Climate Change And Freshwater Zooplankton: What Does It .
Aquat EcolDOI 10.1007/s10452-012-9418-8Climate change and freshwater zooplankton: what does itboil down to?Csaba Vadadi-Fülöp Csaba Sipkay Gergely Mészáros Levente HufnagelReceived: 4 June 2012 / Accepted: 1 October 2012Ó Springer Science Business Media Dordrecht 2012Abstract Recently, major advances in the climate–zooplankton interface have been made some of whichappeared to receive much attention in a broaderaudience of ecologists as well. In contrast to themarine realm, however, we still lack a more holisticsummary of recent knowledge in freshwater. Wediscuss climate change-related variation in physicaland biological attributes of lakes and running waters,high-order ecological functions, and subsequent alteration in zooplankton abundance, phenology, distribution, body size, community structure, life historyparameters, and behavior by focusing on communitylevel responses. The adequacy of large-scale climaticindices in ecology has received considerable supportHandling Editor: Piet Spaak.C. Vadadi-Fülöp (&)Hungarian Scientific Research Fund Office,Czuczor u. 10, 1093 Budapest, Hungarye-mail: vadfulcsab@gmail.comC. SipkayDanube Research Institute of the Hungarian Academyof Sciences, Jávorka Sándor u. 14, 2131 Göd, HungaryG. MészárosSzent István University, Páter Károly u. 1, 2100 Gödöll}o,HungaryL. HufnagelDepartment of Mathematics and Informatics,Corvinus University of Budapest, Villányi út 29-43,1118 Budapest, Hungaryand provided a framework for the interpretation ofcommunity and species level responses in freshwaterzooplankton. Modeling perspectives deserve particular consideration, since this promising stream ofecology is of particular applicability in climate changeresearch owing to the inherently predictive nature ofthis field. In the future, ecologists should expand theirresearch on species beyond daphnids, should addressquestions as to how different intrinsic and extrinsicdrivers interact, should move beyond correlativeapproaches toward more mechanistic explanations,and last but not least, should facilitate transfer ofbiological data both across space and time.Keywords Global warming Daphnia Phenology Community dynamics Ecological modelsIntroductionEvidence accumulated over recent decades providedinsights as to how climate change, as a significant partof global change, has already altered and is projectedto further alter phenology, distribution, abundance,and invasion potential of species as well as biodiversity at a global or nearly global scale (Dukes andMooney 1999; Sala et al. 2000; Walther et al. 2002;Parmesan and Yohe 2003; Root et al. 2003; Thomaset al. 2004). Lakes and rivers (Williamson et al. 2008;Adrian et al. 2009; Schindler 2009) as well as ongoingplankton monitoring programmes (Hays et al. 2005)123
Aquat Ecolhave been recognized as sentinels of global change.Why plankton are particularly good indicators ofclimate change is discussed in Hays et al. (2005) andRichardson (2008) in exhaustive details. Cladoceranand ostracod remains preserved in the sediment recordalso have been used to reconstruct past climaticchanges in lakes (Chivas et al. 1985; Lotter et al. 1997;Battarbee 2000). Zooplankton are key components ofaquatic food webs. Analogous with Drosophila ingenetics, Daphnia is known as a model organism inaquatic ecology (Lampert 2006, 2011). Therefore, abetter understanding of the role of climate change inaltering zooplankton dynamics, structure, and functionis of high scientific and economic value. However, westill lack a more holistic summary of recent knowledgeof climate change from a freshwater zooplankterperspective.Temperature is one of the most important factorsaccounting for variation in nearly all biological ratesand times, including metabolic rate (Gillooly et al.2001), population growth (Savage et al. 2004), lifespan (Gillooly et al. 2001), and developmental time(Gillooly et al. 2002). Metabolic theory (Brown et al.2004) suggests that metabolic rate, as a function ofbody size and temperature, controls ecological processes at all levels of organization from life historycharacteristics to species interactions and ecosystemprocesses. Also, virulence varies with temperaturesaltering the outcome of host–parasite interactions(Mitchell et al. 2005). Temperature rise, however, isnot the only phenomenon accompanying climatechange, variation in precipitation, wind and subsequent floods, droughts, altered