Climate change and the phytoplankton spring bloom: warming and overwintering zooplankton have similar effects on phytoplankton

Indoor mesocosms were used to study the combined effect of warming and of different densities of overwintering mesozooplankton (mainly copepods) on the spring development of phytoplankton in shallow, coastal waters. Similar to previous studies, warming accelerated the spring phytoplankton peak by ca. 1 day °C^?1 whereas zooplankton did not significantly influence timing. Phytoplankton biomass during the experimental period decreased with warming and with higher densities of overwintering zooplankton. Similarly, average cell size and average effective particle size (here: colony size) decreased both with zooplankton density and warming. A decrease in phytoplankton particle size is generally considered at typical footprint of copepod grazing. We conclude that warming induced changes in the magnitude and structure of the phytoplankton spring bloom cannot be understood without considering grazing by overwintering zooplankton.     

Climate change and the timing, magnitude, and composition of the phytoplankton spring bloom

In this article, we show by mesocosm experiments that winter and spring warming will lead to substantial changes in the spring bloom of phytoplankton. The timing of the spring bloom shows only little response to warming as such, while light appears to play a more important role in its initiation. The daily light dose needed for the start of the phytoplankton spring bloom in our experiments agrees well with a recently published critical light intensity found in a field survey of the North Atlantic (around 1.3 mol photons m^?2 day^?1). Experimental temperature elevation had a strong effect on phytoplankton peak biomass (decreasing with temperature), mean cell size (decreasing with temperature) and on the share of microplankton diatoms (decreasing with temperature). All these changes will lead to poorer feeding conditions for copepod zooplankton and, thus, to a less efficient energy transfer from primary to fish production under a warmer climate.     

Climatic effects on the breeding phenology and reproductive success of an arctic-nesting goose species

Climate warming is pronounced in the Arctic and migratory birds are expected to be among the most affected species. We examined the effects of local and regional climatic variations on the breeding phenology and reproductive success of greater snow geese ( Chen caerulescens atlantica ), a migratory species nesting in the Canadian Arctic. We used a long-term dataset based on the monitoring of 5447 nests and the measurements of 19 234 goslings over 16 years (1989–2004) on Bylot Island. About 50% of variation in the reproductive phenology of individuals was explained by spring climatic factors. High mean temperatures and, to a lesser extent, low snow cover in spring were associated with an increase in nest density and early egg-laying and hatching dates. High temperature in spring and high early summer rainfall were positively related to nesting success. These effects may result from a reduction in egg predation rate when the density of nesting geese is high and when increased water availability allows females to stay close to their nest during incubation recesses. Summer brood loss and production of young at the end of the summer increased when values of the summer Arctic Oscillation (AO) index were either very positive (low temperatures) or very negative (high temperatures), indicating that these components of the breeding success were most influenced by the regional summer climate. Gosling mass and size near fledging were reduced in years with high spring temperatures. This effect is likely due to a reduced availability of high quality food in years with early spring, either due to food depletion resulting from high brood density or a mismatch between hatching date of goslings and the timing of the peak of plant quality. Our analysis suggests that climate warming should advance the reproductive phenology of geese, but that high spring temperatures and extreme values of the summer AO index may decrease their reproductive success up to fledging.     

Predicted warming and browning affect timing and magnitude of plankton phenological events in lakes: a mesocosm study

1. Aquatic ecosystems in Northern Europe are expected to face increases in temperature and water colour (TB) in future. While effects of these factors have been studied separately, it is unknown whether and how a combination of them might affect phenological events and trophic interactions. 2. In a mesocosm study, we combined both factors to create conditions expected to arise during the coming century. We focused on quantifying effects on timing and magnitude of plankton spring phenological events and identifying possible mismatches between resources (phytoplankton) and consumers (zooplankton). 3. We found that the increases in TB had important effects on timing and abundance of different plankton groups. While increased temperature led to an earlier peak in phytoplankton and zooplankton and a change in the relative timing of different zooplankton groups, increased water colour reduced chlorophyll‐a concentrations. 4. Increased TB together benefitted cladocerans and calanoid copepods and led to stronger top‐down control of algae by zooplankton. There was no sign of a mismatch between primary producers and grazers as reported from other studies. 5. Our results point towards an earlier onset of plankton spring growth in shallow lakes in future with a stronger top‐down control of phytoplankton by zooplankton grazers.     

