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

Statistical method for testing the neutral mutation hypothesis by DNA polymorphism.

The influence of diet on the distribution of nitrogen isotopes in animals was investigated by analyzing animals grown in the laboratory on diets of constant nitrogen isotopic composition. 
The isotopic composition of the nitrogen in an animal reflects the nitrogen isotopic composition of its diet. The δ^(15)N values of the whole bodies of animals are usually more positive than those of their diets. Different individuals of a species raised on the same diet can have significantly different δ^(15)N values. The variability of the relationship between the δ^(15)N values of animals and their diets is greater for different species raised on the same diet than for the same species raised on different diets. Different tissues of mice are also enriched in ^(15)N relative to the diet, with the difference between the δ^(15)N values of a tissue and the diet depending on both the kind of tissue and the diet involved. The δ^(15)N values of collagen and chitin, biochemical components that are often preserved in fossil animal remains, are also related to the δ^(15)N value of the diet. 
The dependence of the δ^(15)N values of whole animals and their tissues and biochemical components on the δ^(15)N value of diet indicates that the isotopic composition of animal nitrogen can be used to obtain information about an animal's diet if its potential food sources had different δ^(15)N values. The nitrogen isotopic method of dietary analysis probably can be used to estimate the relative use of legumes vs non-legumes or of aquatic vs terrestrial organisms as food sources for extant and fossil animals. However, the method probably will not be applicable in those modern ecosystems in which the use of chemical fertilizers has influenced the distribution of nitrogen isotopes in food sources. 
The isotopic method of dietary analysis was used to reconstruct changes in the diet of the human population that occupied the Tehuacan Valley of Mexico over a 7000 yr span. Variations in the δ^(15)C and δ^(15)N values of bone collagen suggest that C_4 and/or CAM plants (presumably mostly corn) and legumes (presumably mostly beans) were introduced into the diet much earlier than suggested by conventional archaeological analysis.

Influence of Diet On the Distribtion of Nitrogen Isotopes in Animals

Evolution and ecology of influenza A viruses.

A DNA test to sex most birds.

Unlike exercising mammals, migratory birds fuel very high intensity exercise (e.g., flight) with fatty acids delivered from the adipose tissue to the working muscles by the circulatory system. Given the primary importance of fatty acids for fueling intense exercise, we discuss the likely limiting steps in lipid transport and oxidation for exercising birds and the ecological factors that affect the quality and quantity of fat stored in wild birds. Most stored lipids in migratory birds are comprised of three fatty acids (16:0, 18:1 and 18:2) even though migratory birds have diverse food habits. Diet selection and selective metabolism of lipids play important roles in determining the fatty acid composition of birds which, in turn, affects energetic performance during intense exercise. As such, migratory birds offer an intriguing model for studying the implications of lipid metabolism and obesity on exercise performance. We conclude with a discussion of the energetic costs of migratory flight and stopover in birds, and its implications for bird migration strategies.

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Flying, fasting, and feeding in birds during migration: a nutritional and physiological ecology perspective

Birds prepare their eggs from recently ingested nutrients ('income' breeders) or from body stores ('capital' breeders)1. As summers are short at Arctic latitudes, Arctic migrants have been presumed to bring nutrients for egg production from their previous habitats, so that they can start breeding immediately upon arrival1,2,3. But we show here that eggs laid by 10 different wader species from 12 localities in northeast Greenland and Arctic Canada are produced from nutrients originating from tundra habitats, as inferred from carbon stable-isotope ratios in eggs, natal down, and juvenile and adult feathers.

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Ornithology. Arctic waders are not capital breeders.

It is increasingly acknowledged that migratory birds, notably waterfowl, play a critical role in the maintenance and spread of influenza A viruses. In order to elucidate the epidemiology of influenza A viruses in their natural hosts, a better understanding of the pathological effects in these hosts is required. Here we report on the feeding and migratory performance of wild migratory Bewick's swans (Cygnus columbianus bewickii Yarrell) naturally infected with low-pathogenic avian influenza (LPAI) A viruses of subtypes H6N2 and H6N8. Using information on geolocation data collected from Global Positioning Systems fitted to neck-collars, we show that infected swans experienced delayed migration, leaving their wintering site more than a month after uninfected animals. This was correlated with infected birds travelling shorter distances and fuelling and feeding at reduced rates. The data suggest that LPAI virus infections in wild migratory birds may have higher clinical and ecological impacts than previously recognised.