mixing regimes allrepresent major forcing on the abiotic and biotictemplate.This paper aimed at summarizing the observed andprojected future effects of climate change on freshwater zooplankton from the ecologist’s perspective.Thus, we focus on the community level responses, andphysiological attributes are discussed in more limiteddetails. We dedicated an extra section for modelingstudies, as this promising stream of ecology is ofparticular applicability in climate change researchowing to the inherently predictive nature of this field.Not surprisingly, the literature is skewed toward lakeplankton, but wherever possible, we draw on examplesfrom running waters as well.Figure 1 summarizes the documented and projected direct and indirect effects of climate change on123freshwater zooplankton within an environmental andtrophic framework. First, we have a look at thepossible effects of large-scale climatic fluctuations onzooplankton community dynamics. Then, we turn tothe trophic level responses and begin to considervariation in community attributes one after the other.We devote an extra section to ultraviolet radiation(UVR) and subsequent variation in zooplanktonphysiology and community dynamics and demonstratewhy and how this has been linked to climate change.We close with a brief review of some sophisticatedmodeling approaches favored by ecologists in order togain more information than empirical and experimental studies can provide.Large-scale climatic fluctuationsHere, we briefly discuss some well-known large-scaleclimate patterns focusing on the North Atlantic Oscillation (NAO) and then begin to consider the effects ofthose on freshwater zooplankton communities.The NAO is a large-scale climatic oscillation ofatmospheric mass between the subtropical Azores highand the subpolar Iceland low, ranging from centralNorth America to Europe and much into Northern Asia.In a large part of the northern hemisphere, climatevariability is assumed to be influenced by the NAO,particularly in the winter term (Hurrell 1995). Themeasure of this oscillation, the winter NAO index, is thedifference of normalized sea level pressures betweenLisbon, Portugal, and Iceland, Stykkisholmur (Hurrell1995). The NAO index exhibited a positive trend towardlarger values, that is, a positive phase over the 70s, 80s,and 90s (Fig. 2), bringing to milder and wetter winters towestern and northern Europe, whereas colder and drierconditions to the Mediterranean region, northern Canada, and Greenland (Hurrell 1995). A similar phenomenon is the El Nino–Southern Oscillation (ENSO)affecting climate variability worldwide, but particularlyin the tropical Pacific (Allan et al. 1996) and the PacificDecadal Oscillation (PDO) accounting for major variation in the North Pacific climate (Mantua et al. 1997).North Atlantic Oscillation has been recognized as adriving force behind variation in phenology, abundance, distribution, and interspecific relationshipsacting through direct, indirect, and integrated effectsin variable groups and environments (Ottersen et al.2001; Stenseth et al. 2002). Although the NAO
Aquat EcolFig. 1 Conceptual modelof the possible direct andindirect effects of climatechange on freshwaterzooplankton. Solid arrowsindicate linkage betweencomponents while dashedarrows indicate feedbacks.Dotted lines indicatesynchronizing effects ofNAO, ENSO, and PDO (formore details, see the text).Plus and minus signsindicate potential positiveand negative effects onzooplankton. Note, that thismodel does not represent thefood web and theenvironment in detail, rathersketches relevant tracks ofclimate change-relatedimpacts from a zooplankterperspectiveFig. 2 a The winter (December–March) NAO index throughout the twentieth century and beyond. Data were obtained fromthe Climatic Research Unit, University of East Anglia (sourcehttp://www.cru.uea.ac.uk/cru/data/nao/). b The positive phaseof the NAO with the trendlineaccounts for a major variation of winter climate, it hasa memory effect in the community, lasting to almosthalf a year (Straile 2000; Gerten and Adrian 2000;Blenckner et al. 2007). Large-scale climate indices canhave even higher explanatory power than localweather variables (Hallett et al. 2004), because theyintegrate several weather components both in spaceand time and thus can be considered as appropriate‘‘weather packages’’ (Stenseth and Mysterud 2005).Contrasting results obtained from local weather variables and large-scale