Geographical shift of zooplankton communities and decadal dynamics of the Kuroshio–Oyashio currents in the western North Pacific

We investigated temporal and spatial variations in the zooplankton community structure in the Oyashio and Transition region of the subarctic western North Pacific from 1960 to 1999 using principal component analysis (PCA) and zooplankton samples from the historical Odate Collection. In particular, we examined the influence of Kuroshio and Oyashio decadal dynamics on geographical variations in the zooplankton community. The first principal component (PC1) closely represented the interannual variation in cold-water, large copepod species, while the second PC (PC2) represented the variation in warm-water, small copepod species. Using Rodionov's regime-shift method, we detected a significant increase in the PC score after 1976 and 1981 for PC1 and PC2, respectively. After the shift years, (1) warm-water species increased in the Transition zone, (2) the distribution center of the cold-water species shifted southward, and (3) copepod abundance and species diversity increased in the Transition zone as a result of (1) and (2). The timing of these shifts in the zooplankton community roughly coincided with the North Pacific climatic regime shift in 1976/1977. From the mid-1970s to the early 1980s, the southern boundary of the Oyashio shifted southward and increased geostrophic transport was observed in the Kuroshio, indicating spin up of the Kuroshio–Oyashio system. Change in atmospheric circulation during the 1976/1977 regime shift is thought to have caused the spin up of these currents, which subsequently affected the regional zooplankton community through advective processes.     

Earlier onset of the spring phytoplankton bloom in lakes of the temperate zone in a warmer climate

The decoupling of trophic interactions is potentially one of the most severe consequences of climate warming. In lakes and oceans the timing of phytoplankton blooms affects competition within the plankton community as well as food–web interactions with zooplankton and fish. Using Upper Lake Constance as an example, we present a model‐based analysis that predicts that in a future warmer climate, the onset of the spring phytoplankton bloom will occur earlier in the year than it does at present. This is a result of the earlier occurrence of the transition from strong to weak vertical mixing in spring, and of the associated earlier onset of stratification. According to our simulations a shift in the timing of phytoplankton growth resulting from a consistently warmer climate will exceed that resulting from a single unusually warm year. The numerical simulations are complemented by a statistical analysis of long‐term data from Upper Lake Constance which demonstrates that oligotrophication has a negligible effect on the timing of phytoplankton growth in spring and that an early onset of the spring phytoplankton bloom is associated with high air temperatures and low wind speeds.     

Distribution-wide effects of climate on population densities of a declining migratory landbird

1. Increases in global temperatures have created concern about effects of climatic variability on populations, and climate has been shown to affect population dynamics in an increasing number of species. Testing for effects of climate on population densities across a species' distribution allows for elucidation of effects of climate that would not be apparent at smaller spatial scales. 2. Using autoregressive population models, we tested for effects of the North Atlantic Oscillation (NAO) and the El Ni?o Southern Oscillation (ENSO) on annual population densities of a North American migratory landbird, the yellow-billed cuckoo Coccyzus americanus, across the species' breeding distribution over a 37-year period (1966-2002). 3. Our results indicate that both the NAO and ENSO have affected population densities of C. americanus across much of the species' breeding range, with the strongest effects of climate in regions in which these climate systems have the strongest effects on local temperatures. Analyses also indicate that the strength of the effect of local temperatures on C. americanus populations was predictive of long-term population decline, with populations that were more negatively affected by warm temperatures experiencing steeper declines. 4. Results of this study highlight the importance of distribution-wide analyses of climatic effects and demonstrate that increases in global temperatures have the potential to lead to additional population declines.     

