Single species dynamics under climate change

We propose a general mathematical model describing the growth and dispersal of a single species living in a 1-D spatially discrete array of habitat patches affected by a sustained and directional change in climate. Our model accounts for two important characteristics of the climate change phenomenon: (1) Scale dependency: different species may perceive the change in the environment as occurring at different rates because they perceive the environment at different scales, and (2) measure dependency: different species measure the environment differently in the sense that they may be sensible to or cue in on different aspects of it (e.g., maximum temperature, minimum temperature, accumulated temperature) which is associated with their physiological, ecological, and life history attributes, which renders some characteristics of the environment more biologically relevant than others. We show that the deterioration in the quality of habitable patches as a consequence of climate change drives the species to extinction when dispersal is not possible; otherwise, we proof and provide a numerical example that, depending on the velocity of climate change, the scale at which a species measures it, and the particular attribute of the environment that is more biologically relevant to the species under analysis, there is always a migration strategy that allows the persistence of the species such that it tracks its niche conditions through space, thus shifting its geographic range. Our mathematical analysis provides a general framework to analyze species’ responses to climate change as a relational property of a given species in interaction with a change in climate. In particular, we can analyze the persistence of species by taking into account the ways in which they measure and filter the environment. Indeed, one of our main conclusions is that there is not a single climate change but many, as it depends on the interaction between a particular species and climate. Thus, the problem is more complex than assumed by analytically tractable models of species responses to climate change.     

Thermal stratification, nutrient dynamics, and phytoplankton productivity during the onset of spring phytoplankton growth in Lake Baikal, Russia

Lake Baikal, Russian Siberia, was sampled in July 1990 during the period of spring mixing and initiation of thermal stratification. Vertical profiles of temperature, dissolved nutrients (nitrate and soluble reactive phosphorus), phytoplankton biomass, and primary productivity were determined in an eleven-station transect encompassing the entire 636 km length of the lake. Pronounced horizontal variability in hydrodynamic conditions was observed, with the southern region of the lake being strongly thermally stratified while the middle and north basins were largely isothermal through July. The extent of depletion of surface water nutrients, and the magnitude of phytoplankton biomass and productivity, were found to be strongly correlated with the degree of thermal stratification. Horizontal differences likely reflected the contribution of two important factors: variation in the timing of ice-out in different parts of the lake (driving large-scale patterns of thermal stratification and other limnological properties) and localized effects of river inflows that may contribute to the preliminary stabilization of the water column in the face of intense turbulent spring mixing (driving meso-scale patterns). Examination of the relationships between surface water inorganic N and P depletion suggested that during the spring and early summer, phytoplankton growth in unstratified portions of the lake was largely unconstrained by nutrient supplies. As summer progressed, the importance of co-limitation by both N and P became more apparent. Uptake and regeneration rates, measured directly using the stable isotope ¹⁵N, revealed that phytoplankton in stratified portions of the lake relied primarily on NH₄ as their N source. Rates of NH₄ regeneration were in approximate equilibrium with uptake; both processes were dominated by organisms <2 µm. This pattern is similar to that observed for oligotrophic marine systems. Our study underscores the importance of hydrodynamic conditions in influencing patterns of biological productivity and nutrient dynamics that occur in Lake Baikal during its brief growing season.     

Spatial distribution of phytoplankton in Lake Baikal, Spring 1991

• 1 Studies carried out in Lake Baikal in late spring (late May-early June) 1991 showed marked differences in the species composition and abundance of phytoplankton in different regions of the lake. The south and north basins were characterized by small forms of algae. The middle basin, Maloe More and the shallow waters of the Selenga had species with large cells including Aulacoseira islandica subsp. helvetica, Dinobryon cylindricum and D. divergens.
• 2 Areas of high biomass were correlated with shallow waters and river inputs. This was especially apparent in the region of the Selenga delta. The vertical distribution of phytoplankton indicated the non-synchronous start of the spring homothermy throughout the lake. Phytoplankton were concentrated in the upper 100 m layer with subsurface maxima resulting from the sinking of large algae.
• 3 The concentration of phytoplankton biomass in general at this time characterized the lake as moderately productive.

