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*Corresponding author Komzina str., 10, Togliatti, [email protected] 445003,Russia;e-mail: of the Oka riverbasinare presented.of theOka content anditschangesintheforestscarbon issuggested.warming The mapsonthe offuture scenarios greenhouseknown under biotic regulation cycle ofthecarbon changing climate are described. Possible stabilizing thecontinentalbiosphere under offoresttypes formations capableof of forest associationsandzonal/regional modelsforempirical-statistical identification analyticalandcartographic constructing inforestcycle ecosystems. for Methods evaluation ofbioticregulation ofthecarbon methodsfor of landscape-ecological This paperpresents theresults ofapplication of carbon poolsindifferentof carbon componentsof is placedonaprocedure for determination 2006; etc.]. However, whileastrong emphasis Climate..., 1996;Aberetal, 2001;Celniker, both inRussiaandabroad [Gorshkov, 1994; highlighted inquite anumberofpapers system. These problems have already been inthesoil–vegetation–atmospherecycle studying thebioticregulation ofthecarbon climate isconnected, inmany respects, with of thecontinentalbiosphere underchanging Understanding mechanismsofsustainability prediction. modeling, approach, ecological cartographic climatic changes, statistical empirical Erland G.Kolomyts 1 KEY WORDS: ABSTRACT INTRODUCTION 6 PREDICTIVE MODELING PREDICTIVE WARMING: LANDSCAPE-ECOLOGICAL ECOSYSTEMS UNDER GLOBAL CARBON BALANCE OF FOREST 9 Institute ofEcologyInstitute ofthe Volga Basin,RussianAcademy River ofSciences, ecosystem, biogeocoenoses, 1 *,Gennady S.Rozenberg*,Gennady of factors effecting bioticregulationof factors ofCO predictions oftheclimatogenic dynamics lesser extent. Thus, there no are actually climatic changesare illustrated to amuch to global cycle components ofthecarbon assessment ofresponses ofdifferent biotic other ecosystems, problems ofquantitative content inforests,carbon wetlands, and andthecurrent cycle ofthecarbon functioning phytobiota andinsoil, aswell asonthe providing for ofthecontinental thestability The forest cover isoneoftheleadingfactors developed insuchstatement oftheproblem. inthebiosphere hasnotbeenyet cycle concept ofbioticregulation ofthecarbon heterogeneity ofbiogeocoenoses. The iscreated bystructural cycles primarily de Haes, ofbiological 1994].Spatialdiversity andUdo Canada, andUSA,respectively [Klijn phase” inclassificationsofAustralia-Britain, “site”,categories “ecoelement”, and “landtype accepted inRussiacorresponds to the unit ofgeographical ecology elementary functional “core” [Sochava, 1974]. This ofnaturalenvironment, i.e., part active its topological level, isthemostcomplexand (landscape facieses),i.e. ecosystems ofthe the Le Chatelier’s principle. Biogeocoenoses of naturalenvironment inaccordance with thatprovidesthe biological stability cycle studies oflocal-regional mechanismsof biotic regulation may beachieved through Further development oftheconcept content intheatmosphere. 1 , LarisaS.Sharaya 1 003.08.2011 14:38:41 2 3

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7 0 developments [Climate..., 1995;Aberetal, developed. prognosticpoorly International dynamics underchanging climate have been The problems offorecasting forest ecosystem severalhumus may decades. take litter, coarsewoody debris, andmobilesoil presenceduration ofcarbon intheforest are ofthesameorder, however thetotal forests [8].Changesin decompositionrates within ayear ofbroadleaf inthesub-zone years inthetaigazone andare completed thefirstseveralproduction appearduring intheratesClimatogenic ofphytomass shifts changesisrelativelyof thesefunctional small. production anddegradation). Atemporal lag (phytomass insmallbiological cycles shifts changing climate bythe is determined under Biotic regulation cycle ofthecarbon thevegetationchange during period. ecological reconstructionsistemperature significant. offuture The leadingclimaticfactor temperaturewith theaverage isnon- January average Julytemperature, whileitsassociation itselfdependsmainlyonthe thisfactor turn, In parameters. andmainfunctional distribution predeterminingfactor ecosystem territorial ecological moisture whichisthemostpowerful system passesthrough thesummersoil ecosystems withthebackground climatic ofregionalchannel ofconnections andlocal established thatthemain been empirically the Volga has basin[Kolomyts, River 2008].It forest ecosystems intheheadwater area of regional environmental forecast studiesof This problem wasaddressed intheprevious increasing greenhouse effect. is considered oneofthemaincauses [Gorshkov, 1994].Global-scaledeforestation temperature regime oftheatmosphere andthe cycle 1972], includingthecarbon [Sukachev,natural biogeochemicalcycles most potent regulators andstabilizers of biosphere. Forest ecosystems are the APPROACH TOPROBLEM THE THE LANDSCAPE-ECOLOGICAL BACKGROUND experimental test sites of the Middle and test sitesexperimental oftheMiddle outbytheauthorsatsix carried surveys landscape-ecological oflarge-scale Materials resolution. 2001], buttheyhave ahigherlevel ofspatial instance, atsimulationmodeling[Aber etal, obtained are probably notasaccurate as, for most informative characteristics. The results possible to withasmallnumberofthe work concept [Sukachev, 1972]. italso They make biogeocoenoses oftheSukachev’s in theaspect systems astheindivisiblecomplexformations behavior ofgeo-(eco-) models describe [Rozenberg, 1984;Kolomyts, 2008]. These systems much more complicated geo-(eco-) of discrete modelsof empirical-statistical presented herein isbasedonconstruction prediction The locallandscape-ecological ecosystems to climaticchanges. that ensure theresistance ofterrestrial the atmosphere andto identifymechanisms and surface exchange theearth between the phytobiota effect onthecarbon to findtheorigins ofthemechanisms regional conditions. itpossible This makes biogeocoenoses underdifferent zonal/ of bythespatialdistribution determined ofthisregulationdiverse types mechanism allowsoneto identify landscape position.It regulation phenomenafrom abroader We offer theapproach to analysisofbiotic topological oftheecosystems. diversity biomes,sites intypical ratherthantheentire onlytypical limited becausetheydescribe areas. However, thesemodelsare spatially homogeneous ofstructurally characteristics cover onthebasisoffunctional selected with comparatively smallareas oftheforest of landscapemosaic[Forman, 1995]deal studied.been poorly models The so-called ecosystems underclimaticchangeshas addition,behavior oflocal climatic factors. In forest communitiesto thesamebackground ofresponsesfor of thespatialvariability cannot reveal basicmechanismsaccounting 2001; etc] are mainlysmall-scaleand, hence, FIELD DATA FOR MODELING 003.08.2011 14:38:41 3 . 0 8 . 2 0 1 1

