Thorsten Blenckner

Regional and local impact on species diversity - from pattern to processes

Abstract. The impact of regional factors (such as speciation or dispersal) on the species richness in local communities (SL) has received increasing attention. A prominent method to infer the impact of regional factors is the comparison of species richness in local assemblages (SL) with the total number of species in the region (SR). Linear relations between SR and SL have been interpreted as an indication of strong regional influence and weak influence of interactions within local communities. We propose that two aspects bias the outcome of such comparisons: (1) the spatial scale of local and regional sampling, and (2) the body size of the organisms. The impact of the local area reflects the scales of ecological interactions, whereas the ratio between local and regional area reflects the inherent moment of autocorrelation. A proposed impact of body size on the relation is based on the high dispersal and high abundance of small organisms. We predict strongest linearity between SR and SL for large organisms, for large local areas (less important ecological interactions) and for sampling designs where the local habitat area covers a high proportion of the regional area (more important autocorrelation). We conducted a meta-analysis on 63 relations obtained from the literature. As predicted, the linearity of the relationship between SL and SR increased with the proportion of local to regional sampling area. In contrast, neither the body size of the organisms nor the local area itself was significantly related to the relation between SL and SR. This indicated that ecological interactions played a minor role in the shape of local to regional richness plots, which instead was mainly influenced by the sampling design. We found that the studies published so far were highly biased towards larger organisms and towards high similarity between the local and regional area. The proposed prevalence of linear relationships may thus be an artefact and plots of SL to SR are not a suitable tool with which to infer the strength of local interactions.

A conceptual model of climate-related effects on lake ecosystems

Climatic variation and change affect the dynamics of organisms and ecosystem processes. Many studies in the past have analyzed and discussed various climate-driven effects on different components of the lake ecosystem. Only a few synthesis papers have been published in this field. In this overview, a conceptual model has been developed to help explain why lakes respond individually to climate. The model consists of two main components, a so-called Landscape Filter comprising the features of geographical position, catchment characteristics and lake morphology, and a so-called Internal Lake Filter, comprising the features of lake history and biotic/abiotic interactions. The application of this conceptual model on published literature findings illustrates the strength in this encompassing perspective. An assessment of current climate research methods is presented with some perspectives given.

Seasonality of chlorophyll and nutrients in Lake Erken – effects of weather conditions

The seasonality of nutrients and chlorophyll a in Lake Erken (Sweden) was monitored during 1994 to 2001 (warm period) and compared to the time period 1975 to 1979 (cold period). The coupling to weather conditions and potentially influencing factors, such as water temperature, light conditions, and stratification were evaluated. During the warm period the ice cover period was considerably shorter and the ice breakup about one month earlier than in the cold period. The decrease in thickness and duration of snow cover resulted in considerably better light conditions during winter, favouring phytoplankton growth under the ice. The nitrate concentrations were much lower in late winter during the warm period. There were elevated phosphate and ammonium concentrations in the hypolimnion during August. Significantly higher phosphate and chlorophyll a levels were noticed in autumn during the warm period, compared to the cold period.

The Impact of the Changing Climate on the Thermal Characteristics of Lakes

Meteorological forcing at the air-water interface is the main determinant of the heat balance of most lakes (Edinger et al., 1968; Sweers, 1976). Year-to-year changes in the weather therefore have a major effect on the thermal characteristics of lakes. However, lakes that differ with respect to their morphometry respond differently to these changes (Gorham, 1964), with deeper lakes integrating the effects of meteorological forcing over longer periods of time. Other important factors that can influence the thermal characteristics of lakes include hydraulic residence time, optical properties and landscape setting (e.g. Salonen et al., 1984; Fee et al., 1996; Livingstone et al., 1999). These factors modify the thermal responses of the lake to meteorological forcing (cf. Magnuson et al., 2004; Blenckner, 2005) and regulate the patterns of spatial coherence (Chapter 17) observed in the different regions (Livingstone, 1993; George et al., 2000; Livingstone and Dokulil, 2001; Jarvinen et al., 2002; Blenckner et al., 2004)

The Impact of Variations in the Climate on Seasonal Dynamics of Phytoplankton

Phytoplankton, an assemblage of suspended, primarily autotrophic single cells and colonies, forms part of the base of the pelagic food chain in lakes. The responses of phytoplankton to anthropogenic pressures frequently provide the most visible indication of a long-term change in water quality. Several attributes related to the growth and composition of phytoplankton, such as their community structure, abundance as well as the frequency and the intensity of blooms, are included as indicators of water quality in the Water Framework Directive. The growth and seasonal succession of phytoplankton is regulated by a variety of external as well as internal factors (Reynolds et al., 1993; Reynolds, 2006). Among the most important external factors are light, temperature, and those associated with the supply of nutrients from point and diffuse sources in the catchment. The internal factors include the residence time of the lakes, the underwater light regime and the mixing characteristics of the water column. The schematic diagram (Fig. 14.1) shows some of the ways in which systematic changes in the climate can modulate these seasonal and inter-annual variations. The effects associated with the projected changes in the rainfall are likely to be most pronounced in small lakes with short residence times (see George et al., 2004 for some examples). In contrast, those connected with the projected changes in irradiance and wind mixing, are likely to be most important in deep, thermally stratified lakes.

