Declining Ruff Philomachus pugnax populationsa response to global warming? CHRISTOPH ZOCKLER UNEP-WCMC, 219 HuntingdonRoad, CambridgeCB3 0DL, UK, e-maih [email protected] Z6ckler, C. Declining Ruff Philomachuspugnaxpopulations:a responseto global warming?Wader Study Group Bull. 97: 19-29. Ruff breedingpopulationshavedeclinedwidely andin all habitatsacrosstemperateEurasia.Of an estimated populationof 2.2-2.8 millionbirds,98% arenow confinedto habitatsin theArctictundra.Only 8,000-14,000 femalesstill breedin wet grassland habitatsin Europe.Althoughdrainageandagriculturalintensification have damagedor destroyedmanygrassland habitats,theydo notexplainwhy Ruffshavecontinuedto declineeven wherehabitatshavebeenimprovedand in placeswhereotherwet grasslandspeciesare stableor increasing. The hypothesisthattheproblemis limitedto oneor a few flywaysis notfully supported by the availabledata thoughthereis somesuggestion of a greaterdeclinein W Europe/WAfrica populations.For mostregions, systematicmonitoringdataare lacking.Neverthelessthe emergingpictureis that the populationhasshifted northwardsandeastwardsandhasretreatedfrom thewet grasslandhabitatsformerlyoccupiedalongthe southern edgesof its range.It is suggested that the causesare probablyof a globalnatureand may be linked with climate change.It is unclearwhetherthe total populationhas declinedor only shiftednorth and east.More co-ordinatedandsystematic monitoringof breedingandwinteringpopulationswill be necessary beforea full understanding of thesechangescan be reached. INTRODUCTION AND METHODS mainly in Central and EasternEurope. It shouldbe noted, however,that althoughinformationfrom Europeansitesis Despiterecentdeclines,the Ruff Philomachus pugnaxis still fairly complete,few dataare availablefor many regionsof widespreadand one of the commonestwader speciesof the tundraon eitherpopulationsize or trends.Nevertheless Eurasia.The breedingpopulationhasrecentlybeenestimated someobservations from theseregionsare discussed below. at 2.3-2.8 million birds (Z6ckler in prep.). This roughly In Europe,theRuff wasformerlymuchmorewidespread equatesto thelatestestimatefor thewinteringpopulationin thanit is today,rangingasfar southasAustria,Hungary,and Africa of more than 2 million birds (Trolliet & Girard 2001, Bavaria. Even in the 19th century,however, it was already van Gils & Wiersma 1996). However,recentobservationsof in seriousdecline.First it retreatedfrom its breedingsitesin decliningnumbers,particularlyin wet grassland habitats,are south-centralEurope.Then, by the 1940s,the oncecoherent alarming.Thispaperpresentsanoverviewof thecurrentsitu- distributionbecamepatchyandrestrictedto coastalareasand ationfor breedingRuff in Eurasiaas a baseline for future a few wet grasslandsitesinland (e.g. Glutz et al. 1975). The decline continuedthroughthe 1970s and 1980s with conpopulationmonitoring. Most datain this reportcomefrom two sources:recently centrationsin fewer andfewer sitesalongthe coastand even publishedregionaloverviewsfor countriesor regionswithin fewer sitesinland(Melter 1995, van Dijk 2000, HOtkeret al. countries(for sourcesseeTables 1 & 2), andthroughdirect 2001, Chylareckypers. comm.). Neverthelesssomecoastal sitescontinuedto supportsubstantialpopulations;particupersonalcommunicationwith local and countryexpertsin theInternational WaderStudyGroup.The figuresdiffer sub- larly newly reclaimed salt marshes and mudflats in the stantiallyfromthoseof Piersma(1986), andalsofrom those Waddensea.After a period of desalination,many of these of Heathet al. (2000). It is thereforeimportantto providean polderswere newly colonisedand for someyearsheld sigupdatedoverview of current populationstatus,including nificant populations,but later thesecrashedas the habitat breedingareasoutsideEurope,whereverpossible.The dis- changed.ExamplesincludetheLauwersmeer(van Dijk pers. cussion section draws international, national and local attencomm.), the Hauke Haien Koog andotherpoldersalongthe coastline(H6tker et al. 