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

Fresh waters are particularly vulnerable to climate change because (i) many species within these fragmented habitats have limited abilities to disperse as the environment changes; (ii) water temperature and availability are climate-dependent; and (iii) many systems are already exposed to numerous anthropogenic stressors. Most climate change studies to date have focused on individuals or species populations, rather than the higher levels of organization (i.e. communities, food webs, ecosystems). We propose that an understanding of the connections between these different levels, which are all ultimately based on individuals, can help to develop a more coherent theoretical framework based on metabolic scaling, foraging theory and ecological stoichiometry, to predict the ecological consequences of climate change. For instance, individual basal metabolic rate scales with body size (which also constrains food web structure and dynamics) and temperature (which determines many ecosystem processes and key aspects of foraging behaviour). In addition, increasing atmospheric CO2 is predicted to alter molar CNP ratios of detrital inputs, which could lead to profound shifts in the stoichiometry of elemental fluxes between consumers and resources at the base of the food web. The different components of climate change (e.g. temperature, hydrology and atmospheric composition) not only affect multiple levels of biological organization, but they may also interact with the many other stressors to which fresh waters are exposed, and future research needs to address these potentially important synergies.

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Climate change and freshwater ecosystems: impacts across multiple levels of organization

Ecological studies often require studying the common structure of a pair of data tables. Co-inertia analysis is a multivariate method for coupling two tables. It is often neglected by ecologists who prefer the widely used methods of redundancy analysis and canonical correspondence analysis. We present the co-inertia criterion for measuring the adequacy between two data sets. Co-inertia analysis is based on this criterion as are canonical correspondence analysis or canonical correlation analysis, but the latter two have additional constraints. Co-inertia analysis is very flexible and allows many possibilities for coupling. Co-inertia analysis is suitable for quantitative and/or qualitative or fuzzy environmental variables. Moreover, various weighting of sites and various transformations and/or centering of species data are available for this method. Hence, more ecological considerations can be taken into account in the statistical procedures. Moreover, the principle of this method is very general and can be easily extended to the case of distance matrices or to the case of more than two tables. Simulated ecological data are used to compare the co-inertia approach with other available methods.

Co-inertia analysis and the linking of ecological data tables

Summary

1. Relationships between probabilities of occurrence for fifteen diadromous fish species and environmental variables characterising their habitat in fluvial waters were explored using an extensive collection of distributional data from New Zealand rivers and streams. Environmental predictors were chosen for their likely functional relevance, and included variables describing conditions in the stream segment where sampling occurred, downstream factors affecting the ability of fish to move upriver from the sea, and upstream, catchment-scale factors mostly affecting variation in river flows.



2. Analyses were performed using multivariate adaptive regression splines (MARS), a technique that uses piece-wise linear segments to describe non-linear relationships between species and environmental variables. All species were analysed using an option that allows simultaneous analysis of community data to identify the combination of environmental variables best able to predict the occurrence of the component species. Model discrimination was assessed for each species using the area under the receiver operating characteristic curve (ROC) statistic, calculated using a bootstrap procedure that estimates performance when predictions are made to independent data.



3. Environmental predictors having the strongest overall relationships with probabilities of occurrence included distance from the sea, stream size, summer temperature, and catchment-scale drivers of variation in stream flow. Many species were also sensitive to variation in either the average and/or maximum downstream slope, and riparian shade was an important predictor for some species.



4. Analysis results were imported into a Geographic Information System where they were combined with extensive environmental data, allowing spatially explicit predictions of probabilities of occurrence by species to be made for New Zealand's entire river network. This information will provide a valuable context for future conservation management in New Zealand's rivers and streams.

Using multivariate adaptive regression splines to predict the distributions of New Zealand's freshwater diadromous fish

www.freshwaterecology.info – An online tool that unifies, standardises and codifies more than 20,000 European freshwater organisms and their ecological preferences ☆

1. The influence of 11 environmental variables on benthic macroinvertebrate communities was examined in seven glacier-fed European streams ranging from Svalbard in the north to the Pyrenees in the south. Between 4 and 11 near-pristine reaches were studied on each stream in 1996–97.

2. Taxonomic richness, measured at the family or subfamily (for Chironomidae) levels for insects and higher levels for non-insects, increased with latitude from Svalbard (3 taxa) to the Pyrenees (29 taxa).

3. A Generalized Additive Model (GAM) incorporating channel stability [Pfankuch Index (PFAN)], tractive force, Froude number (FROU), water conductivity (COND), suspended solids (SUSP) concentration, and maximum temperature explained 79% of the total deviance of the taxonomic richness per reach. Water temperature and the PFAN of stability made the highest contribution to this deviance. In the model, richness response to temperature was positive linear, whereas the response to the PFAN was bell-shaped with an optimum at an intermediate level of stability.

4. Generalized Additive Models calculated for the 16 most frequent taxa explained between 25 (Tipulidae) and 79% (Heptageniidae) of the deviance. In 10 models, more than 50% of the deviance was explained and 11 models had cross-validation correlation ratios above 0.5. Maximum temperature, the PFAN, SUSP and tractive force (TRAC) were the most frequently incorporated explanatory variables. Season and substrate characteristics were very rarely incorporated.

5. Our results highlight the strong deterministic nature of zoobenthic communities in glacier-fed streams and the prominent role of water temperature and substrate stability in determining longitudinal patterns of macroinvertebrate community structure. The GAMs are proposed as a tool for predicting changes of zoobenthic communities in glacier-fed streams under climate or hydrological change scenarios.