climate indices may improve ourunderstanding of what features of weather mayaccount for the pattern observed.The NAO has been linked to zooplankton successional events (earlier CWP: clear-water phase in highNAO years) (Straile 2000; Gerten and Adrian 2000;Wagner and Benndorf 2007) and abundances (Straileand Geller 1998; George 2000; Blenckner et al. 2007;Straile and Müller 2010).There is now ample evidence of large-scale coherent response of zooplankton to NAO (Straile 2002;Blenckner et al. 2007). ENSO also has been found tosynchronize the dynamics of several zooplankton taxain north-temperate lakes (Rusak et al. 2008). PDO andENSO have been linked to Daphnia abundanceincrease in Castle Lake (Park et al. 2004). In responseto the positive phase of the PDO, spring breakup dates123
Aquat Ecolhave shifted earlier in Lake Aleknagik over recentdecades and, as a result, daphnids increased inabundance in summer (Schindler et al. 2005). Byextending the duration of the stratification period,ENSO and PDO influenced phytoplankton and zooplankton phenologies in Lake Washington (Winderand Schindler 2004a).The observed behavior of the NAO index, that is, atrend toward a more positive phase, has resulted in aweakening winter stress allowing earlier populationgrowth, reduced mortality (Straile and Stenseth 2007),and overwintering populations (Adrian and Deneke1996). However, fall warming can have severeconsequences for zooplankton overwintering successthrough early emergence of resting stages, the result ofwhich can be the loss of the cohort, and throughswitching from sexual to asexual reproduction resulting in delayed production and reduced number ofephippial resting eggs, respectively (Chen and Folt1996).Climate variability, such as El Nino-relateddroughts, may alter zooplankton population dynamicsby triggering the emergence of resting stages residingin lake sediments (Arnott and Yan 2002). This may beof significant risk by depleting the egg bank and thusreducing the number of potential colonists whenenvironmental conditions improve (Arnott and Yan2002). On the other hand, diapausing strategies canhelp certain species to withstand unfavorable conditions under future environmental change (Hairston1996).Food web-related effects: a zooplankterperspectiveBottom–up effectsThere is now convincing empirical evidence forincreasing dominance of cyanobacteria in response toclimate change (Robarts and Zohary 1987; Adrian andDeneke 1996; Mooij et al. 2005; Shatwell et al. 2008;Paul 2008; Paerl and Huisman 2009; Wagner andAdrian 2009). It has received strong experimental (DeSenerpont Domis et al. 2007a; Jöhnk et al. 2008) andtheoretical (Elliott et al. 2005; Elliott 2010) supports aswell; however, some studies have not found betterperformance of cyanobacteria under rising temperatures(e.g., Moss et al. 2003). Cyanobacteria in general have123an arsenal of properties which enable these algae tothrive under climate change and outcompete otherphytoplankton including (1) buoyancy regulation, (2)N-fixing abilities, (3) UVR tolerance, and (4) betterperformance under elevated temperatures (Spencer andKing 1987; Robarts and Zohary 1987; Paul 2008).Increased cyanobacteria dominance does not favorherbivorous zooplankton for some reasons: (1) cyanobacteria are poor-quality food for zooplankton (Haney1987; Brett and Müller-Navarra 1997; Wilson et al.2006), (2) their filaments cause mechanical interferencewith the feeding apparatus of filter feeders (‘‘clogging’’)(Webster and Peters 1978; Porter and McDonough1984), (3) they can be toxic for consumers (Hietala et al.1997; Hansson et al. 2007a), but the direct harmfuleffects of cyanobacterial toxins to zooplankton haverecently been questioned (Wilson et al. 2006). It hasbeen shown that the mechanical interference of filaments with the filtering combs of daphnids decreaseswith increasing viscosity and declining temperature(Abrusán 2004). Note, however, that small zooplankters(cyclopoid copepods, small-bodied cladocerans, androtifers) can escape from competition with large-bodiedherbivores, for example, Daphnia, and can even benefitfrom cyanobacteria dominance (Fulton and Paerl 1988;Hansson et al. 2007a; Dupuis and Hann 2009a). Also,calanoid copepods have found to be better suited tocyanobacterial blooms than do cladocerans (Haney1987).Thermal stratification and related vertical mixingstrongly determine nutrient and light availability forphytoplankton (Diehl et al. 2002). In