2001)(Figs 1-2). tion to a speciesthatis apparentlydecliningall overEurope, Schleswig-Holstein The recentdeclinein the specieswas first notedamong and possiblyalsoin many partsof Asia. It alsofocuseson possiblereasonsfor the decline,particularlythe likely imthe small populationon the southernedge of the breeding pact of climatechange. range.The decreasein Belgium,FranceandBritain led to its currentextinctionin thesecountriesduring the 1980s and 1990s.Also, furthereastalongthe southernboundaryof the RESULTS range in the stepperegion of Bashkortostan,Tomkovich In wet grasslandsitesandin naturalhabitatsacrossEurasia, (1992) describedthe Ruff asextinct.The declineover the last thepredominanttrendin the breedingruff populationin re- 20 yearsin theNetherlandsfrom 1,500to a maximumof 150 centyearshasbeenoneof decline(Tables1 & 2). Sitesfor- femalesandin Germanyfrom 450-600 to a maximumof 100 merly occupiedaroundthe peripheryof the rangeare now (Tables 1 & 2) is alarmingandhasresultedin a lot of attenvacatedwith the remaining populationnow concentrated tion in thesecountrieswhich are on the edgeof the species' 19 Buffetin 97 April 2002 20 Wader Study Group Bulletin Table 1. Population size(females)andtrendsof the breedingRuffpopulation incountriesandsub-national regionswithmainlywetgrassland habitats.Data for countriesand sub-nationalregionare in bold;data for Landerin Germanyand specificsites in The Netherlands, Polandand Estoniaare in regularfont.Trendsare basedon eitherdata in tableor listedreference(DEC = decline;INC = increase;STA = stable; EXT = extinct). WetJSaltgrassland ca. 1980 ca. 1990 ca. 2000 Trend Source 1-50 4 - 10 300-1,500 500-4,000 DEC DEC INC? ? Glushenkov et al. (1999) Grishanov (1998) Tomkovich 1992, V. Sotnikov - estimate Lebedeva - estimate, populations Russian oblasts ChuvashiaRepublic Kaliningradskaya Kirov region Kostromaregion DEC Lebedeva 0-1 3,500-4,000 100-200 10-100 5-50 DEC ? DEC DEC DEC Zubakin et al. (1998) Mischenko & Sukhanova(1998) Ivanchev, Kotyukov (1999) 10,0007 2,000 - 2,400 2,0007 2,200 85-180 STA? DEC > 150 50 50 200 350-400 150-300 50 20-30 50 100-200 50-100 11 DEC DEC DEC DEC DEC 540 10-15 500 200-300 DEC DEC DEC Nikiforov & Mongin (1998), Kozulin (pers.comm.) Veromann 1980, Heath et al. (2000), L6hmus et al. (2001) M•igi & Kaisel (1999) Viksne (1983, 1997), Priedniks et al. (1989) Tomkovich 1992, Heath et al. (2000) Chylarecki P. & Winiecki (2001) Witkowski, Dyrcz, Lontkowski& Stawarczyk(in prep.), Nawrocky (pers. comm.) Lebed & Knysh (1999), Gorban& Shidlovski(1999) S6renssen(1999), Widemo (pers. comm.) Thorup(pers.comm.),Frikke ( 1991), Koskimies(1992), Grell (1998) Z6ckler (1998a), H'filterleinet al. 2000, H6tker et a/.2001 Mordovia 1-10 Moscowregion Novgorodregion Ryazan'region Smolenskregion Vladimir region Belarus Estonia Mats alu Latvia Lithuania Poland Biebrza Ukraine South Sweden Denmark 150 500 Germany 500-600 Schleswig-Holstein -200 Mecldenburg-Vorpommern 50-80 Brandenburg ~ 10 Niedersachsen Bremen Nordrhein-Westfalen The Netherlands 50 -100 - estimate Lebedeva - estimate Lebedeva - estimate 300 200 30 75 50 10 1 DEC DEC DEC DEC ~200 10 45 5-10 15 0 DEC DEC H•ilterlein 0-2 0-2 0 EXT Nehls et al. 2001 1,250 -1,500 600 100-150 DEC H•ilterlein et al. 2000 Kube pers.comm.,H•ilterleinet al. 2000 Ryslavy 2001 et al. 2000 Seitz 2001 Osieck& Hustings1994, Crampet al. 1983 van Dijk pers. comm. Lauwersmeer UK France 350 3-21 Total 200 1-7 5-13 6 0 0 DEC EXT EXT 10,900-17,100 8,090-13,890 DEC van Dijk pers. comm. Sharrocket al. 1981, Ogilvie et al. 2001 Duboiset al. (1991), Trolliet (pers.comm.) dueto the species'highlynomadicbehaviour. distribution.In 1991, HOtkerestimatedthe Ruff population fluctuations of thetwelvecountriesthatat thetimeformedtheEuropean However, at several sites Ruffs have declined when other that some Unionat 1,900breedingfemales.Now onlya quarterof that wadershave performedwell and this suggests of ArcticRuff mayalsobe in decline.Declines number,475, remain.Countrypopulations