     

Nutrient limitation of the primary production of phytoplankton in Lake Baikal

Nutrient limitation of the primary production of phytoplankton at some stations in southern and central Lake Baikal was studied by nutrient enrichment experiments in August 2002. Chlorophyll (Chl.) a concentrations ranged from 0.7 to 5.8μgl⁻¹. Inorganic nutrient concentrations were low: soluble reactive phosphorus ranged from 0.05 to 0.20μmoll⁻¹, ammonia from 0.21 to 0.41μmoll⁻¹, and nitrite plus nitrate from 0.33 to 0.37μmoll⁻¹. In the five enrichment experiments, phosphate spikes and phosphate plus nitrate spikes always stimulated primary production. Nitrate spikes also stimulated primary production in four of the experiments. Significant differences were detected between the controls and phosphate spikes and between the controls and phosphate plus nitrate spikes. Thus, the first limiting nutrient is thought to be phosphorus, but once phosphorus is supplied to the surface water, the limiting nutrient will quickly shift from phosphorus to nitrogen.     

An HPLC analysis of the summer phytoplankton assemblage in Lake Baikal

1. The enormous size and spatial heterogeneity of Lake Baikal require rapid methods for large sample sets. We therefore tested the applicability of a novel, high‐performance liquid chromatography (HPLC)‐based, combination of methods for analysing phytoplankton. In July 2001, samples were collected in a transect across the lake at various depths down to 30 m. Phytoplankton (>3 μm) and autotrophic picoplankton (APP) were counted under light and epifluorescence microscopes, respectively. Pigments were analysed with HPLC. 2. The pigment data allowed the contributions of the dominant phytoplankton groups to the total chlorophyll a (Chl a) in the lake to be estimated by multiple linear regression and by the CHEMTAX matrix factorisation program. Three marker pigments, fucoxanthin, lutein and zeaxanthin, were shown to be useful indicators of the abundance and spatial distribution of certain phytoplankton groups. The relative contributions of the various phytoplankton groups to the total Chl a in the lake determined using these marker pigments were similar, but not identical, to those determined by cell counts. 3. Pigment analyses of isolated strains from Lake Baikal and some European lakes confirmed that phycoerythrin‐containing Cyanobacteria with very high amounts of zeaxanthin were responsible for the low Chl a/zeaxanthin ratios of the water samples. A picoplanktonic species of Eustigmatophyceae was isolated from the lake. Its high violaxanthin content, responsible for very low Chl a/violaxanthin ratios of some water samples, can be used to estimate the contribution of this group to total Chl a.     

Long-term trends in seasonal plankton dynamics in Lake Mead (Nevada-Arizona,USA) and implications for climate change

Where biodiversity conservation and environmental preservation are significant concerns, rapid assessment and monitoring of the colonization and spread of non-native species are essential for timely decision-making and response. We developed and evaluated the adequacy of a rapid assessment protocol (RAP) for detecting non-native fish species in 74 Eastern Brazilian lakes. The RAP consists of a single field day employing two surveyors to conduct interviews with local sport fishers, visual surveys of fish, angling with lures, and gillnetting. We compared our results with those from separate, intense, large sampling effort (LSE) field surveys. Despite requiring less than 1/100th of the field effort, the RAP was able to detect the presence of most non-native fish species that were reported in the same lakes by LSE surveys. Information from local sport fishers was invaluable, particularly for detecting species that were in low abundance, but was not available for lakes within a forest preserve area. Non-native introductions commonly lead to rapid and widespread invasion and adverse effects on native biota. Our results strongly suggest that the RAP could function as a cost-effective tool for efficiently assessing the presence of non-native fishes in lakes and monitoring their colonization and spread over large geographic areas.     

Retrospective of diatom diversity dynamics in thanatocenoses under climate change

The species diversity of planktonic diatoms in thanatocenoses of bottom sediments in Lake Bol’shoe Miassovo (Southern Urals) has been studied and a reconstruction of the thermal regime of the lake for the last 1000 years is made. It is established that the restructuring of the diatom assemblage and decrease in species diversity in thanatocenoses are caused by rapid climate warming that began in the 19th century in the Southern Urals.     