deep lakes, algalgrowth in spring is initiated by the stratification of thewater column allowing algal cells to be exposed tohigher light levels (Reynolds 1984; Gaedke et al.1998). Many freshwater systems have experiencedincreased stratification (reduced mixing) and a longerduration of stratification period (King et al. 1997;Gaedke et al. 1998; Livingstone 2003; Winder andSchindler 2004a; Coats et al. 2006; Peeters et al.2007a). As sinking velocities are a function of cellsize, reduced vertical mixing favors small-sizedspecies, species with buoyancy regulation (e.g., cyanobacteria) and motile flagellates over diatoms andgreen algae with higher sinking velocities (Streckeret al. 2004; Huisman et al. 2004; Winder and Hunter2008) and thus may alter community structure ofphytoplankton. All those are of crucial importance forconsumers, as diatoms are known as high-quality food
Aquat Ecolfor zooplankton because of their high content ofHUFA (highly unsaturated fatty acids) (Brett andMüller-Navarra 1997), whereas cyanobacteria mayhave several negative effects on grazers as discussedearlier in this section. Increased stratification andreduced mixing also can cut nutrient flux from deeperlayers. As a consequence, nutrient conditions in theepilimnion can be limiting for algae (George et al.1990) and decreased phytoplankton production canlead to the starvation of zooplankton; for instance, inEsthwaite Water, strongly reduced Daphnia abundance in ‘‘calm years’’ was assumed to be the result offood limitation because shallow mixing hamperedalgal growth through nutrient depletion (George2000). Daphnids, however, may benefit directly fromstratification through a positive response to increasedwater temperatures and/or indirectly via increasedphytoplankton productivity (Straile and Geller 1998).River flows are projected to decrease or increaseworldwide depending on the climate of the catchments(Arora and Boer 2001; Kundzewicz et al. 2007;Johnson et al. 2009) and thus may shape bottom–upeffects on zooplankton. Depending on the future flowregime, phytoplankton abundance may considerablyincrease or decrease in running waters (Phlips et al.2007), often with an increasing contribution ofcyanobacteria under low-flow conditions (Joneset al. 2011).2001; Gyllström et al. 2005; Schindler et al. 2005;Jeppesen et al. 2009, 2010). Furthermore, fish maybecome smaller and more omnivorous with increasingtemperatures implying higher predation on zooplankton (Jeppesen et al. 2010). An important considerationhere is that high densities of planktivorous fish andrelated strong predation pressure, as indirect effects,have the potential to mask the effect of climate changeon zooplankton (Schindler et al. 2005). As a result ofclimate change and resulted changes in lake physicalproperties, an increase in thermal habitat for cold-,cool-, and warm-water fish was recognized for largelakes and partly for small lakes as well (De Stasio et al.1996; Fang et al. 1999). Global warming may changethe habitat overlap between planktivorous fish andtheir prey both spatially and temporally, but theoutcome of this predator–prey interaction may varywith species and localities (De Stasio et al. 1996;Helland et al. 2007). High predation pressure by fishmay constrain zooplankton vertical migration underclimate warming (De Stasio et al. 1996). In shallowlakes, fish kills are expected to increase due todecreasing water levels (McGowan et al. 2005) andshortening of the ice cover period will, in turn, favorfish (Fang et al. 1999; Jackson et al. 2007).Top–down effectsPhenology is the study of seasonally recurring eventsin nature and it has been widely used as indicator ofclimate change (Walther et al. 2002; Root et al. 2003;Visser and Both 2005). Phenological response toglobal warming is now fairly acknowledged anddocumented in a variety of plant and animal speciesworldwide (see Parmesan and Yohe 2003; Root et al.2003; Parmesan 2007; Thackeray et al. 2010). Whenthe time lag between population peaks of prey andpredator increases, a mismatch between food availability and food requirement may arise (Cushing1969). Climate change is increasingly recognized asthe driver of such mismatches between predator andprey by triggering phenological events in an asynchronous way (Durant et al. 2007).This decoupling is a result of different phenologicalresponses of interacting species. Such decoupling wasreported from Lake Windermere (George and Harris1985) and Lake