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7 1 under different zonal/regional conditions of the Middle and Upper Volga Regions, according to the the to according Regions, Volga Upper and Middle the of conditions zonal/regional different under Fig. 1. Landscape couplings of the predicted carbon balance dynamics in forest biogeocoenoses biogeocoenoses forest in dynamics balance carbon predicted the of couplings 1. Landscape Fig. ΔС(Fa) balance (t/ha); –carbon T,... E, sun –sunny, TE, slopes: sites –local (see exposed solar inthetext); scenarios of HadCM3 model. shad andneutral –shady 003.08.2011 14:38:41 3 . 0 8 . 2 0 1 1

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7 2 (Hadley Centre ofClimate Research, UK) [Climate..., 1996]–HadCM3,version A2 ocean generalcirculation models (AOGCMs) global coupledatmospheremost extreme to 2100wereperiod from taken oneofthe The regional for climate scenarios the forest litter mass, humus insoilorganic-mineral layers, mass (brushwood anddead-wood), (2) root mass, phytomass, tree-shrub balance: (1)skeletal ha a particular (Fig. 1).Eachofthesetest sites characterizes forest-steppethe southern to mixed forests wide rangeofzonal plantformations, from forecast analysis. These test sites containa in amethodspecificallydeveloped for the Upper Volga Region (1987–1998)were used The following phytomass parameters ( regions, considerablydistant. often backgroundreflecting conditionofother other geotopes were considered extrazonal, a given ecoregion). The biogeocenosesof ofzonal for bioclimaticnorm a localtype occupies eluviallocationsandrepresents within thebiogeocenoses(thisgroup (placor)groupflat interfluve wasisolated regional level couplings[Polynov, 1956]. The ofthemesoreliefat types characteristic A similarsystem oflandscapecouplingsis 1964]. [Glazovskaya, (Saq), according to theclassification (TA) to theaccumulative (A) andsuperaqual andthetransaccumulative slope transit(T) oflocalsites (geotopes)types through the from theeluvial(E)andtranseluvial(TE) direction ofedaphicmoistening increase: landscape couplings(microcatenae) inthe were located alongthevector ofthelocal in eachofthetest sites. The faciesgroups been isolated basedon45–50sampleareas offorestdiversity andtest sites types have Six faciesgroups thatrepresent general Town, etc. Farmstead,Shchelokovsky Green By-Sura, corresponding conventional name(Zhyguli, ) were usedfor calculationofthecarbon regional ecosystems BR ; ML (3)total green mass, ; (5) debris – dead skeletal –deadskeletal ; (5)debris , with the HU WD BV . ; (6) ; (6) ; (4) ; (4) BS t / ;

the 21 in theMiddle Volga Region bytheendof 7.0 [Climate ...,1996]. The modelforecasts a5.5– beganattheend ofthisperiod warming represented thebaseline. Recentglobal formeteorological 1881–1985 observation 2050–2075, and2075–2100. of The period 1985–2010, 2010–2030,2030–2050, (20–25yrsduration): included five intervals The entire forecast inouranalysis period areas. both nemoralandboreal forests withinlarge endurance andcausethedisintegration of changes thatmay exceed thelevel oftree ecosystems feedback to globalclimatic oneto evaluatethis modelpermits theforest [Pope etal, 2000].Ecological by prediction C unchanged; therefore, thehumidifyfactor the annualprecipitation willremain virtually when biotareleases CO source,sink into thecarbon i.e., thesituation the transformation ofbiotafrom thecarbon violation oftheLe Chatelier’s is principle terrestrial andoceanbiota.Anapparent utilizationbythe enhancement ofcarbon atmosphere isaccompaniedbyanequal spontaneous increase ofCO state ofenvironment willbestableifany following postulate [Gorshkov, 1994]:the natural environment isexpressed bythe The Le Chatelier’s for of principle stability parameters offorestfunctional ecosystems. analysis ofclimatogenic dynamicsofthe asthebasisforassumed thisscenario our in theentire Middle Volga Region. We have climatictrend willbedeveloping thermo-arid Volga region. Thus, untiltheyear of2100,the and from ofthe 1.37to 0.65insub-taiga forest-steppesouthern oftheZhyguli region topoecosystems underdifferent zonal/ offorestof metaboliccharacteristics assessed usinghydro-thermal ordination The potential ofbioticregulation hasbeen influence. under external METHODS OFMETHODS FORECAST ANALYSIS hum ° increase ofthemeanJulytemperature willdecrease from 0.95to 0.36inthe st century. According to thisscenario, 2 into theatmosphere 2 content inthe 003.08.2011 14:38:42 3 . 0 8 . 2 0 1 1

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bold font. **) *) ( is notalways high,thoughquite significant As canbeseenfrom Table 1,thecorrelation ofthe0–50smsoillayer.moisture reserves depth of50cmandbysummerproductive bysoiltemperature atthe was performed changes ofbiological cycle). The ordination simulation ofanticipated climatogenic regional andlocalconditions(anempirical S so-called forestry method based onthe forestry so-called different bioticcomponents, we usedthe To content in assesschanges ofthecarbon under theinfluenceofgeophysical trend. ofchangesgiven tendency parameters general can beinterpreted onlyasacertain 7 P y 3 parameters of diff x y =b

y =b y =b

— coeffi (calculation R Model type Model type 1 y = formula) *) n – —coeffi Table 1. Examples of calculation models that describe changes of biogeocoenotic biological cycle cycle biological biogeocoenotic of changes describe that models calculation of Table 1. Examples + b soiltemperature at50cmdepth( + b + b + b exp 0.05). 0 0 0 +b +b +b 1 2 2 2 /

cients ofstandard deviation.Signifi cantarguments inequationsofmultiple regression are showninthe x x x x (b 2 1 1 1 2 1 1 2 2 cient ofcorrelation; P–signifi cancelevel (the Pearson criterion); ) x 0 x x + 1 2 1 +