Intercomparison, interpretation, and assessment of spring phenology in North America estimated from remote sensing for 1982–2006

Shifts in the timing of spring phenology are a central feature of global change research. Long‐term observations of plant phenology have been used to track vegetation responses to climate variability but are often limited to particular species and locations and may not represent synoptic patterns. Satellite remote sensing is instead used for continental to global monitoring. Although numerous methods exist to extract phenological timing, in particular start‐of‐spring (SOS), from time series of reflectance data, a comprehensive intercomparison and interpretation of SOS methods has not been conducted. Here, we assess 10 SOS methods for North America between 1982 and 2006. The techniques include consistent inputs from the 8 km Global Inventory Modeling and Mapping Studies Advanced Very High Resolution Radiometer NDVIg dataset, independent data for snow cover, soil thaw, lake ice dynamics, spring streamflow timing, over 16 000 individual measurements of ground‐based phenology, and two temperature‐driven models of spring phenology. Compared with an ensemble of the 10 SOS methods, we found that individual methods differed in average day‐of‐year estimates by ±60 days and in standard deviation by ±20 days. The ability of the satellite methods to retrieve SOS estimates was highest in northern latitudes and lowest in arid, tropical, and Mediterranean ecoregions. The ordinal rank of SOS methods varied geographically, as did the relationships between SOS estimates and the cryospheric/hydrologic metrics. Compared with ground observations, SOS estimates were more related to the first leaf and first flowers expanding phenological stages. We found no evidence for time trends in spring arrival from ground‐ or model‐based data; using an ensemble estimate from two methods that were more closely related to ground observations than other methods, SOS trends could be detected for only 12% of North America and were divided between trends towards both earlier and later spring.     

Trends and temperature response in the phenology of crops in Germany

The phenology of 78 agricultural and horticultural events from a national survey in Germany spanning the years 1951–2004 is examined. The majority of events are significantly earlier now than 53 years ago, with a mean advance of 1.1–1.3 days per decade. The mean trends for 'true phases', such as emergence and flowering, of annual and perennial crops are not significantly different, although more trends (78% vs. 46%) are significant for annual crops. We attempt to remove the influence of technological advance or altered farming practices on phenology by detrending the respective time series by linear regression of date (day number) on year. Subsequently, we estimate responses to mean monthly and seasonal temperature by correlation and regression in two ways; with and without removing the year trend first. Nearly all (97%) correlation coefficients are negative, suggesting earlier events in warmer years. Between 82% and 94% of the coefficients with seasonal spring and summer temperatures are significant. The conservative estimate (detrended) of mean temperature response against mean March–May temperature (−3.73 days °C⁻¹) is significantly less than the full estimate (−4.31 days °C⁻¹), the 'true' size of phenological temperature response may lie in between. Perennial crops exhibited a significantly higher temperature response to mean spring temperature than the annual crops.     

The effects of Daphnia on nutrient stoichiometry and filamentous cyanobacteria: a mesocosm experiment in a eutrophic lake

1. Stoichiometric theory predicts that the nitrogen : phosphorus (N : P) ratio of recycled nutrients should increase when P‐rich zooplankton such as Daphnia become dominant. We used an enclosure study to test the hypothesis that an increased biomass of Daphnia will increase the relative availability of N versus P sufficiently to decrease the abundance of filamentous cyanobacteria. The experiment was conducted in artificially enriched Lake 227 (L227) in the Experimental Lakes Area (ELA), north‐western Ontario, Canada. Previous studies in L227 have shown that the dominance of filamentous, N‐fixing cyanobacteria is strongly affected by changes in the relative loading rates of N and P. 2. We used a 2 × 2 factorial design with the addition or absence of D. pulicaria and high or low relative loading rates of N and P (+NH₄, –NH₄) in small enclosures as treatment variables. If Daphnia can strongly affect filamentous cyanobacteria by altering N and P availability, these impacts should be greatest with low external N : P loading rates. The phytoplankton community of L227 was predominantly composed of filamentous Aphanizomenon spp. at the start of the experiment. 3. Daphnia strongly reduced filamentous cyanobacterial density in all enclosures to which they were added. The addition of NH₄ had only a small impact on algal community composition. Hence, we conclude that Daphnia did not cause reductions in cyanobacteria by altering the N : P ratio of available nutrients. 4. Despite the lack of evidence that Daphnia affected filamentous cyanobacteria by altering the relative availability of N and P, we found changes in nutrient cycling consistent with other aspects of stoichiometric theory. In the presence of Daphnia, total P in the water column decreased because of an increase in P sedimentation. In contrast to P, a decrease in suspended particulate N was offset by an increase in dissolved N (especially NH₄). Hence, dissolved and total N : P ratios in the water column increased with Daphnia as a result of differences in the fate of suspended particulate N versus P. There was minimal accumulation and storage of P in Daphnia biomass in the enclosures. 5. Our experiment demonstrated that Daphnia can strongly limit filamentous cyanobacterial abundance and affect the biogeochemical cycling of nutrients. In our study, changes in nutrient cycling were apparently insufficient to cause the changes in phytoplankton community composition that we observed. Daphnia therefore limited filamentous cyanobacteria by other mechanisms.     