Macrobenthic invertebrate richness and composition along a latitudinal gradient of European glacier‐fed streams

1. A study on glacial stream ecosystems was carried out in six regions across Europe, from Svalbard to the French Pyrenees. The main aim was to test the validity of the conceptual model of Milner & Petts (1994) with regard to the zonation of chironomids of glacier-fed rivers along altitudinal and latitudinal gradient. 2. Channel stability varied considerably, both on the latitudinal and altitudinal scale, being lowest in the northern regions (Svalbard, Iceland and Norway) and the Swiss Alps. Water temperature at the upstream sites was always <2 °C. 3. There was a prominent difference in taxonomic richness between the Alpine and the northern European regions, with a higher number of taxa in the south. In all regions, the chironomid community was characterized by the genus Diamesa and the subfamily Orthocladiinae. Of a total of 63 taxa recorded, two (Diamesa bertrami and Orthocladius frigidus) were common in all the regions except Svalbard. 4. On the basis of cluster analysis, seven distinct groups of sites were evident amongst glacial-fed systems of the ®ve regions (Pyrenees excluded). This classi®cation separated the glacier-fed streams on geographical, latitudinal and downstream gradients. 5. Canonical Correspondence Analysis (CCA) of environmental variables was carried out using 41 taxa at 105 sites. Slope, water depth, distance from source, water temperature and the Pfankuch channel stability index were found to be the major explanatory environmental variables. The analysis separated Diamesinae and typical upstream orthoclads from the other chironomids by low temperature and high channel instability. 6. In all six regions, Diamesa was present closest to the glacier. Within 200 m of the glacier snout, other genera of Diamesinae were found together with Orthocladiinae. Pioneer taxa like Diamesa species coexisted with later colonizers like Eukiefferiella minor/®ttkaui in relatively unstable channels. 7. The longitudinal succession of chironomid assemblages across altitudinal and latitudinal gradients in glacial streams followed the same pattern, with similar genera and groups of species. The general aspects of the conceptual model of Milner & Petts (1994) were supported. However, Diamesa species have wider temperature limits than predicted and other Diamesinae as well as Orthocladiinae colonize metakryal habitats. Correspondence: Brigitte

Chironomid (Diptera: Chironomidae) communities in six European glacier-fed streams

1. Benthic macroinvertebrate distribution was examined in relation to channel characteristics (including stability), substratum, hydraulic variables, primary production (chlorophylla) and coarse particular organic matter (CPOM) in an alpine glacial stream, the Mutt (Upper Rhone valley, Switzerland). Co-inertia analysis and canonical correspondence analysis were used to identify the major environmental gradients in¯uencing community variations. 2. The Mutt (length: 3.6 km, altitudinal range: 1800±3099 m a.s.l.) exhibited typical characteristics of a kryal stream. Average summer temperature remained below 2 °C immediately downstream from the snout but was on average 5 °C higher 1700 m downstream. Seasonal variations in water sources were evidenced by the high latesummer(September) contribution of groundwater with increased conductivity. 3. Sixty-six taxa were recorded from the five reaches sampled at three periods (snowmelt,ice melt and low water in late summer) in 1996 and 1997, of which 29 were Chironomidae. Three taxa of Diamesinae were the first colonizers of the stream below the glacier, but 16 taxa, including Ephemeroptera, Plecoptera and Trichoptera, were already recorded 200 m downstream. Water depth, channel slope and Pfankuch's Index of channel stability were strongly correlated with the longitudinal faunal gradient. Maximum temperature, current velocity and water conductivity were also correlated, but to a lesser extent. 4. The rapid incorporation of non-chironomid taxa into the stream community represented a departure from Milner & Petts's (1994) conceptual model of invertebrate succession downstream of glacial margins. The results confirmed that glacial stream communities are primarily driven by physical determinants.

Macroinvertebrate community structure in relation to environmental variables in a Swiss glacial stream

Changes in the chironomid communities in Lake Geneva in relation with eutrophication, over a period of 60 years

The study of the invertebrate assemblages in a small glacial catchment of the Swiss Alps (the Mutt) was designed to categorise sites in relation to their water sources and assess the contribution of small headwater tributaries to the biodiversity of such Alpine catchments. Physico-chemical characteristics and macroinvertebrate drift were monitored in seven streams during three contrasting hydrological periods in June. August and September 1997. Physico-chemical characteristics clearly differentiated kryal (glacial) and krenal (groundwater-fed) streams. The glacial waters showed a broader amplitude in environmental variation, depending on altitude, geology and season. Macroinvertebrate drift composition was significantly correlated with the environmental conditions. In kryal streams, where harsh conditions prevailed, diversity and abundance were low, while the krenal streams showed highly diversified and dense macroinvertebrate communities, favoured by physico-chemical constancy. However, the low number of taxa found at the same time in these groundwater-fed tributaries and in the main Mutt stream indicated a low contribution from these small streams to the communities of the main stream. Nevertheless, the habitat heterogeneity generated by different water origins in the tributaries favoured the coexistence of diversified macroinvertebrate communities in a rather small catchment area. Therefore these tributaries played an important part in the biodiversity of the glacial catchment.

Brigitte Lods-Crozet

Papers

Macrobenthic invertebrate richness and composition along a latitudinal gradient of European glacier‐fed streams

Chironomid (Diptera: Chironomidae) communities in six European glacier-fed streams

Macroinvertebrate community structure in relation to environmental variables in a Swiss glacial stream

Changes in the chironomid communities in Lake Geneva in relation with eutrophication, over a period of 60 years

Invertebrate drift and physico-chemical variables in the tributaries of the Mutt, a Swiss glacial stream