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

▪ Abstract Lentic freshwater habitats in temperate regions exist along a gradient from small ephemeral ponds to large permanent lakes. This environmental continuum is a useful axis for understanding how attributes of individuals ultimately generate structure at the level of the community. Community structure across the gradient is determined by both (a) physical factors, such as pond drying and winter anoxia, that limit the potential breadth of species distributions, and (b) biotic effects mediated by ecological interactions, principally predation, that determine the realized success of species. Fitness tradeoffs associated with a few critical traits of individuals often form the basis for species turnover along the gradient. Among species that inhabit temporary ponds, distributions are often constrained because traits that enhance developmental rate and competitive ability also increase susceptibility to predators. In permanent ponds, changes in the composition of major predators over the gradient limit ...

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Mechanisms creating community structure across a freshwater habitat gradient

 – Among the species in the family Salmonidae, those represented by the genera Salmo, Salvelinus, and Oncorhynchus (subfamily Salmoninae) are the most studied. Here, various aspects of phenotypic and life-history variation of Atlantic salmon Salmo salar L., brown trout Salmo trutta L., and Arctic charr Salvelinus alpinus (L.) are reviewed. While many strategies and tactics are commonly used by these species, there are also differences in their ecology and population dynamics that result in a variety of interesting and diverse topics that are challenging for future research. Atlantic salmon display considerable phenotypic plasticity and variability in life-history characters ranging from fully freshwater resident forms, where females can mature at approximately 10 cm in length, to anadromous populations characterised by 3–5 sea-winter (5SW) salmon. Even within simple 1SW populations, 20 or more spawning life-history types can be identified. Juveniles in freshwater can use both fluvial and lacustrine habitats for rearing, and while most smolts migrate to sea during the spring, fall migrations occur in some populations. At sea, some salmon undertake extensive oceanic migrations while other populations stay within the geographical confines of areas such as the Baltic Sea. At the other extreme are those that reside in estuaries and return to freshwater to spawn after spending only a few months at sea. The review of information on the diversity of life-history forms is related to conservation aspects associated with Atlantic salmon populations and current trends in abundance and survival. Brown trout is indigenous to Europe, North Africa and western Asia, but was introduced into at least 24 countries outside Europe and now has a world-wide distribution. It exploits both fresh and salt waters for feeding and spawning (brackish), and populations are often partially migratory. One part of the population leaves and feeds elsewhere, while another part stays as residents. In large, complex systems, the species is polymorphic with different size morphs in the various parts of the habitat. Brown trout feed close to the surface and near shore, but large individuals may move far offshore. The species exhibits ontogenetic niche shifts partly related to size and partly to developmental rate. They switch when the amount of surplus energy available for growth becomes small with fast growers being younger and smaller fish than slow growers. Brown trout is an opportunistic carnivore, but individuals specialise at least temporarily on particular food items; insect larvae are important for the young in streams, while littoral epibenthos in lakes and fish are most important for large trout. The sexes differ in resource use and size. Females are more inclined than males to become migratory and feed in pelagic waters. Males exploit running water, near-shore and surface waters more than females. Therefore, females feed more on zooplankton and exhibit a more uniform phenotype than males. The Arctic charr is the northernmost freshwater fish on earth, with a circumpolar distribution in the Holarctic that matches the last glaciation. Recent mtDNA studies indicate that there are five phylogeographic lineages (Atlantic, Arctic, Bering, Siberian and Acadian) that may be of Pleistocene origin. Phenotypic expression and ecology are more variable in charr than in most fish. Weights at maturation range from 3 g to 12 kg. Population differences in morphology and coloration are large and can have some genetic basis. Charr live in streams, at sea and in all habitats of oligotrophic lakes, including very deep areas. Ontogenetic habitat shifts between lacustrine habitats are common. The charr feed on all major prey types of streams, lakes and near-shore marine habitats, but has high niche flexibility in competition. Cannibalism is expressed in several cases, and can be important for developing and maintaining bimodal size distributions. Anadromy is found in the northern part of its range and involves about 40, but sometimes more days in the sea. All charr overwinter in freshwater. Partial migration is common, but the degree of anadromy varies greatly among populations. The food at sea includes zooplankton and pelagic fish, but also epibenthos. Polymorphism and sympatric morphs are much studied. As a prominent fish of glaciated lakes, charr is an important species for studying ecological speciation by the combination of field studies and experiments, particularly in the fields of morphometric heterochrony and comparative behaviour.

https://onlinelibrary.wiley.com/doi/pdf/10.1034/j.1600-0633.2003.00010.x

Atlantic salmon Salmo salar L., brown trout Salmo trutta L. and Arctic charr Salvelinus alpinus (L.): a review of aspects of their life histories

Chloride concentrations are increasing at a rate that threatens the availability of fresh water in the northeastern United States. Increases in roadways and deicer use are now salinizing fresh waters, degrading habitat for aquatic organisms, and impacting large supplies of drinking water for humans throughout the region. We observed chloride concentrations of up to 25% of the concentration of seawater in streams of Maryland, New York, and New Hampshire during winters, and chloride concentrations remaining up to 100 times greater than unimpacted forest streams during summers. Mean annual chloride concentration increased as a function of impervious surface and exceeded tolerance for freshwater life in suburban and urban watersheds. Our analysis shows that if salinity were to continue to increase at its present rate due to changes in impervious surface coverage and current management practices, many surface waters in the northeastern United States would not be potable for human consumption and would become toxic to freshwater life within the next century.