Washington (Winder and Schindler2004b). In the latter case, an advancement in the onsetClimate warming-related change in habitat structure caninduce strong cascading effects in pelagic food webs. Arecent example of how this works comes from the studyof Manca and DeMott (2009). Climate warming linkedchange in thermal refuge from fish predators allowed thepredatory cladoceran Bythotrephes longimanus (Leydig1860) to strongly increase in abundance and suppressthe population of Daphnia hyalina galeata. Warmingcan increase the risk of predation by prolonging theperiod of predators spending in the open water. Forexample, Leptodora kindtii (Focke, 1844) predationon Daphnia galeata (Sars, 1864) advanced by 13 daysper degree warming in Bautzen reservoir, Germany(Wagner and Benndorf 2007).Climate warming may allow increased predationpressure on zooplankton due to improving growingconditions and decreasing winter mortality of fish(Moore et al. 1996; Mehner 2000; Benndorf et al.Shift in phenology123
Aquat Ecolof thermal stratification in spring resulted in a forwardshift of the phytoplankton bloom. While the rotiferKeratella cochlearis (Gosse, 1851) appeared to adaptits phenology, Daphnia pulicaria (Forbes, 1893) wasnot able to do so and declined in abundance. Theauthors proposed that the different life history strategyand thus the different hatching cues used by thecladoceran may explain the mismatch. For the marinealternatives, see Edwards and Richardson (2004) andRichardson (2008). In order to interpret these shifts inphenology, Visser and Both (2005) called for ayardstick, a measure that will reflect a species successor failure to match its environment under climatechange. Such a yardstick can be the shift in phenologyof food abundance. Decoupling of the trophic relationship between the keystone herbivore Daphnia andits algal prey can result in the absence of the springclear-water phase as shown by De Senerpont Domiset al. (2007b) with a seasonally forced predator–preymodel.Climate change-related change in lake physicalproperties and subsequent change in algal numbersand community structure might explain much o
Springer Science Business Media Dordrecht 2012 Abstract Recently, major advances in the climate– zooplankton interface have been made some of which appeared to receive much attention in a broader audience of ecologists as well. In contrast to the . in the tropical Pacific (Allan et al. 1996) and the Pacific Decadal Oscillation (PDO .
Plankton samples were preserved in 4% buffered formalin. A total of 108 zooplankton samples were collected from March 2011 to February 2012. Zooplankton abundance was standardized to the number of individuals per 100 m3 (ind 100 m-3). The identification of each zooplankton species in a diverse zooplankton community is
what extent diVerent zooplankton, especially proto-zooplankton, feed on the diVerent life forms of Phae-ocystis in situ. To better understand the mechanisms controlling zooplankton grazing in situ, future studies J. C. Nejstgaard (&) UNIFOB AS, Department of Biology, University of Bergen, Bergen High Technology Centre, 5020 Bergen, Norway
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The Offshore Zooplankton and Ichthyoplankton Characterization Plan is an effort to collect data that will be used to describe the plankton assemblage in the northern Gulf of Mexico (GOM) and to inform the evaluation of potential impacts from the Deepwater Horizon Oil Spill (DHOS) to the zooplankton and ichthyoplankton in the region.
Two new integrating samplers for zooplankton, phytoplankton, and water chemistry. Archiv fur Hydrobiologie 75: 244-249. Lewis, W.M. Jr. 1978. Comparison of temporal and spatial variation in the zooplankton of a lake by means of variance components. Ecology 59: 666-671. Lewis, W.M. Jr. 1980. Evidence for stable zooplankton community structure .
www.helcom.fi Baltic Sea trends Indicators HELCOM core indicator report July 2018. Zooplankton mean size and total stock (MSTS) Key Message . This core indicator evaluates zooplankton community structure to
2.1 Ekosistem Kolam . Zooplankton memegang peranan penting dalam jaring makanan di perairan . Pada proses selanjutnya zooplankton merupakan sumber makanan alami bagi larva ikan dan mampu mengantarkan ke jenjang trolik yang lebih tinggi. Menurut Nybakken (1992), hanya satu golongan zooplankton yang penting, yaitu Subkelas Copepoda .
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