+ + At a weak connection, the latter thelatter At aweak connection, Prioksko-Terrasny Shchelokovsky Shchelokovsky Shchelokovsky Shchelokovsky Shchelokovsky Shchelokovsky Regional eco- erent regional ecosystems in the Middle Volga Region under trends of changes in soil soil in changes of trends under Region Volga Middle the in ecosystems regional erent temperature and soil moisture content during the vegetation period vegetation the during content moisture soil and temperature (see Fig. 1) Kerzhenets systems By-Sura Zhyguli Zhyguli reserve Zeleny Zeleny Gorod Gorod Hutor Hutor Hutor Parame ters (see inthe t 50 ); text)

WD WD WD HU HU HU HU HU ML ML ML x BV BR BR BS BS BS 2

– stored soilmoisture in0–50сmlayer (W-50). 3.3097 .00 .6 90.49 0 0.769 0.00403 0.9472 –34.93 4396.306207.0 0 0.642 64.63 –4.389 .853.8056016.0 0 0.506 35.08 0.0845 53 .1900020570.4•10 0.577 0.00042 0.0159 25.31 4. 14751209602.21 0 0.936 5.142 –114.7 642.1 40 031 .09 .9 17.75 0 0.699 0.00091 –0.3611 44.08 28 028 .04 .2012.78 0 0.72 0.00149 –0.2588 22.85 71 –2.333 27.18 –0.2402 16.34 .6 62 .1 12.25 39.3 0 0 0.614 0.611 26.26 15.42 1.768 3.706 19 –0.7415 21.93 4. .96–.01 .5 0.7•10 0.557 –0.00312 0.3956 244.6 5.838 8.712 25011 .4 9.31 0 0.649 0 0.1712 12.5 0 5.99 .4009 .0608108.03 0 0.851 0.0006 0.0091 8.84 b 0 different of theforecast periods for each the soil-plantcover andtheatmosphere for flowsbetween possible changesofthecarbon This balanceequationwasusedto calculate +ΔC( follows: plant-atmosphere system wascalculated as change ΔC( for oftime[Celniker, longperiods 2006]. The components cycle assessments ofthecarbon method gives thebest results atbalance dynamics oflive anddeadphytomass. This Coeffi b cients 1 ΔС( HU Fa 0.04896 0.07426 –0.7135 –0.0678 ) –ΔC( Fa 55.75 ) =ΔC( ) of the carbon flowinthesoil- ) ofthecarbon b 2 BS WD ) –ΔC( .9 20.29 0 0.696 .1 6.51 0 0.717 .1 0 0.713 .6 3.09 0 0.664 .8 0.1•10 0.582 RPS ) +ΔC( characteristics **) BV Statistical ) –ΔC( ML ) + –3 –3 –3 BR 17.92 16.32 ). (1) 71.4 y 003.08.2011 14:38:42 3 . 0 8 . 2 0 1 1

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7 4 becomes asource ofadditionalCO or,by globalwarming onthecontrary, induced inthebiological cycle shifts of CO of theequationprovide for anincrease At positive values, thefirstthree members may have bothpositive andnegative values. oftheequation(1) part member oftheright ecosystems (mentionedabove). Each group offaciesinallexaminedsixregional Topography atresolution 3 Mission from theNASA Morphometric oftherelief. ecosystems withmorphometry oflocal characteristics and functional of thestructural statistically significant linkages (1 : 50–100,000) wasbasedonpredetermined, thelocallevel, mapping large-scale At et al, 2002]. parameters introduced byP. [Shary A.Shary values. Herein, we used asystem of18 are confinedto sixbasicmorphometric methods ofaquantitative relief description the analyzed territory. The currently known ofsignificantly theboundaries extending calculated) onrepresentative test sites, thus the analysis)ofdatameasured (or (substantiated duringand extrapolation ofrelief allowinterpolationproperties Multidimensional environment-forming etal, 2002]. analysis [Shary, 2001;Shary oftheforecast part used inthecartographic were New methodsofgeomorphometry emissions. of CO ecosystems consumesadditionalamount which showswhetherthisgroup offorest the atmosphere byΔС( isdescribed exchange biogeocoenosesand between the overall balanceofchangescarbon the results are quite theopposite. Thus, At negative valuesoftheabove parameters, deposit). live phytomass plays therole ofitssink(i.e., source,as anadditionalcarbon whilethe thiscase, thedeadphytomassIn poolacts members provide for adecrease ofthisflow. to theatmosphere, whiletheotherthree 2 2 emissionfrom thesoil-plantcover from theatmosphere dueto the parameters were derived data (SRTM3, Shuttle Radar ShuttleRadar data (SRTM3, '' arc seconds arc seconds Fa ), 2

the total C sources – ofthebaselineperiod ecoregion: 12.6 significance at resolution 30 ShuttleRadar (SRTM30, Topography Mission, project result ofimplementationSRTM elevationsobtainedasa surface of earth thematrices NASA satellite dataontherelief: Regional mapswere usingthe constructed Table 3. other predictors are given inthenote to Here, 25.3 • the changeintotal Cstocks by2050– extrapolation. extrapolation. The methodutilizes the using ourmethodofinduction-hierarchic out maps at1:2,500,000scalewas carried level offorecast of withconstruction Transition from thelocalto theregional 2001]. using program “Analytical GISEco” [Shary, plant formations andtheircombinations for predictive for periods performed the of changesinthetotal Ccontent hadbeen statistical analysisofspatialdifferentiation advance,Russia witha900mgrid. the In Kavraysky forVII INTprojection theEuropean for thisbasinwastransformed into the attheregional level.territory The matrix asthemodel basinwastaken river The Oka version2_1/SRTM30/). on equator, http://dds.cr.usgs.gov/srtm/ or 90equator,on m correlation coefficient equations (withthecorresponding Spearman instance, thefollowing multipleregression projectionforKruger the9th6 spacing) of50mrecalculated into theGauss- withacellsizeelevations asmatrices (grid on srtm/version2_1/SRTM3/) C Q base H Δ +18.0• – mean curvature; designations of –meancurvature; • C =222.6–55.6 H 2050 =–99.8.+47.1• ; ( P r s ) were obtainedfor theZhyguli =0.87, H '' ; ( arc secondsor900m r s =0.88, P <10 http://dds.cr.usgs.gov/ r • S