The dynamics of food chains under climate change and nutrient enrichment

Warming has profound effects on biological rates such as metabolism, growth, feeding and death of organisms, eventually affecting their ability to survive. Using a nonlinear bioenergetic population-dynamic model that accounts for temperature and body-mass dependencies of biological rates, we analysed the individual and interactive effects of increasing temperature and nutrient enrichment on the dynamics of a three-species food chain. At low temperatures, warming counteracts the destabilizing effects of enrichment by both bottom-up (via the carrying capacity) and top-down (via biological rates) mechanisms. Together with increasing consumer body masses, warming increases the system tolerance to fertilization. Simultaneously, warming increases the risk of starvation for large species in low-fertility systems. This effect can be counteracted by increased fertilization. In combination, therefore, two main drivers of global change and biodiversity loss can have positive and negative effects on food chain stability. Our model incorporates the most recent empirical data and may thus be used as the basis for more complex forecasting models incorporating food-web structure.     

Photosynthetic potential of phytoplankton in the deep water of Lake Baikal,Russia

We examined the photosynthetic activity of the phytoplankton community collected from the surface to a depth of 1000m in the south basin of Lake Baikal. Experiments were conducted in June (mixing period) and August (stratified period). The carbon fixation rate was measured by the use of a ¹³C tracer after the incubation of samples under light conditions in the upper water column. Photosynthetic fixation of ¹³C was detected for samples collected from a depth of 500m, indicating the viability of phytoplankton in deep water. The community composition was dominated by Bacillariophyceae in deep water. The finding of lower activity at a depth of 200m than that at a depth of 500m in August suggests that the spring diatom bloom could be a significant source of viable populations at a depth of 500m. Photosynthetic activity was not detected in samples collected at a depth of 1000m.     

Plant community dynamics in a calcareous grassland under climate change manipulations

This study investigates the effects of field manipulations of local climate to determine the potential impact of climate change on plant community dynamics in a calcareous grassland. The experimental site is located in a grassland at the Wytham estate, Oxfordshire, UK. The one hectare study area is within a 10 ha abandoned arable field on Jurassic corallian limestone. Two climate change scenarios were used: warmer winters with increased summer rainfall and warmer winters with summer drought. Plant cover and species richness were significantly increased in plots receiving supplemented summer rainfall, while the amount of litter was significantly reduced. Litter formation was significantly increased by winter warming and drought. The responses of the plant community to the climate manipulations were related to the life-history attributes of the dominant species. Seedling recruitment was limited by microsite availability, which also varied in the different climate manipulations. The results are discussed in terms of successional dynamics. They suggest that warmer winters may delay succession, as gap formation in the sward will provide sites for colonisation of annuals, thereby enabling their persistence in the sward. Under wetter conditions during summer, perennial grasses tend to close the sward, thereby inhibiting the establishment of later successional species.     

Assessing the vulnerability of endemic diatom species in Lake Baikal to predicted future climate change: a multivariate approach

Diatoms in Lake Baikal exhibit significant spatial variation, related to prevailing climate, lake morphology and fluvial input into the lake. Here we have assessed the threats to endemic planktonic diatom species (through the development of empirical models), which form a major component of primary production within the lake. Multivariate techniques employed include redundancy analysis (RDA) and Huisman–Olff–Fresco (HOF) models. Our analyses suggest that eight environmental variables were significant in explaining diatom distribution across the lake, and in order of importance these are snow thickness on the ice, water depth, duration of days with white ice, suspended matter in the lake, days of total ice duration, temperature of the water surface in July, concentration of zooplankton and suspended organic matter. Impacts on dominant phytoplankton diatom species are highlighted using t‐value biplots. Predictions of future climate change on Lake Baikal are likely to result in shorter periods of ice cover, decreased snow cover across the lake in spring, increased fluvial input into the lake, and an increase in the intensification of surface water stratification during summer months. All these factors are likely to impact negatively on the slow‐growing, cold‐water endemics such as Aulacoseira baicalensis and Cyclotella minuta, which currently dominate diatom assemblages. Instead, taxa that are only intermittently abundant, at present, in offshore areas (e.g. Stephanodiscus meyerii) are likely to become more frequent. However, given the climatic gradient across the lake, the timing and extent of changes in community structure are likely to vary. Moreover, palaeolimnological records show that Lake Baikal diatom assemblages have been dynamic throughout the Holocene, with both endemic and cosmopolitan species exhibiting periods of dominance. Effects of climate change on the entire lake ecosystem may yet be profound as the structure of the pelagic food web may change from one based on endemic diatom taxa to one dominated by nondiatom picoplankton, and as limnological functioning (e.g. stratification and mixing) affects deepwater oxygen availability, nutrient cycling and trophic linkages.     

