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

Freshwater biodiversity is the over-riding conservation priority during the International Decade for Action - 'Water for Life' - 2005 to 2015. Fresh water makes up only 0.01% of the World's water and approximately 0.8% of the Earth's surface, yet this tiny fraction of global water supports at least 100000 species out of approximately 1.8 million - almost 6% of all described species. Inland waters and freshwater biodiversity constitute a valuable natural resource, in economic, cultural, aesthetic, scientific and educational terms. Their conservation and management are critical to the interests of all humans, nations and governments. Yet this precious heritage is in crisis. Fresh waters are experiencing declines in biodiversity far greater than those in the most affected terrestrial ecosystems, and if trends in human demands for water remain unaltered and species losses continue at current rates, the opportunity to conserve much of the remaining biodiversity in fresh water will vanish before the 'Water for Life' decade ends in 2015. Why is this so, and what is being done about it? This article explores the special features of freshwater habitats and the biodiversity they support that makes them especially vulnerable to human activities. We document threats to global freshwater biodiversity under five headings: overexploitation; water pollution; flow modification; destruction or degradation of habitat; and invasion by exotic species. Their combined and interacting influences have resulted in population declines and range reduction of freshwater biodiversity worldwide. Conservation of biodiversity is complicated by the landscape position of rivers and wetlands as 'receivers' of land-use effluents, and the problems posed by endemism and thus non-substitutability. In addition, in many parts of the world, fresh water is subject to severe competition among multiple human stakeholders. Protection of freshwater biodiversity is perhaps the ultimate conservation challenge because it is influenced by the upstream drainage network, the surrounding land, the riparian zone, and - in the case of migrating aquatic fauna - downstream reaches. Such prerequisites are hardly ever met. Immediate action is needed where opportunities exist to set aside intact lake and river ecosystems within large protected areas. For most of the global land surface, trade-offs between conservation of freshwater biodiversity and human use of ecosystem goods and services are necessary. We advocate continuing attempts to check species loss but, in many situations, urge adoption of a compromise position of management for biodiversity conservation, ecosystem functioning and resilience, and human livelihoods in order to provide a viable long-term basis for freshwater conservation. Recognition of this need will require adoption of a new paradigm for biodiversity protection and freshwater ecosystem management - one that has been appropriately termed 'reconciliation ecology'.

https://www.cambridge.org/core/services/aop-cambridge-core/content/view/96AA265E3D88C95FB802E7A73CCB4332/S1464793105006950a.pdf/div-class-title-freshwater-biodiversity-importance-threats-status-and-conservation-challenges-div.pdf

Freshwater biodiversity: importance, threats, status and conservation challenges

H umans have long been fascinated by the dynamism of free-flowing waters. Yet we have expended great effort to tame rivers for transportation, water supply, flood control, agriculture, and power generation. It is now recognized that harnessing of streams and rivers comes at great cost: Many rivers no longer support socially valued native species or sustain healthy ecosystems that provide important goods and services (Naiman et al. 1995, NRC 1992).

/pdf/the-natural-flow-regime-3j2upj9j41.pdf

The Natural Flow Regime

▪ Abstract Local habitat and biological diversity of streams and rivers are strongly influenced by landform and land use within the surrounding valley at multiple scales. However, empirical associations between land use and stream response only varyingly succeed in implicating pathways of influence. This is the case for a number of reasons, including (a) covariation of anthropogenic and natural gradients in the landscape; (b) the existence of multiple, scale-dependent mechanisms; (c) nonlinear responses; and (d) the difficulties of separating present-day from historical influences. Further research is needed that examines responses to land use under different management strategies and that employs response variables that have greater diagnostic value than many of the aggregated measures in current use. In every respect, the valley rules the stream.      H.B.N. Hynes (1975)

http://deq.idaho.gov/media/525804-Landscapes_Riverscapes_Influence_landuse_stream_ecosystems_Allan_2004.pdf

Landscapes and Riverscapes: The Influence of Land Use on Stream Ecosystems

The flow regime is regarded by many aquatic ecologists to be the key driver of river and floodplain wet- land ecosystems. We have focused this literature review around four key principles to highlight the important mech- anisms that link hydrology and aquatic biodiversity and to illustrate the consequent impacts of altered flow regimes: Firstly, flow is a major determinant of physical habitat in streams, which in turn is a major determinant of biotic com- position; Secondly, aquatic species have evolved life history strategies primarily in direct response to the natural flow regimes; Thirdly, maintenance of natural patterns of longitu- dinal and lateral connectivity is essential to the viability of populations of many riverine species; Finally, the invasion and success of exotic and introduced species in rivers is facilitated by the alteration of flow regimes. The impacts of flow change are manifest across broad taxonomic groups including riverine plants, invertebrates, and fish. Despite growing recognition of these relationships, ecologists still struggle to predict and quantify biotic responses to altered flow regimes. One obvious difficulty is the ability to distin- guish the direct effects of modified flow regimes from im- pacts associated with land-use change that often accom- panies water resource development. Currently, evidence about how rivers function in relation to flow regime and the flows that aquatic organisms need exists largely as a series of untested hypotheses. To overcome these problems, aquatic science needs to move quickly into a manipulative or experimental phase, preferably with the aims of restora- tion and measuring ecosystem response.

/pdf/basic-principles-and-ecological-consequences-of-altered-flow-4u5bupofua.pdf

Basic principles and ecological consequences of altered flow regimes for aquatic biodiversity.