andthelevel of GA the earth surface surface the earth –6

P – 25.6 ); (2) <1–) (3) <10–6). GA ° zone. For • +

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couplings (E the linksofchainregional landscape mesocatenae wasbrokeninto mesosites, i.e., formation represented asapoly-vector setof (microcatena). Then, eachrangeofagiven spatially ordered system oflocalz theregional characterizes This series inagiven series and extrazonal ecoregion. biogeocoenoses from theirflatinterfluve groupmap wascorrelated of withacertain distinguished onthesmall-scalegeobotanic ofplantformations Each type/subtype The procedure wasasfollows. ofextrapolation ecosystem changes. asamodelofthemaintrends of may serve the predicted climate changes. These rows (see above) thatcorrespond to thevector of biogeocoenoses andextrazonal interfluve are composedofthevector rows offlat environment. These regional systems on thebackground regional-bioclimatic local geomorphological andedaphyc factors oftheinfluence on theidentifiedpatterns systems were of localzonality isolated based changes [Kolomyts, 2008]. The regional a form oftheirresponse to globalclimatic systems as oflocalgeo-(eco-) poly-zonality establishedphenomenon ofempirically mosaic was, then,supplemented with the Thus obtainedtheregional phytocatenae of acorresponding geobotanicalrange. groups ateachtest sites into mesogeotopes aggregated thebiogeocoenosesofallisolated imageofmicrocatenae,a homomorphic we couplings [Polynov, 1956]amesocatena is B.B. Polynov’s conceptionoflandscape Assuming thatinaccordance withthe Prioksko-Terrasny reserve. Green Town, andtheVyksa, Kerzhenets, Farmstead,of thetest sites Shelokovsky regression equationsappliedto thedata such mapwascompiledusingmultiple area oftheregion. For riverbasin, theOka of mesoscalesites for theentire forested possible to create anintermediate map statistics were applied. Then, itwas shady). Newmethodsofgeomorphometric g solar-exposure two with divisionofthe TE and T locationsinto 7 5 → TE → T radations (sunny and → TA, A;seeabove), onality onality the comingcentury, thecontent ofcarbon and 180–243t/ha,respectively. Therefore, in the third regions were 140–190,134–2,112, baseline valuesfor thefirst,second, and and inthebiogeocenosesgeneral. The phytomass flowsinthelive skeletal the carbon potential rangeofclimatogenic changesof +(115÷120) t/ha. These valuesrepresent the Δ Volga region. valuesof changes The extreme broadleaf forest Novgorod oftheNizhnyi to +(95-100)t/hain TE andEofconiferous- Farmsteadthe ZhiguliandShchelokovsky 62) valueswillrangefromin itsextreme –(37– contentchanges ofthetotal [ΔC( carbon 2100, 2).By (Table ofproductivity the shifts biomass, represents of thegreatest part phytomass (BS),mainlyincrement ofwoody biogenesis part. tree-shrub The skeletal butinitsautotrophic ofthecycle, part changes occurnotinthedecomposition of time, themaximal(byabsolute values) same time, sufficientlylongperiods during toshould react climaticchangesfirst.Atthe 2009]; therefore, thedecompositionpart [Kolomyts, part changes thantheproductivity asarule,cycle, ismore sensitive to climatic 1. ofthebiological The decompositionpart and Rozenberg, 2009]. publications [e.g., Kolomyts, 2008;Kolomyts inmoredescribed detailinourprevious areprediction given below(theyare The mainresults oftheempirical-statistical Predictions by Models Analytical surveys. offieldlandscape-ecological materials predictive mappingbaseddirectlyonthe fundamentally novel methodofregional isa of induction-hierarchicextrapolation environment. algorithmThe described ofthegeographicregional types/subtypes representative membersofdifferent zonal- aslocal of thebiogeocoenosesserving baseline orpredictive metabolicparameters DISCUSSION MODELING RESULTS AND THEIR C ( Fa t / ha ) will comprise from) willcomprise –(78÷100) t/hato inEand TA ofoak-linden forests of BS 003.08.2011 14:38:43 )] 3 . 0 8 . 2 0 1 1

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7 6 baseline period in fl in period baseline Table 2. The forecasted component and balance changes in the carbon content ( content carbon the in changes balance and component forecasted The Table 2. (see inthetext) Parameters aac 3.8–71 2.9–32 3.8–31.32 –31.38 –33.24 –29.09 –27.61 –27.19 –26.17 137.88 –29.0 Balance –29.25 70.9 –31.24 58.15 239.43 43.91 Balance 30.0 14.77 –23.33 201.36 –28.39 –34.42 Balance –37.34 –40.56 –83.11 211.41 –77.29 –69.91 Balance –61.09 –50.51 198.61 Balance WD WD WD WD WD HU HU HU HU HU ML ML ML ML ML BV BV BV BV BV BR BR BR BR BR BS BS BS BS BS at interfl Base values 5.9–71 2.5–77 2.9–27.91 –27.99 –27.76 –27.85 60.19 –27.14 49.82 155.09 38.16 26.61 2.90 13.73 –3.31 134.14 –10.84 –14.77 –18.99 123.42 36 24 34 35 36 –4.09 –3.45 –3.66 –4.73 –12.59 –3.50 –4.71 –11.60 –3.41 2.51 –11.68 –4.16 –2.45 2.48 –9.81 –3.91 –7.17 –4.32 13.69 2.55 –9.44 –7.15 10.15 –4.34 17.08 2.52 –7.21 –4.28 –3.43 –7.19 2.84 –4.30 –3.11 17.06 –7.44 12.79 –4.11 –2.65 14.28 15.66 42.41 –2.28 11.09 –1.71 –14.76 7.99 –13.53 –5.64 11.16 4.44 –11.45 –6.46 39.48 –11.09 –7.82 –9.76 –21.66 –8.04 –8.80 –20.68 31.64 –8.88 –8.45 –19.72 –18.62 –18.06 40.40 –39.66 –8.02 –15.29 –16.13 –35.78 –7.53 –10.54 –14.08 –31.55 –6.98 61.47 –11.90 –27.32 10,24 –23.05 30.25 82.78 .504 02 03 04 –0.64 0.60 –0.40 0.86 –0.31 –11.15 0.82 11.14 –11.85 –0.26 –11.86 1.19 12.69 0.40 –1.88 –12.64 9.58 1.21 7.85 –13.00 –1.87 9.44 4.41 84.7 –1.89 –2.38 6.58 –3.84 –1.88 3.30 –2.05 –3.80 7.48 –1.98 3.01 –1.60 –3.73 6.00 –1.00 2.65 –3.63 –2.79 –0.13 –1.91 2.32 –3.46 –2.45 –1.13 –1.71 5.48 1.90 4.06 –2.03 –0.93 –1.46 6.59 –1.60 8.99 –0.84 –1.23 –1.28 7.78 –0.61 –1.06 –2.26 5.88 6.34 –0.60 4.90 –2.10 4.90 5.17 –1.93 3.56 –1.77 5.24 –1.60 4.39 uve forest biogeocoenoses under diff under biogeocoenoses forest uve Middle and Upper Volga Regions Volga Upper and Middle d 0023 0027 2100 2075 2050 2030 2010 ) Prioksko-Terrasny reserve e ) Kerzhensky reserve ) Kerzhensky c ) Green Town b a ) By-Sura ) Zhyguli Changes in predicted periods Changes inpredicted erent zonal-regional conditions of the the of conditions zonal-regional erent t / ha ) relative to the tothe ) relative 003.08.2011 14:38:43 3 . 0 8 . 2 0 1 1