Long-term dynamics of phytoplankton assemblages: Microcystis-domination in Lake Taihu, a large shallow lake in China

Long-term phytoplankton assemblages in a large shallow Chineselake, Lake Taihu, were presented using the monthly monitoringdata from October 1991 to December 1999. Earlier research results(1960, 1981 and 1988) were applied to discuss the differenttrophic stages of the lake. The species composition in the lakewas more closely related to eutrophication level than to lake-size,shallowness, or turbidity. Each summer, a single peak of phytoplanktonbiovolume appeared in Meiliang Bay. The results of principalcomponents analysis showed a distinct temporal shift in speciescomposition between summer and winter. A clear spatial differencein phytoplankton occurred between Meiliang Bay and the lakecentre. Wind speed and direction affected the horizontal distributionof phytoplankton, especially Microcystis, in the lake. Temperature,underwater light climate, nutrients and grazing by zooplanktonand by fish were discussed to explain the overwhelming dominanceof Microcystis. Four nutrient-phytoplankton stages were identifiedin the lake: an oligo-mesotrophic stage with low algal biomassuntil 1981, a eutrophic situation with blooms of Microcystisduring 1988–1995, hypertrophic conditions with the dominanceof Planctonema and total phosphorus up to 200 mg m^-3 from 1996to 1997 and the restoration period after 1997. The wax and waneof the phytoplankton assemblages were mainly controlled by temperature,wind and turbidity while long-term biomass dynamics were influencedby the level of nutrients.     

Regional, vertical and seasonal distribution of phytoplankton and photosynthetic pigments in Lake Baikal

A 3-year phytoplankton study was carried out in Lake Baikal(Siberia) as part of the CONTINENT project and in conjunctionwith a 60-year long monitoring programme by the Irkutsk StateUniversity. A combination of microscopy and high performanceliquid chromatography (HPLC) pigment analysis was used. Allover the lake, the dominant functional group (by biovolume)was the vernal diatom blooms, due to the dominance of endemicCyclotella species. Chlorophyll a (Chl a) was significantlyhighest at the Selenga and Barguzin inflows (2.39 ± 0.34and 2.49 ± 0.18 nmol L^–1, mean ± 95% CI,respectively) and higher in the South than in the North (1.43± 0.26 and 0.96 ± 0.13 nmol L^–1). This variationof Chl a reflected changes in the phytoplankton composition.Diatoms and Chrysophyceae were the major contributors to thetotal Chl a except in the South (Chlorophyceae) and SelengaDelta (cyanobacterial picoplankton). There were also indicationsof species composition changes due to enhanced P-loading fromthe Selenga River. However, canonical analyses indicated thattemperature and stratification were the major driving forcesfor regional distribution patterns and seasonal succession.It seems likely that further global warming will cause a shiftin the species and group composition towards small cells atthe expense of the large endemic diatom flora.     

