Hypoxia, A.K.A. “The Dead Zone”

Global change and eutrophication of coastal waters

Nutrient changes in the Mississippi River and system responses on the adjacent continental shelf

Coastal eutrophication near the Mississippi river delta

Beyond Science into Policy: Gulf of Mexico Hypoxia and the Mississippi River