Year-round collections of mayflies (Ephemeroptera) from a Colorado mountain stream allowed critical examination of several methods of calculating production for species with different life cycles. Five of the six numerically dominant species exhibited slow seasonal, univoltine life cycles. Baetis tricaudatus was bivoltine. Two species demonstrated well synchronized development, three species were poorly synchronized and a sixth was intermediate. Mean density and biomass data from each sampling date were used to ascertain the goodness-of-fit of each species to the Allen curve. It is proposed that such information can provide quantitative criteria for identifying species with well synchronized development and thereby determine when it is appropriate to directly apply cohort methods while avoiding time intensive body size (e.g. head width) measurements necessary for size-frequency analyses. In addition, these data demonstrate that species specific production varies with gross changes in elevation.

Mayfly production in a Colorado mountain stream: an assessment of methods for synchronous and non-synchronous species

(2002). Spatial patterns of Ephemeroptera, Plecoptera and Trichoptera diversity in fragmented alpine streams. SIL Proceedings, 1922-2010: Vol. 28, No. 3, pp. 1429-1435.

Spatial patterns of Ephemeroptera, Plecoptera and Tricoptera diversity in fragmented alpine streams

H. B. N. Hynes (1970, 1975) has eloquently established stream ecology as a fundamental subdiscipline of limnology. In general terms he describes a river as an expression of the valley through which it flows; production of carbon in lotic habitats is greatly influenced by input of allochthonous nutrients and detritus from the drainage basin. The degree of heterotrophy and/or autotrophy in stream communities is a function of this allochthonous input and the extent of canopy development, which influences the amount of solar energy reaching primary producer biomass in the water. A natural river system may be conceptually perceived as an ecological continuum in which the P/R ratio (i.e., the relationship between community production and respiration) changes from much less than unity in heavily canopied reaches of small headwater tributaries to slightly greater than unity in the slow, meandering areas of the river near its mouth. This idea is being developed in collaborative, holistic studies in very different geographical areas for the purpose of defining more satisfactory methods of quantifying ecological processes and providing ultimate comparative data on lotic ecosystems (Cummins, 1975; this volume).

Dammed Rivers of the World: Symposium Rationale

The initial hypothesis that predation pressure should decrease with decreasing pH in aquatic macrobenthic communities if predatory invertebrates are more sensitive to water acidification than prey invertebrates is tested. Short-term toxicity bioassays were conducted in soft water (average value of total hardness 38.0 mg CaCO3/L) to determine the differential sensitivity of the predator, Dugesia dorotocephala (Turbellaria, Tricladida), and the prey, larvae of Cheumatopsyche pettiti (Insecta, Trichoptera), to low pH. Test pH solutions were prepared with sulfuric acid (H2SO4). Test species were also exposed to high concentrations of sulfate ion (95 mg SO4=/L for D. dorotocephala and 340 mg SO4=/L for C. pettiti) as sulfate toxicity controls, using potassium sulfate (K2SO4). No mortality was observed during these toxicity controls, indicating that toxic effects generated by low pH were fundamentally due to H+ ions. The 72 and 96-h LC50s (as pH values) and their 95% confidence limits were 4.88 (4.72–5.05) and 5.04 (4.89–5.21) for D. dorotocephala, and 3.25 (3.00–3.51) and 3.48 (3.24–3.73) for C. pettiti. Net-spinning caddisfly larvae migrated from their retreat nets and protruded their anal papillae before dying. After short-term bioassays, predation-pressure laboratory experiments were performed for 6 days. The cumulative mortality of C. pettiti by predation of D. dorotocephala decreased with decreasing sublethal pH values. The average predation rates at mean pH values of 7.7, 7.7, 6.6, 6.5, 6.2 and 6.0 were 2.5, 2.0, 1.33, 1.17, 0.67 and 0.33 larvae/day, respectively. The major biotic factor affecting predation pressure appears to be the reduction in the physiological activity of triclads at low pH. It is concluded that predation pressure can decrease in aquatic macrobenthic communities if prey are more tolerant to water acidification than predators.

Differential sensitivity of Dugesia dorotocephala and Cheumatopsyche pettiti to water acidification: ecological implication for predator-prey interactions.

Spatial and seasonal patterns in the densities and species richness of Simuliidae (blackflies) were examined in the glacier-fed Roseg River, Switzerland. We also investigated how selected environmental factors were related to the observed community structure. Overall, 7 blackfly species were found in streams of this glacial flood plain. Total densities and species richness differed significantly among sampling dates, and densities showed a significant downstream increase. Non-metric multidimensional scaling supported these findings and suggested that seasonal changes were affected strongly by extent of glacial influence. Furthermore, our results were indicative that spatio-temporal habitat heterogeneity ameliorated the negative effects of high discharge conditions in summer. This resulted in enhanced overall ecosystem stability in this glacier-fed stream ecosystem, and enabled biotic assemblages to sustain populations under the harsh environmental conditions experienced at this time of the year. Low temperatures and channel instability limited Simuliidae colonization close to the glacier terminus. Suspended particles represented the most important food source for simuliids in the Roseg River.

James V. Ward

Papers

Mayfly production in a Colorado mountain stream: an assessment of methods for synchronous and non-synchronous species

Spatial patterns of Ephemeroptera, Plecoptera and Tricoptera diversity in fragmented alpine streams

Dammed Rivers of the World: Symposium Rationale

Differential sensitivity of Dugesia dorotocephala and Cheumatopsyche pettiti to water acidification: ecological implication for predator-prey interactions.

Diversity, distribution and seasonality of the Simuliidae fauna in a glacial stream system in the Swiss Alps