Diatom succession trends in recent sediments from Lake Baikal and their relation to atmospheric pollution and to climate change

Recent environmental change in Lake Baikal has been attributed to anthropogenic influences on the ecosystem, especially through pollution and cultural eutrophication. These hypotheses are tested in this paper principally by diatom analyses in 20 short sediment cores. Most of the cores were collected with a new type of box corer specifically designed for use on Lake Baikal. Most cores contain a good sediment record but turbidites occur in some sediment profiles which may be best recognized using a combination of techniques, such as radiometric dating and percentage dry weight analyses. The most recent sediments, especially those in the southern basin and in the very north of Baikal, contain a record of anthropogenic contamination in the form of lead and spheroidal carbonaceous particles, which confirms that the southern basin of Baikal is most affected by atmospheric sources of pollution. However, there is no sedimentary diatom evidence indicating offshore water quality deterioration in Baikal owing to air pollution or eutrophication. Small increases in diatoms which indicate nutrient enrichment (e.g. Stephanodiscus minutulus, Synedra acus v. radians and Synedra acus v. acus) may reflect local eutrophication of the shallow waters close to the Selenga Delta and certain coastal sites in the southern basin near to the Baikalsk paper and pulp mill. By using numerical techniques, Lake Baikal can be split into at least four regions on the basis of its surface sediment flora: the south, middle and north basins, and the shallow waters surrounding the Selenga Delta region. Diatom analyses reveal that the endemic flora of Lake Baikal has been constantly changing over at least the last 2000 years and that these fluctuations are probably responses to natural climatic variability. Recent sediments of Baikal may be affected by taphanomic processes (e.g. dissolution) and turbidite deposition, and these must be taken into account when interpreting the sedimentary diatom record. The diatom flora of the lake is currently dominated by several species, such as Aulacoseira baicalensis, A. islandica, Cyclotella minuta and Stephanodiscus binderanus v. baicalensis. All these species, except for C. minuta, have become more common in the lake in approximately the last 130 years, and we hypothesize that these changes may be attributed to a number of different processes linked to an ameliorating climate after the end of the Little Ice Age. The results presented here have important implications for this recently designated World Heritage Site, with regard to future pollution controls and catchment management policies.     

Long-term changes in phytoplankton, zooplankton and salmon related to climate

Recently, large‐scale changes in the biogeography of calanoid copepod crustaceans have been detected in the northeastern North Atlantic Ocean and adjacent seas. Strong biogeographical shifts in all copepod assemblages were found with a northward extension of more than ° in latitude of warm‐water species associated with a decrease in the number of colder‐water species. These changes were attributed to regional increase in sea surface temperature. Here, we have extended these studies to examine long‐term changes in phytoplankton, zooplankton and salmon in relation to hydro‐meteorological forcing in the northeast Atlantic Ocean and adjacent seas. We found highly significant relationships between (1) long‐term changes in all three trophic levels, (2) sea surface temperature in the northeastern Atlantic, (3) Northern Hemisphere temperature and (4) the North Atlantic Oscillation. The similarities detected between plankton, salmon, temperature and hydro‐climatic parameters are also seen in their cyclical variability and in a stepwise shift that started after a pronounced increase in Northern Hemisphere Temperature anomalies at the end of the 1970s. All biological variables show a pronounced change which started after circa 1982 for euphausiids (decline), 1984 for the total abundance of small copepods (increase), 1986 for phytoplankton biomass (increase) and Calanus finmarchicus (decrease) and 1988 for salmon (decrease). This cascade of biological events led to an exceptional period, which is identified after 1986 to present and followed another shift in large‐scale hydro‐climatic variables and sea surface temperature. This regional temperature increase therefore appears to be an important parameter that is at present governing the dynamic equilibrium of northeast Atlantic pelagic ecosystems with possible consequences for biogeochemical processes and fisheries.     

Lipids and fatty acids of two pelagic cottoid fishes (Comephorus spp) endemic to Lake Baikal