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change by ecosystems oftheMiddle Volga region may stored phytomass intheskeletal oftheforest and debris and debris results intheincrease of forest litter mass above ground deadorganic matter. This will decomposition andmineralizationofthe increase andimpede underglobalwarming the zonal forest-steppe ecotones will At thesametime, soilmoisture deficitof be insignificant. balancewillto thechangesincarbon change little;therefore, theircontribution forests watersheds, ofwell-drained will sunny slopesandthatofpinesteppified meadow-steppe communitiesonsteep offorest-biogeocoenoses, particularly sequestrationofxeromorphicrate ofcarbon reduced.markedly Atthesametime, the forestsEA–Saq birch-spruce-pine willbe zone, themosthydromorphic Aand glacial plainsofthenemoral-forest sub- phytomassskeletal within theaqueous- sequestrationintheThe rates ofcarbon willdiminish. carbon sink ofatmospheric uplands (seeFig. 1,A–C). Their role asa lime andoakforests oferoded-loamy mesophilicelm- inflatinterfluve quickly phytomassskeletal willdecrease most communities. depositioninthe Carbon potential offorestof theproductivity an unambiguousandquite cleardecline the middleforest-steppe, there willbe and ofthesouthern Within theterritory ecosystems (see Table. 2). the valueΔ( may beevaluated cycle by of thecarbon Zonal-regional contrastsofbioticregulation balance.which willdirectlyaffect thecarbon of similarmagnitude offorest productivity, an abruptdrop or, inothercases, anincrease willinduce,2. Globalwarming insomecases, the atmosphere. exchange intheircarbon with productivity of theregional mixed andnemoralforests’ points to quite asignificant regulating role compared to thebaselinevalues. This fact greater mineralizationofthe humusmass 7 7 ± BW Fa (25–50)% andmore onaverage ) for the eluvial (flat interfluve) ) for theeluvial(flatinterfluve) . However, there willbeeven ML

result inadditionalrelease ofCO humus massaswell. These processes will its mass(includingtheforest litter) andthe and, correspondingly,further willreduce decomposition ofdeadorganic matter even trendarid willintensify theprocesses of Fig. 1,D).Ontheotherhand, thethermo- will increase aswell (see Table 2, this phytocoenotic sink poolasacarbon and, asaconsequence, thesignificance of of thesetopoecosystems willalsoincrease intheperennialphytomasscarbon skeletal Accordingly, ofatmospheric conservation zone willincrease (andquite significantly). ofthesub-taiga strip and ofthesouthern forest-steppe boundary) zone (atthenorthern coniferous forest biogeocoenosesofthe living organic matter ofthemixed anddark of andreserves productivity The primary the atmosphere (see Table 2, steppe. conifer andmixed The flatinterfluve forests forest- andtypical inthesouthern degradation anddisappearance ofmarginal willinevitablyaccelerate theprocess of It the continentalbiosphere willbedisturbed. one oftheleadingstabilizingmechanisms as cycle the naturalbalanceofcarbon to the atmosphere.of carbon Accordingly, the source ofsignificant additionalemissions southoftheforestThe extreme zone willbe system willbedeveloping ontheRussianPlain. regulation inthesoil-vegetation-atmosphere biotic cycle antithetical phenomenaofcarbon Under global warming, mutually two 3. (see Table2, additional release ofCO with higherdegree ofitsutilizationand abruptly degradation ofdeadorganic matter willincrease temperature andsoildrying rise willbemore explicitandintensive.cycle Soil ofthebiological part Changes inthedetritus amount ofCO whichmustresultproduction, inalower induce adecrease ofthetotal annualforest trend zone, will sub-taiga thethermo-arid forestedOn theoutwash lowlandsofthe soil-plant cover to theatmosphere. HU resulting inadditionalCO e , andFig. 1,F, G). 2 boundbyphotosynthesis. 2 to theatmosphere a 2 ). emissions to 2 from the c , and 003.08.2011 14:38:43 3 . 0 8 . 2 0 1 1

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7 8 pine forests lowlands outwash ofsub-taiga Excessively moistforests dry andextremely zone. ofthesub-taiga strip in thesouthern plains inthenemoral-forest and sub-zone by mixed and coniferous forests ofhigh Le Chatellier’s willbeperformed principle) the greenhouse effect (according to the ecosystems andtheatmosphere reducing exchange theterrestrialcarbon between The mainpositive bioticregulation ofthe balance.carbon play similar negative role inthechangeof forests lowlandswill outwash ofthesub-taiga Fig. 2. The total carbon content in forest ecosystems in the territory of the Zhyguli ecoregion ( ecoregion Zhyguli the of territory the in ecosystems forest in content carbon total The 2. Fig. changes in the total carbon content for the predicted period of 2050 ( 2050 of period predicted the for content carbon total the in changes to HadCM3 model period upto 2050. period The fragments of C balanceofforestsmodels ofthecarbon for the As examples, letusconsiderthecartographic in Cartographic Expression Predictive Scenarios global warming. ofcontinentalbiospherethe stability under boreal forests thatwillessentiallysupport this istheonlygroup offormations of Probably, importance. will beofsecondary at the local level (Fig. 2). In at thelocallevel (Fig.In 2). ΔC( Fa ) mapsoftheZhyguli forests are presented b ), in according t/ha, the baselineperiod, base a ) 003.08.2011 14:38:43