Effects of climatic change on the phenology of butterflies in the northwest Mediterranean Basin

Phenological changes in response to climatic warming have been detected across a wide range of organisms. Butterflies stand out as one of the most popular groups of indicators of climatic change, given that, firstly, they are poikilothermic and, secondly, have been the subject of thorough monitoring programmes in several countries for a number of decades. Here we provide for the first time strong evidence of phenological change as a consequence of recent climatic warming in butterflies at a Spanish site in the northwest Mediterranean Basin. By means of the widely used Butterfly Monitoring Scheme methodology, three different phenological parameters were analysed for the most common species to test for trends over time and relationships with temperature and precipitation. Between 1988 and 2002, there was a tendency for earlier first appearance dates in all 17 butterfly species tested, and significant advances in mean flight dates in 8 out of 19 species. On the other hand, the shape of the curve of adult emergence did not show any regular pattern. These changes paralleled an increase of 1–1.5°C in mean February, March and June temperatures. Likewise, a correlation analysis indicated the strong negative effect of spring temperature on phenological parameters (i.e. higher temperatures tended to produce phenological advances), and the opposite effect of precipitation in certain months. In addition, there was some evidence to indicate that phenological responses may differ between taxonomic lineages or species with similar diets. We discuss the consequences that these changes may have on species' population abundances, especially given the expected increase in aridity in the Mediterranean Basin caused by current climatic warming. We predict that varying degrees of phenological flexibility may account for differences in species' responses and, for multivoltine species, predict strong selection favouring local seasonal adaptations such as diapause phenomena or migratory behaviour.     

Effects of decadal climate change on zooplankton over the last 50 years in the western subarctic North Pacific

Decadal‐ to multi‐decadal variations have been reported in many regional ecosystems in the North Pacific, resulting in an increasing demand to elucidate the link between long‐term climatic forcing and marine ecosystems. We detected phenological and quantitative changes in the copepod community in response to the decadal climatic variation in the western subarctic North Pacific by analyzing the extensive zooplankton collection taken since the 1950s, the Odate Collection. Copepod species were classified into five seasonal groups depending on the timing of the annual peak in abundance. The abundance of the spring community gradually increased for the period 1960–2002. The spring–summer community also showed an increasing trend in May, but a decadal oscillation pattern of quasi‐30‐year cycles in July. Phenological changes coincided with the climate regime shift in the mid‐1970s, indicated by the Pacific decadal oscillation index (PDO). After the regime shift, the timing of the peak abundance was delayed one month, from March–April to April–May, in the spring community, whereas it peaked earlier, from June–July to May–June, in the spring–summer community, resulting in an overlap of the high productivity period for the two communities in May. Wintertime cooling, followed by rapid summertime warming, was considered to be responsible for delayed initiation and early termination of the productive season after the mid‐1970s. Another phenological shift, quite different from the previous decade, was observed in the mid‐1990s, when warm winters followed by cool summers lengthened the productive season. The results suggest that climatic forcing with different decadal cycles may operate independently during winter–spring and spring–summer to create seasonal and interannual variations in hydrographic conditions; thus, combinations of these seasonal processes may determine the annual biological productivity.     