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Hampered Foraging and Migratory Performance in Swans Infected with Low-Pathogenic Avian Influenza A Virus

The flight range of migrating birds depends crucially on the amount of fuel stored by the bird prior to migration or taken up en route at stop-over sites. However, an increase in body mass is associated with an increase in energetic costs, counteracting the benefit of fuel stores. Water imbalance, occurring when water loss exceeds metabolic water production, may constitute another less well recognised problem limiting flight range. The main route of water loss during flight is via the lungs; the rate of loss depends on ambient temperature, relative humidity and ventilatory flow and increases with altitude. Metabolite production results in an increased plasma osmolality, also endangering the proper functioning of the organism during flight. Energetic constraints and water-balance problems may interact in determining several aspects of flight behaviour, such as altitude of flight, mode of flight, lap distance and stop-over duration. To circumvent energetic and water-balance problems, a bird could migrate in short hops instead of long leaps if crossing of large ecological barriers can be avoided. However, although necessitating larger fuel stores and being more expensive, migration by long leaps may sometimes be faster than by short hops. Time constraints are also an important factor in explaining why soaring, which conserves energy and water, occurs exclusively in very large species: small birds can soar at low speeds only. Good navigational skills involving accurate orientation and assessment of altitude and air and ground speed assist in avoiding physiological stress during migration.

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Metabolic constraints on long-distance migration in birds

-During directional flight trained Eurasian Kestrels (Falco tinnunculus) in the laboratory expended 13.8 W. Free-living birds expended 14.6 W during nonsoaring flight that included both directional flight and wind hovering. The former value was obtained by combining food balance and indirect calorimetry techniques, the latter by doubly labeled water (D2180). Because the energy-expenditure rates are so similar for directional flight alone and for wind hovering and directional flight combined, we argue that the value for either mode of kestrel flight may be used to analyze time-energy budgets. We predicted avian flight costs (ef) from an equation based on published data on flight costs in 14 species (body mass 3.8-1,000 g): ef = 17.360M1 013b0 -4236S, 1.926 W where M is body mass (g), b, is wing span (cm), and sw is wing area (cm2). Inclusion of the morphological data with body mass significantly improved the prediction of flight cost [r2 = 0.84 vs. r2 = 0.75 without b, and sw, F, (2,18) = 5.34, P < 0.05]. Received 8 May 1986, accepted 29 November 1986. QUANTIFICATION of the components of daily energy expenditure will improve the understanding of the behavior of free-living birds. This can be achieved by combining time-activity budgets with estimates of the various cost factors (e.g. basal metabolism, thermoregulation, flight; Koplin et al. 1980, Mugaas and King 1981). Energy expenditure during flight is increased greatly relative to during other behaviors (e.g. Berger and Hart 1974). Therefore, even though little time may be devoted to flight, an error in the estimate of the flight cost will produce a considerable error in the total daily energy expenditure estimated from time-activity models. Available allometric equations that predict flight cost in birds are based solely on body mass (e.g. Berger and Hart 1974, Kendeigh et al. 1977, Butler 1980) and provide only a rough estimate of power consumption. Energy expenditure during flight varies among birds of the same mass, according to flight pattern and the aerodynamic and behavioral properties of the species (Nisbet 1967, Utter and LeFebvre 1970, Hails 1979, Dolnik 1982, Flint and Nagy 1984). The Eurasian Kestrel (Falco tinnunculus) has two distinct modes of powered flight: (1) directional flapping flight and (2) wind hovering, which is flapping flight against the wind with zero ground speed (Videler et al. 1983). We determined energy expenditure in kestrels during powered flight. We used this information to quantify total daily energy expenditure of the Eurasian Kestrel throughout its annual cycle (Masman 1986). We used two techniques to determine flight cost. The energy budgets of trained, free-flying kestrels in the laboratory were reconstructed by monitoring daily metabolizable energy intake, oxygen consumption during rest, and time spent flying per day, from which the energy expenditure during directional flight was estimated. In the field we measured daily energy expenditure of free-living kestrels by a doubly labeled water (D2180) technique (Lifson et al. 1955). Time spent flying was recorded simultaneously. Energy expenditure during combined directional flight and wind hovering was estimated from the correlation between daily CO2 production and time spent in flight.

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Marcel Klaassen

Papers

Flying, fasting, and feeding in birds during migration: a nutritional and physiological ecology perspective

Ornithology. Arctic waders are not capital breeders.

Hampered Foraging and Migratory Performance in Swans Infected with Low-Pathogenic Avian Influenza A Virus

Metabolic constraints on long-distance migration in birds

Energy-expenditure during free flight in trained and free-living eurasian kestrels (falco-tinnunculus)