Matured females of two Lake Baikal endemic fish species, Comephorus baicalensis and Comephorus dybowski, have been investigated for lipid of the whole body and specific tissues (liver, muscles, ovaries), phospholipid classes and fatty acids of neutral and polar lipids. Total lipid in the body (38.9% fresh weight), liver (23.5%) and muscles (14.5%) of C. baicalensis were greater than those of C. dybowski (4.7, 8.7 and 2.6%, respectively); only their ovaries were similar (5.3 and 5.6% lipid, respectively). In both species, phosphatidylcholine and phosphatidylethanolamine were the major phospholipids, ranging from 60.7 to 75.1% of total phospholipid and 14.5–25.7%, respectively. In most cases, monounsaturated fatty acids (MUFA) were the major fatty acid group in C. baicalensis, whereas polyunsaturated fatty acids (PUFA) were the major group in C. dybowski. The MUFA 18:1(n-9) prevailed over other fatty acids in C. baicalensis and varied from 19% in polar lipids of muscles to 56.1% in neutral lipids of muscles. In polar lipid of C. dybowski, the PUFA 22:6(n-3) prevailed over other fatty acids in muscles and ovaries, while 16:0 dominated polar liver lipids and neutral lipids of all tissues. Other major fatty acids included 16:1(n-7), 18:1(n-7), and 20:5(n-3). Values of the (n-3)/(n-6) fatty acid ratio for neutral lipids of C. baicalensis (0.5–0.9) are well below the range of values characteristic either for marine or freshwater fish, while these values for polar lipids (1.6–1.8) are in the range typical of freshwater fish. Neutral lipid fatty acid ratios in C. dybowski (2.5–3.1) allow it to be assigned to freshwater fish, but polar lipids (2.8–3.7) leave it intermediary between freshwater and marine fish.     

Predicting the influence of wolf-provided carrion on scavenger community dynamics under climate change scenarios

Climate change poses an immediate threat to the persistence and distribution of many species, yet our ability to forecast changes in species composition is hindered by poor understanding of the extent to which higher trophic‐level interactions may buffer or exacerbate the adverse effects of warming. We incorporated species‐specific consumption data from 240 wolf‐killed elk carcasses from Yellowstone National Park into stochastic simulation models to link trends in the El Niño Southern Oscillation (ENSO) to food procurement by a guild of scavengers as a function of gray wolf reintroduction. We find that a shift in ENSO towards the El Niño (warming) phase of the cycle coincident with increasing global temperatures reduces carrion for scavengers, particularly those with strong seasonal patterns in resource use such as grizzly bears. Wolves alleviate these warming‐induced food shortages by rendering control over this crucial resource to biotic rather than abiotic factors. Ecosystems with intact top predators are likely to exhibit stronger biotic regulation and should be more resistant to climate change than ecosystems lacking them.     

Biotic interactions may overrule direct climate effects on spring phytoplankton dynamics

To improve our mechanistic understanding and predictive capacities with respect to climate change effects on the spring phytoplankton bloom in temperate marine systems, we used a process‐driven dynamical model to disentangle the impact of potentially relevant factors which are often correlated in the field. The model was based on comprehensive indoor mesocosm experiments run at four temperature and three light regimes. It was driven by time‐series of water temperature and irradiance, considered edible and less edible phytoplankton separately, and accounted for density‐dependent grazing losses. It successfully reproduced the observed dynamics of well edible phytoplankton in the different temperature and light treatments. Four major factors influenced spring phytoplankton dynamics: temperature, light (cloudiness), grazing, and the success of overwintering phyto‐ and zooplankton providing the starting biomasses for spring growth. Our study predicts that increasing cloudiness as anticipated for warmer winters for the Baltic Sea region will retard phytoplankton net growth and reduce peak heights. Light had a strong direct effect in contrast to temperature. However, edible phytoplankton was indirectly strongly temperature‐sensitive via grazing which was already important in early spring at moderately high algal biomasses and counter‐intuitively provoked lower and later algal peaks at higher temperatures. Initial phyto‐ and zooplankton composition and biomass also had a strong effect on spring algal dynamics indicating a memory effect via the broadly under‐sampled overwintering plankton community. Unexpectedly, increased initial phytoplankton biomass did not necessarily lead to earlier or higher spring blooms since the effect was counteracted by subsequently enhanced grazing. Increasing temperature will likely exhibit complex indirect effects via changes in overwintering phytoplankton and grazer biomasses and current grazing pressure. Additionally, effects on the phytoplankton composition due to the species‐specific susceptibility to grazing are expected. Hence, we need to consider not only direct but also indirect effects, e.g. biotic interactions, when addressing climate change impacts.