Toward a synthetic understanding of the role of phenology in ecology and evolution

Phenology affects nearly all aspects of ecology and evolution. Virtually all biological phenomena—from individual physiology to interspecific relationships to global nutrient fluxes—have annual cycles and are influenced by the timing of abiotic events. Recent years have seen a surge of interest in this topic, as an increasing number of studies document phenological responses to climate change. Much recent research has addressed the genetic controls on phenology, modelling techniques and ecosystem-level and evolutionary consequences of phenological change. To date, however, these efforts have tended to proceed independently. Here, we bring together some of these disparate lines of inquiry to clarify vocabulary, facilitate comparisons among habitat types and promote the integration of ideas and methodologies across different disciplines and scales. We discuss the relationship between phenology and life history, the distinction between organismal- and population-level perspectives on phenology and the influence of phenology on evolutionary processes, communities and ecosystems. Future work should focus on linking ecological and physiological aspects of phenology, understanding the demographic effects of phenological change and explicitly accounting for seasonality and phenology in forecasts of ecological and evolutionary responses to climate change.     

A bioassay study of effects of zooplankton,iron and NTA on the phytoplankton productivity of a marl lake

Additions of iron and NTA had minor stimulatory effects on the phytoplankton productivity of samples of water from two calcareous Michigan lakes in autumn and winter contrary to effects at other seasons. Added Daphnia pulex significantly reduced phytoplankton productivity by grazing at this time of year, in all tests at all levels of addition from 4–32 animals/l, greatly in excess of any possible beneficial effects of increased nutrient availability.     

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.     

Combined effects of climate and biotic interactions on the elevational range of a phytophagous insect

1. The ranges of many species have expanded in cool regions but contracted at warm margins in response to recent climate warming, but the mechanisms behind such changes remain unclear. Particular debate concerns the roles of direct climatic limitation vs. the effects of interacting species in explaining the location of low latitude or low elevation range margins. 2. The mountains of the Sierra de Guadarrama (central Spain) include both cool and warm range margins for the black-veined white butterfly, Aporia crataegi, which has disappeared from low elevations since the 1970s without colonizing the highest elevations. 3. We found that the current upper elevation limit to A. crataegi's distribution coincided closely with that of its host plants, but that the species was absent from elevations below 900 m, even where host plants were present. The density of A. crataegi per host plant increased with elevation, but overall abundance of the species declined at high elevations where host plants were rare. 4. The flight period of A. crataegi was later at higher elevations, meaning that butterflies in higher populations flew at hotter times of year; nevertheless, daytime temperatures for the month of peak flight decreased by 6.2 degrees C per 1 km increase in elevation. 5. At higher elevations A. crataegi eggs were laid on the south side of host plants (expected to correspond to hotter microclimates), whereas at lower sites the (cooler) north side of plants was selected. Field transplant experiments showed that egg survival increased with elevation. 6. Climatic limitation is the most likely explanation for the low elevation range margin of A. crataegi, whereas the absence of host plants from high elevations sets the upper limit. This contrasts with the frequent assumption that biotic interactions typically determine warm range margins, and thermal limitation cool margins. 7. Studies that have modelled distribution changes in response to climate change may have underestimated declines for many specialist species, because range contractions will be exacerbated by mismatch between the future distribution of suitable climate space and the availability of resources such as host plants.     

Influence of climate change on the abundance, distribution and phenology of woodland bird species in temperate regions