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7 b. The changes in the total carbon stocks for theforecast for inthetotal changes b. stocks The carbon 9 fragmentary large tracts of forest among agricultural 10 among large offorest land; areas. tracts –forest ofsteppes area fragmentary 3–9 –thenorthern 2 –0 1 –10.2–15.0; 2–15.0–19.3; 3–43–130; 4–130–160; ofsteppes area 7–thenorthern 5–160–200; 6–200–423; a. The distribution of the carbon content ofthecarbon distribution (1985). The a. attheendofbaseline period (t/ha): stocks carbon The ò Fig. 3. The fragments of the maps on the total carbon content and its forecasted changes changes forecasted its and content carbon total the on maps the of fragments The 3. Fig. 5.5; 3 –(–115) ò (–89); 4–(–89) within the territory of the Oka river basin. river Oka the of territory the within ò (–38); 5–(–38) ò (–12); 6–(–12) period of2050period relative to (t/ha): thebaseline 1–(–1.3) ò 0; 7–0 ò 38; 8–38 ò 63; 9–63 ò 003.08.2011 14:38:46 115. 1; 2– 3 . 0 8 . 2 0 1 ò 1

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8 0 characterized bythemaximalCOcharacterized well asthebottoms ofsmallvalleys, willbe theforecastIn period, theselocations, as the Zhyguli withtheleastcatchment area. the gentlenear-watershed shadyslopesof stocksthe highesttotal areof carbon typical Note: Predictors inregression equationsare: regions cycle ofthepositive bioticcarbon pine andsmall-leaved forests. The other Volga Uplandthatisoccupied bycomplex oftheBy- part ofthenorth-western typical from +(65–100)to –(80–136)t/ha),is varying patchy of(ΔС( distribution A particularly balance.and negative valuesofthecarbon contrast andfrequent patchiness ofpositive The forecast mapshows alsothepronounced (see Fig. 1,E the pools,phytomass includingtheskeletal content inmostof changes inthecarbon be ofany significance dueto relatively low However, thescaleofthisregulation willnot regulation. cycle positive (asawhole)carbon will perform forest cover ofthisterritory Table 3.Underglobalwarming, themodern for ofthemapsare construction presented in (Fig. 3).Multipleregression equations used basin river mentioned above, bytheOka The regional level isrepresented, as willbeinsignificantcarbon too. are minimalandtheforecasted release of On steep slopes, stocks theinitialсarbon southern azimuth(180º),sundeclination35º; southern thatdefimaximal curvature nesthe “ridge” landscape forms; ruso ln omtosRegression equations Groups ofplantformations Table 3. The equations of connection between carbon content changes and the most important important most the and changes content carbon between connection of equations Table The 3. а broadleaf forests forests ofbasin) (allsecondary e forests areas) ontheircontemporary f pine forests d c b ) Primary and secondary forests ofbasin(all andsecondary ) Primary rala oet ΔС( ) Broadleaf forests ) Secondary substitutingfor spruce, pine,) Secondary and environment factors in forest formations of the Oka river basin to the year of 2050 (ΔС( 2050 of year tothe basin river Oka the of formations forest in factors environment pueadbodefsrc oet ΔС( ) Spruceandbroadleaf-spruce forests ieadbodefpn oet ΔС( ) Pine andbroadleaf-pine forests ) Secondary substitutingfor pineandbroadleaf/ ) Secondary , F). 2 emissions. Z T –elevation; –average temperature ofJuly. Fa ) MCA ΔС( ΔС( ΔС( Q maximum carbon depositionmay beuptomaximum carbon intense growth ofstands (Fig. 4, where thetemperature willinducean rise atrelativelydeposit carbon dry TE mesosites, and broadleaf-spruce forests willactively regulation (Fig.cycle 4, Table 4).Spruce riverbasinto thetotal bioticcarbon Oka separate groups offorest formations ofthe of to assessthe contribution isnecessary It of Cartographic Forecast Territorial Generalizations cycle. carbon positive bioticregulation ofthe most part, for marked, the land willexhibitaweakly forests ofagricultural onthevastterritories bankofthe right Volga river. Fragmentary downstreamriver area andtheneighboring bankofthe Klyazma left lowland-outwash forestsspruce andspruce-birch ofthe Central RussianUpland, andto broadleaf- ofthe forests boundary ofthenorth-west to oak-limecharacteristic andbirch-aspen (0 to –12;occasionally, up to –38 t/ha) willbe negative Weakly marked are mainlyprevalent. broadleaf, spruce, andbirch-aspen forests there,slopes oftheSmolensk-Moscow ridge; andtheeastern Meshchera of Ryazanskaya regulation are theswampyforested lowland –insolationofslopeswiththeSunpositionat Fa Fa Fa Fa Fa Fa –catchment areas; –catchment ) ) ) ) ) ) 2050 2050 2050 2050 2050 2050 = 28.8 = 37.5 = 83.1 = –76.2 = –24.0 = 76.7 MCA MCA k Z max T Z +46.7 +21.0 + 42.9 +76.6 +23.9 –18.4 GA T MCA +24.0 MCA –degree ofslope; MCA GA k max +12.1 –7.1 –10.9 +9.7 –7.2 Q –63.8 Q T GA T T +12.1 –61.6 –7.8 +12.7 Δ +5.1 C( Fa Fa k ) ) values 2050 max a – ). The ) 003.08.2011 14:38:48 3 . 0 8 . 2 0 1 1

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8 b) small-leaved formations Secondary inthe place offore 1 a) formations: Aboriginal 1– spruce andbroadleaf-spruce leaf-pine; 3 – broadleaf. E, TE, T, of mesosites (see TE, (see E, 3–broadleaf. Fig. sun exposure inthetext); andshad – solar 1)leaf-pine; types Fig. 4. The changes in the total C stocks in the groups of plant formations of the Oka river river Oka the of formations plant of groups the in Cstocks total the in changes The 4. Fig. basin predicted for 2050 under different conditions of plain mesorelief plain of conditions different under 2050 for predicted basin forest; 5–flood-plain forest; sts: 1–sts: spruce andbroadleaf-spruce; andbroad- 2–pine ; 2 – pine and broadleaf-pine; 3 – broadleaf; 4 – paludal 4–paludal 3–broadleaf; andbroadleaf-pine; ; 2–pine 003.08.2011 14:38:48 3 . 0 8 . 2 0 1 1