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Increased salinization of fresh water in the northeastern United States

IN no domain of zoological science have the con tributions of American naturalists been more splendid than in that of vertebrate palaeontology, and in the list of pioneer workers in this domain an honoured place will always be accorded to Samuel Wendell Williston, who, born and nurtured in obscurity, had by the time of his death in 1918 won his way to recogni tion as the chief authority upon the extinct amphibians and reptiles. Apart from his specialist papers upon fossil vertebrates and upon modern dipterous flies, Williston was well known to the general zoologist for his excellent book on “Water Reptiles of the Past and Present,” published in 1914, and during the last year of his life he was busily engaged in the preparation of a second work of a general character on “The Reptiles of the World, Recent and Fossil.” The volume now under review comprises the main part of this general work, so far as it had been completed at the time of the author's death, put into shape and edited at the competent hands of Prof. W. K. Gregory. The Osteology of the Reptiles. By Prof. Samuel Wendell Williston. Arranged and edited by Prof. William King Gregory. Pp. xiii + 300. (Cambridge, Mass.: Harvard University Press; London: Oxford University Press, 1925.) 18s. 6d. net.

The Osteology of the Reptiles

The paper considers the extent to which salinity determines the structure of biological communities (composition and species richness and diversity) in saline lakes, ie inland bodies of water with salinities in excess of 3 g l-1 It also considers the extent to which oxygen, ionic composition, pH, hydrological patterns (degree of permanence and impermanence of water), geographical position, palaeoclimatic events, chance, human intervention, and biological interactions especially predation determine biological communities in salt lakes It suggests that salinity is less significant as a determinant of community structure in salt lakes than has been assumed

Salinity as a determinant of the structure of biological communities in salt lakes

Many inland waters are becoming more saline from human activities, particularly in semi-arid and arid regions. The causes and distribution of anthropogenic salinisation, the salinisation of freshwater lakes, rivers and streams, and increases in the salinity of large, permanent saline lakes are discussed. The impacts of anthropogenic salinisation are far-reaching, increasing, deleterious and largely irreparable. Environmental, social and environmental costs are high. Attention is drawn to the importance of anthropogenic salinisation and its impacts. The need for better recognition of the costs of salinisation and for more effective management is stressed.

Anthropogenic salinisation of inland waters

High and often variable salinity is an obvious feature of salt lakes. Correspondingly, salinity is usually assumed to be an important ecological determinant in such lakes. An investigation of the macroinvertebrate fauna of 79 lakes (salinities from 0.3 to 343 g 1−1) in the Western District of Victoria, Australia, examined this assumption. Over the total range of salinity, species richness and composition are highly correlated with salinity. However, these relationships become nonsignificant over intermediate ranges of salinity. Furthermore, many taxa have very broad tolerances to salinity at these intermediate ranges, implying that factors other than salinity may determine their distribution. An appreciation of scale (that is, the range of salinity over which observations are considered) resolves the paradox that, despite these broad tolerances by most taxa, species richness and composition strongly reflect salinity over the entire salinity range.

Salinity as a determinant of salt lake fauna: a question of scale

Despite the obvious ubiquity, ecological importance and limnological interest of temporary waters, most limnological texts have little to say about them. Dussart (1966), Schwoerbel (1974), Golterman (1975), Moss (1980), Goldman & Horne (1983), and Wetzel (1983), to mention just a few recent and otherwise comprehensive texts of North American or European provenance, make scant if any reference. Cole’s (1983) text does better, but only those authors who live in or have lived in tropical or arid countries make extensive reference (e.g. Bayly & Williams, 1973; Beadle, 1981). Temporary waters are especially widespread in arid and semi-arid regions. Even the most cursory glance at maps of world climate will indicate the vast extent of these regions in Africa, Australia, the Middle East and Central Asia, parts of South America, and in W. North America. The paucity of permanent waters in arid and semi-arid regions means, of course, that temporary waters have an ecological significance much greater there than in more temperate regions. For example, throughout most of continental Australia, waterfowl are closely dependent for survival on the occurrence of temporary water-bodies (Frith, 1967). The limnological interest of temporary waters relates above all to the nature of the ecological stresses that assail their biota, and the nature of corresponding adaptation.

Biotic adaptations in temporary lentic waters, with special reference to those in semi-arid and arid regions

Australia is the world's driest inhabited continent. Water is our limiting resource. It might therefore be thought that our water resources would be the subject of the most intensive study. Certain aspects, it must be conceded, have received much attention, notably the availability of water in terms of actual quantity. The size of the surface water and the groundwater resource is well understood and indeed receives about as much study as can reasonably be expected in a country with as sparse a population and level of scientific manpower as ours. Although the importance of understanding the water resource in terms of quantity is widely accepted, what has not been generally appreciated is that for this resource to be 'available' to human society for all the different uses to which it is put, it is not sufficient that there exists within easy reach of the end users a certain total volume of water. For that water to fulfil its functions-for agriculture, industry, the home, recreation, biological conservation-it must be in a certain state: it must conform to certain chemical, physical and biological criteria, and what has not been sufficiently appreciated in Australian society is that the condition a water is in depends very much on the ecology of the waterbody in which it resides. There are waterbodies in the world, for example high-altitude glacial lakes, which are naturally so pristine that their water could be used for any purpose without treatment.

William David Williams

Papers

Salinity as a determinant of the structure of biological communities in salt lakes

Anthropogenic salinisation of inland waters

Salinity as a determinant of salt lake fauna: a question of scale

Biotic adaptations in temporary lentic waters, with special reference to those in semi-arid and arid regions

Limnology in Australia