There is now overwhelming evidence that an increase in the concentration of greenhouse gases in the Earth's atmosphere has caused global temperatures to increase by 0.6 °C since 1900 and further increases of between 1.4 and 5.8 °C are predicted over the next century. Changes in climatic conditions have already influenced the demography, phenology and distribution of a wide range of plant and animal taxa. This review focuses on the impacts, both observed and potential, of climate change on birds breeding in temperate woodlands of the Western Palaearctic, a significant proportion of which are currently declining. Changes in ambient temperatures and patterns of precipitation may have direct and indirect effects on the survival rates and productivity of bird species, thus influencing population sizes. For some species or populations, the timing of events such as egg-laying and return from the wintering grounds is also changing in relation to shifts in the peak of food availability during the breeding season. The degree to which different individuals are able to track these temporal changes will have a significant bearing on population sizes and distributions in the future. Unless active management steps are taken, the relatively low dispersal rates of tree species may lead to a decrease in the total area of some woodland habitat types as losses at the southern edge of the range are likely to occur much more quickly than expansion at the northern edge. In addition, the dispersal rates of many woodland birds are themselves low, which could affect their ability to move to new habitat patches if currently occupied areas become unsuitable. Thus, woodland birds may be particularly susceptible to the impacts of climate change.     

Modelling the effects on phytoplankton communities of changing mixed depth and background extinction coefficient on three contrasting lakes in the English Lake District

1. The process‐based phytoplankton community model, PROTECH, was used to model the response of algal biomass to a range of mixed layer depths and extinction coefficients for three contrasting lakes: Blelham Tarn (eutrophic), Bassenthwaite Lake (mesotrophic) and Ullswater (oligotrophic). 2. As expected, in most cases biomass and diversity decreased with decreasing light availability caused by increasing the mixed depth and background extinction coefficient. The communities were generally dominated by phytoplankton tolerant of low light. Further, more novel, factors were identified, however. 3. In Blelham Tarn in the second half of the year, biomass and diversity did not generally decline with deeper mixing and the community was dominated by nitrogen‐fixing phytoplankton because that nutrient was limiting to growth. 4. In Bassenthwaite Lake, changing mixed depth influenced the retention time so that, as the mixed depth declined, the flushing rate in the mixed layer increased to the point that only fast‐growing phytoplankton could dominate. 5. In the oligotrophic Ullswater, changing the mixed depth had a greater effect through nutrient supply rather than light availability. This effect was observed when the mixed layer was relatively shallow (<5.5 m) and the driver for this was that the inflowing nutrients were added to a smaller volume of water, thus increasing nutrient concentrations and algal growth. 6. Therefore, whilst changes in mixed depth generally affect the phytoplankton via commonly recognized factors (light availability, sedimentation rate), it also affected phytoplankton growth and community composition through other important factors such as retention time and nutrient supply.     

Hierarchical regulation of pelagic invertebrates in lakes of the northern Great Plains: a novel model for interdecadal effects of future climate change on lakes

Endorheic lakes of the northern Great Plains encompass a wide range of environmental parameters (e.g., salinity, pH, DOC, Ca, nutrients, depth) that vary 1000‐fold among sites and through the past 2000 years due to variation in basin hydrology and evaporative forcing. However, while many environmental parameters are known to individually influence zooplankton diversity and taxonomic composition, relatively little is known of the hierarchical relationships among potential controls or of how regulatory mechanisms may change in response to climate variation on diverse scales. To address these issues, we surveyed 70 lakes within a 100 000 km² prairie region to simulate the magnitude of environmental change expected to occur over 100–1000 years and to quantify the unique and interactive effects of diverse environmental parameters in regulating pelagic invertebrate community structure at that scale. Multivariate analyses showed that salinity was the principal correlate of changes in invertebrate composition among lakes, with a sequential loss of taxa between salinities of 4 and 50 g total dissolved solids L^?1 until one to two species predominated in highly saline systems. In contrast, changes in the concentrations of Ca²⁺ and other mineral nutrients exerted secondary controls of invertebrate assemblages independent of salinity, whereas lake depth provided a tertiary regulatory mechanism structuring species composition. In contrast to these large‐scale hierarchical patterns, seasonal surveys (May, July, September) of a subset of 21 lakes in each of 2003–2005 revealed that annual meteorological variation had no measurable effect on pelagic invertebrates, despite large differences in temperature, precipitation, and evaporation arising from regional droughts. Together these findings show that pelagic invertebrate communities in saline lakes are resilient to interannual variability in climate, but suggest that lakes of the northern Great Plains may provide a sensitive model to forecast centennial effects of future climate change.