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8 2 4.6–7.5 tht/km and T slopes, especiallythosefacingnorth. The E oak-lime forests ofriver valleysandadjacent deposition sinks.minor carbon The main role incarbon even slopes. Broadleaf forests as willserve forests sites andofgentle offlatinterfluve inthelight-coniferousshould beexpected balanceNegative valuesofthecarbon the TE and TA–A elementsofthemesorelief. will accumulate mainlyinpineforests on the regulation Carbon cycle. ofthecarbon aconsiderablepositiveto contribution make riverbasin,pineforestsareas intheOka will Table 4andFig. 4, than thatofdark-coniferous forests (see willbe1.5 timesstronger,their contribution have positive i.e., cycle, effect onthecarbon Pine andbroadleaf-pine formations willalso balancewillbeminor.carbon to changesintheregionaltheir contribution an insignificant area oftheseformations, (see alsoFig. 4, 70% will exceed emissionsbynearly carbon depositionindark-coniferouscarbon forests decay andhumusmineralization.Overall, intensification oftheprocesses offorest litter emissionsasaresultthe carbon ofdrastic phytomass byskeletal willbesurpassed E and depositioninthe T territories, carbon therefore, 3.6–3.8tht/km exceed itsemissionsfrom peatlitter decay; phytomassin theskeletal willsignificantly significant intensification ofproductivity. groundwater level decrease willinduce neutral slopes, soiltemperature and rise contrasts ofΔС( tois expected have thehighestterritorial The group offloodplainforest formations by 2050. additionally fixed withinswampwatersheds their productivity. The CO a on theLe Chatelier’s (seeFig. principle) 4, regulation (based cycle main positive carbon Paludal forest formations willprovide for the emitters. ). Global warming willdrasticallyincrease). Globalwarming mesorelief asCO componentswillact will beplayed by mesohygrophytic 2 . Atthemore hydromorphic a Fa a); ). ). Ontheshadyand Occupying thegreatestOccupying however, becauseof 2 of carbon willbe ofcarbon 2 sequestration 2

content different insign (seeFig. 4, associated withchangesinthecarbon small-leaved forestsSecondary willbe positive. balanceoftheregiontotal carbon willbe ofthefloodplain forestscontribution to the th. t/km phytomassin theskeletal willbe5.7–6.6 mesocatenae, storage anadditionalcarbon theupper linksof in thesecommunities. In will induceaccelerated growth ofstands on sabulousloamy soils. trend Thermo-arid grow inhigherhydromorphic conditions taiga dark-coniferous forest associations and aspenforests substitutingfor sub- emission of4.4–4.8tht/km and sunny TE sites resulting inadditionalCO decomposition inthecommunitiesofeluvial the sametime, there willbeintensification of may have bothpositive andnegative broadleaf forests underglobalwarming by temperate coniferous, mixed, and Thus, regulation cycle thebioticalcarbon precursors. Δ of pattern ontheabove described impact lime forests arelatively willmake little aspen forests growing placeofoakand In balance (see Table 4andFig. 4, there willbeasignificant negative carbon place ofpineandbroadleaf-pine forests, small-leaved associationsexistinthe On lesshydromorphic sandysoils, where to theLe Chatelier’s principle. regulation according cycle positive carbon in types dark-coniferous taigaandsub-taiga role oftheboreal forest cover consistingof growing timber. This seemsto betheleading inthe to additionaldepositionofcarbon of restorative successions, mustcontribute presencethe permanent ofinitialstages spruce andbroadleaf-spruce forests i.e., of birch andaspenforestsof secondary for basin, i.e., upto 7.5th.t/km maximal inrelation river to theentire Oka depositionvaluesthereThe carbon willbe CONCLUSION C(Fa) oftheiraboriginal dynamicstypical 2 by2050. ofsubstitution The fact 2 (seeFig. 4, 2 . The resultant . The b b ). Birch- ). Birch a 003.08.2011 14:38:49 3 ). At . 0 8 . 2 2 0

1 1

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of HadCM3), doubling of atmospheric COof HadCM3),doublingatmospheric climate model(theprecursorUKMO-1987 shown that,according of to thescenario (CENTERE) and wasTER [Aber etal, 2001]. It by predictive biogeochemical modelsCEN content wereand thecarbon calculated productivity changes inthenetprimary coniferous, mixed, andotherUSAforests, other predictive regional estimations. For figures are quite comparableto thedataof content. of thebaselinecarbon These may befrom –(21 basin,averageriver weighted by area, main water catchment area ofthe Volga balanceoftheforeststhe carbon ofthe and thebiosphere asawhole.2050, By effects onthestate ofterrestrial ecosystems the rangeof–(1.5 results inrelative balancevaluesin carbon 8 3 .Fodpan91 4.866 44 .71.45 2.39 2.17 0.84 -4.45 -0.17 6.62 1.01 2.13 149.28 35.18 9219 2343 1.8 formations All groups offorest 8. Flood-plain –4.96 7. Paludal broadleaf substituting for 6.76 6. Secondary mixed light-coniferous and substituting for 84.68 5. Secondary 4770 mixed dark-coniferous and substituting for 4. Secondary 3. Broadleaf mixed broadleaf-pine 2. Pine and mixed broadleaf-spruce 1. Spruceand Forests (groups of plant formations) Table 4. Changes of the total carbon stocks (mln t) in plant formations of the Oka river basin by2050 basin river Oka the of formations (mln t) plant in stocks carbon total the of Table Changes 4. ò 34 468 4.4-0.24.22.97 43.82 -101.52 -3.15 145.34 -10.03 1476.83 8.86 83947 -51 25.98 4.69 40.97 –14.79 22.6 318.7 40.77 16670 -23.17 293.26 45.77 17770 481.76 26230 Area, 1.8) to +(7.8 ò 917.304 11 07 -0.99 -0.76 -1.18 0.42 76.83 3.28 4911 1.22 -1.8 3.02 37.14 2034 km 27)% to +(11 2 Total carbon (baseline) ò 12.5)%. ò 17)% cuuainemission accumulation 2