Regional and Supra-Regional Coherence in Limnological Variables

Limnologists and water resources managers have traditionally perceived lakes as discrete geographical entities. This has resulted in a tendency for scientific lake studies to concentrate on lakes as individuals, with little connection either to each other or to large-scale driving forces. Since the 1990s, however, a shift in the prevailing paradigm has occurred, with lakes increasingly being seen as responding to regional, rather than local, driving forces. The seminal work on regional coherence in lake behaviour was that of Magnuson et al. (1990), who showed that many features of lakes within the same region respond coherently to drivers such as climate forcing and catchment processes. From this study it emerged that the degree of coherence among lakes is greatest for those properties most directly affected by climate forcing. Specifically, the physical properties of lakes tend to vary in a more coherent way than their chemical and biological properties (see also Kratz et al., 1998). Further overviews of the topics of coherence and climate-driven variability, focusing

Lake ice phenology

In Chapter 5 of this book, it is shown that the formation of ice on the surface of a lake (‘ice-on’) and its thawing and ultimate disappearance (‘ice-off’) are complex phenomena governed by mechanisms that involve many interacting meteorological (and some non-meteorological) forcing factors. Linking ice phenology – the timing of ice-on and ice-off – to climatic forcing might therefore be expected to be a difficult task. This task, however, is simplified considerably by the fact that air temperature is the dominant variable driving ice phenology (Williams, 1971; Ruosteenoja, 1986; Vavrus et al., 1996; Williams and Stefan, 2006), and is also correlated to some extent with other relevant meteorological driving variables such as solar radiation, relative humidity and snowfall.

The influence of calcium on the chlorophyll–phosphorus relationship and lake Secchi depths

The basic aim of this study was to analyse the influence of calcium on the Chl–TP relationship and to apply the findings to improve dynamic (mechanistically-based) modelling of phosphorus and lake eutrophication. We have analysed long-term data from 73 lakes. The influences of calcium found in these statistical analyses have been integrated into a dynamic foodweb model, the LakeWeb-model, which also includes a mass-balance model for phosphorus. Differences in the model outcome between simulations without and with considerations to the role of calcium are discussed. We can conclude that calcium is an important factor influencing both the Chl–TP relationship and Secchi depths in mesotrophic and eutrophic lakes. Our results also indicate that lakes with long-term median Ca-concentration between 10–30 mg/l function as hardwater lakes. The results also stress the importance of taking a holistic view of lakes since the bedrock, soils and land-use activities in the catchment influence the calcium concentration in lakes and therefore the phosphorus cycle, water clarity and the productivity of a given lake. The predictive power of the Chl–TP regression increases markedly if hardwater lakes are omitted from the model domain. For lake foodweb and mass-balance modelling, we show that the inclusion of the presented calcium moderator clearly improved the predictions of lake TP-concentrations in water and sediments, chlorophyll and Secchi depths in Lake Erken, a hardwater lake in Sweden.

The Impact of Climate Change on Lakes in Northern Europe

In Northern Europe, most lakes are characterized by extended periods of winter ice cover, high spring inflow from snow melt and brown water produced by the transport of dissolved organic carbon (DOC) from the surrounding catchments. In this chapter, the potential impact of climate change on the dynamics of these lakes is addressed by: (i) Describing the historical responses of the lakes to changes in the weather. (ii) Summarizing the results of modelling studies that quantify the impact of future changes in the climate on the lakes and the surrounding catchments. Many existing water quality problems could well be exacerbated by the effects of climatic change. It is therefore important to assess the holistic responses of the individual lakes to the combined effects of local changes in the catchment and regional changes in the weather (Hall et al., 1999; Anderson et al., 2005). Overall, the response of individual lakes to climate change can be very different (Blenckner et al., 2004). For example, mountain lake catchments are affected differently from those at lower altitudes. In addition, the landscape position of a particular lake influences hydrological flow regime (Kratz et al., 1997). Furthermore, the response of lakes to climatic variation is also modified by physical lake features such as morphometry and water clarity which, in turn, is also affected by the concentration of the dissolved organic carbon (see for example Fee et al., 1996). Also, the alignment of the lake in relation to the main wind direction is important for the timing of the ice break-up and mixing regime. Even, the environmental changes experienced by the lake in the past can affect the magnitude of the response to climatic variation. Lakes in a recovery phase from eutrophication, acidification, toxic components or any other strong human disturbance, might respond differently to climatic variability and change owing to their specific history and food web structure.

The Impact of the Changing Climate on the Supply and Re-Cycling of Phosphorus

For more than half a century, phosphorus (P) has been a major focus of limnological research. Early studies by Rodhe (1948) identified P as a key factor limiting the growth of algae. Recently, Istvanovics (2008) reviewed the cycling of phosphorus in lakes and its role in eutrophication. Historical increases in P loading have been observed in lakes distributed throughout Europe. In urban areas, the main factor responsible for the increase has been the influx of P from municipal and industrial point sources (Forsberg, 1987 In rural areas, the increase in loading has principally been due to changes in land-use and the increased use of artificial fertilizers (Forsberg, 1987). Over the years, water companies and local administrations have successfully reduced the quantities of P reaching lakes from point sources but reducing the load attributable to diffuse sources has proved more difficult. Another factor influencing the seasonal availability of P to lakes and reservoirs is the internal recycling of the element from both shallow and deep sediments. During the 1980s research on the role of internal P loading from lake sediments developed rapidly and new knowledge was gained about factors regulating this process in both shallow and deep lakes (Bostrom et al., 1982; Cullen and Forsberg, 1988). Mobilisation as well as transport processes were identified and the classical Einsele-Mortimer theory (Einsele, 1938; Mortimer, 1941; Mortimer, 1942) that explained the reduction of iron and the dissolution of phosphate under anaerobic conditions was expanded to include the microbial breakdown of organic phosphorus as a significant factor. More recently, increasing attention has been paid to the effects that year-to-year variations in the climate have on both the supply and the internal recycling of phosphorus.