in theforest vegetation cover. lower content estimates changes ofcarbon to beextreme, itscalculationsgive much realistic and, thoughHadCM3isconsidered balancearemodeling ofthecarbon quite one cansee, theresults ofourpredictive model (see Table As 4,column "Deposit"). by2050according to HadCM3 the territory i.e. five timesgreater thanforecasted for this parameter willbeequalto 231mlnt, approximate river basin), area oftheOka 23.1 Gt.For onepercent ofthisarea (i.e., an a65-yearduring (1985-2050)isabout period that thetotal Cdepositionintheforests [Zalihanov etal, 2006],itmay beassumed sinkintheRussianforestsof thecarbon as awhole. rates Basedonthemodern ontheforestsome materials cover ofRussia Let uscompare theobtainedresults with Change ofthecarboncontent of thons millions Total  % 003.08.2011 14:38:49 3 . 0 8 . 2 0 1 1

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8 4 2. Celniker, 2. Yu.L. (2006). CO Aber J., NelsonR.P., St.,etal(2001).Forest1. Mcnulty Processes andGlobalEnvironmental Change: 17. Zalihanov,17. M.Ch.,Losev, Shelekhov, A.M.(2006). Naturalecosystems astheimportant K.S., 16. Sukachev, V.N. Studies. Selected V.1. (1972).Fundamentals ofForest Typology andBio- 15. Sochava, V.B. (1974). Geotopology asaDivisionofGeosystem Science// Topological as- 14. Shary, P.A. Sharaya L.S.,Mitusov, A.V. (2002).Fundamental quantitative methodsofland P.A.Shary 13. (2001).Analytical GISEco. Website: http//www.esti-map.ru. Rozenberg,12. inPhytocenology. G.S.(1984).Models Moscow. Nauka, 265p. (inRussian). 11. Pope, V.D., G Polynov, B.B.10. Studies. (1956).Selected USSRAcademy ofSciencesPress, Moscow. 751p. (in Kolomyts, E.G.,Rozenberg,9. G.S.(2009).Forecasted andcarbon changesofbiological cycle Kolomyts, E.G.(2008).Local mechanismsofglobal changesinnaturalecosystems. 8. Nauka, F.Klijn, 7. andUdodeHaes, H.(1994).Ahierarchical approach to ecosystems anditsimplica- Gorshkov, V.G.6. (1994).Physical andbiological basisoflife stability. Biota,Environment. Man, 5. M.A.(1964).GeochemicalFundamentalsGlazovskaya, of Typology of andMethodology Forman, 4. R.T.T. (1995).LandMosaics: The EcologyLandscapeandRegion. Uni- Cambridge 3. Climate Change1995(1996). The Science ofClimaticChange/J.T. Houghton, L.G.Meira REFERENCES Predicting andMultipleStressors theEffects //BioScience. ofIndividual V. 51.№9.P. 735–751. 634. (inRussian). resource //BulletinoftheRussian Academy ofmankind. ofSciences. V.76, №7.pp. 612– cenology. Leningrad. Nauka, 418p. (inRussian). pp. Novosibirsk. 3–86(inRussian). ofGeosystem Science.Nauka, pects analysis//Geoderma. V.107.surface №1–2.pp. 1–32. inHadleyCentreetrizations climate model–HadCM3//Climate Dynamics. V.16. pp. 123–146. Russian). cal Economic&Statistics. V. 15,NoF09,Pp. 1–19. balance intemperate forest ofEcologi- Journ. //Intern. ecosystems underglobalwarming Moscow. 427p. inEnglish,pp. (inRussian,Abstract 374–380). tion for ecological landclassification//LandscapeEcology. V.9. №2.pp. 89–104. Springer-Verlag, 340p. Berlin. Landscape Research. State Press, University Moscow Moscow. 230p. (inRussian). vers. Press, 315р. Cambridge, UK. Filho., andB.A. Callanderetal. (Eds). Univ.The Cambridge Press, 572p. Cambridge, UK. ForestIn Moscow. Science. Nauka, pp. Russian). 213–229(In allani M.L.,Rowntree P.R., andStratton, R.A.(2000). ofnewphysical param- The impact 2 exchange inforest biogeocoenoses//Concepts ofBiocenology 003.08.2011 14:38:50 3 . 0 8 . 2 0 1 1

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8 5 present andfuture. Sheistheauthorof36scientificpublications. ErlandG. Gennady S.Rosenberg Gennady LarisaS.Sharaya national and international technicalnational andinternational magazines. He publishedsixmonographs andalarge numberofpapersin formal analysisusingcomputerfield datafor technology. further andprocessingprediction. Hedeveloped methodsofcollecting systems, modeling ofgeo(eco-) andeco-geographical area ofinterest isproblems oflandscapeecology, empirical (Tolyatti), RussianAcademy ofSciences. Dr. Kolomyts’s main Landscape Ecology, ofEcologythe Institute Volga Basin ofSciencedegree.Doctor Dr. of isHeadofLaboratory Kolomyts 1976,hereceived State in1960.In his University Moscow scientific publications. and regional ecology. Heistheauthorofmore than700 (Togliatti). Dr. Rosenberg’s mainfieldofresearch istheoretical ofEcologythe Director oftheInstitute Volga Basin River oftheRussianAcademyMember ofSciences(2000)and ofSciencedegree.Doctor Dr. Rosenberg isCorresponding 1984,hereceived in1971.In his State (Ufa) University Bashkir technology and functional parameterstechnology ofecosystems andfunctional inthe mapping oflocalandregional ecosystems usingGIS environmental factors. Dr. Sharaya isdeveloping methodsfor oflandscapeformsquantitative as characteristics interestmethods. istheapplicationof Herparticular offorestvariability ecosystems using geomorphometric associated withinvestigations ofspatialandtemporal Russian Academy ofSciences. Herresearch interests are Ecology, ofEcologythe Institute Volga Basin(Tolyatti), ofLandscape 1985. SheisSeniorResearcher attheLaboratory State University,Moscow in1976andobtainedherPhD in

Kolomyts graduated from theFaculty ofBiology, graduated from theFaculty ofGeography, graduated from theFaculty ofBiology, 003.08.2011 14:38:50 3 . 0 8 . 2 0 1 1

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