listedasstableby populations Hftker (1991), suchas thosein Britain and France,in fact arereportedfor most(10/14) of thefew tundrasitesthatare becameextinctshortlyafterwards, withnobreedingrecords regularlymonitored(Table3). Thesedatagiveanindication of a widespread declinebutthisneedsto be verified.A resince1996 and 1997 respectively. Long-termmonitoring in Finlandhasalsoshowna substan- cent Russian initiative of the International Wader Study tialdeclineoverthepast20 years.However,withabout30,000 Grouphas addressedthe issueof trendinterpretationin Breedbreeding females,thatpopulation is notendangered (V'als'finen Arcticbreedingbirds,suchasRuff. A co-ordinated approach startedin et al. 1998).No significantchanges havebeennotedin other ingConditionSurveywith a circumpolar partsof Scandinavia or in mostof European Russia,butthese 1997 to monitorbreedingperformancein connectionwith (Solovievet al. populations havenotbeenintensively studied. Mostwetgrass- weatherconditionsandlemmingabundance land areas with time series information confirm the decreas1998)andit ishopedthatthiswill leadto a betterunderstandto tundra-breeding Ruffs. ing trendnoticedin the restof Europe(Tables1 & 2). The ing of whatis happening drastic decline in almost all non-tundra sites is in contrast to thestableor evenincreasing situationin somenorthernArctic DISCUSSION populations (Table3) andrequiresexplanation. The Russiantundrasupportsby far the majorityof the Reasons for the decline total Ruff population,about95%. Trend informationis declineof almostall wet grassland waders scarceand is oftendisguisedby naturalannualfluctuations The widespread resultingfrom coldweatherandpredationaswell asspatial in Europeandin manypartsin Asia urgentlyneedsto be Bulletin 97 April 2002 Zbckler: Declining Ruff populations 21 Table 2, Populationsize (females)and trends of the breedingRuff populationin countriesand sub-nationalregionswith mainly natural habitats.Trends are based on either data in table or listed reference (DEC = decline; INC = increase; STA = stable; EXT -- extinct). Tundra/peatland ca. 1980 ca. 1990 ca. 2000 Trend 15,000 80,000 196,000 15,000 57,000 39,000 15,000 50,0007 30,000 STA DEC? DEC Source habitats Norway Sweden Finland Russian Koskimies 1992, Kalas (1994) Koskimies(1992), Girard & Kirby (1997), Sfrenssen(1999) Girard & Kirby (1997), V•iis•inenet al. (1998) oblasts Murmanskregion 8,000-20,000 DEC Lebedeva,recalculatedfrom Bianki et al. (1993), Tatarinkova (1998) Mikhaleva, (1998), Lebedeva - estimate Karelia 800-5000 Leningradregion Tver region (peatlands) Novgorod region Vologda region Arkhangelsk (exc. Nenetsk AO) Nenetsky Autonomous Region Komi Republic Pskov region (peatland?) 500-5000 Lebedeva 500-5000 - estimate 1,000-10,000 1,500-10,000 STA Nikolaev (1998), Lebedeva - estimate Mischenko & Sukhanova(1998) Butiev, Shitikov, Lebedeva, (1998) Butiev, Shitikov (1998), Morozov (1998), Lebedeva- estimate 1,600 -5,000 STA Morozov (1998) (Vaigachonly) + Lebedeva- estimate 3500-4000 Lebedeva - estimate 1,000-10,000 3,000-4,000 Lebedeva - estimate Yamal INC Gydan Taimyr INC? Yakutia ? Ryabitsev& Alekseeva (1998) ? Chukotka STA Koryak Mountains ? Sachalin ? Soloviev et al. (200lb) Steppe habitats Hungary Kasakhstan 8-10 1-5 Bashkortostan Baikal area Mongolia 0-1 EXT? Heath et al. (2000) 0-1 EXT? Khrokov 0-1 EXT? Ilyichev & Fomin 1979, Tomkovich (pers.comm.) Sumja& Skryabin(1989) Skryabin & Toopitsyn (1998) 10-1007 107 1998 understood as a basisfor conservation action.Severalexplanationshave beenproposed.Often, however,the mostobvious site-specificfactor has been identified as the prime Melter 1995, van Dijk et al. 2000, Tomkovich& Lebedeva 1998, 1999).The breedingpopulationsthatremain,whether in naturalhabitatsor in wet grasslands, rely entirelyon wet cause and conservation action has been taken without conor very wet conditions.All nestsfoundthroughoutthe entire sideringwhat is known aboutwhole populations.Possible rangehavebeenin suchlocations,usuallyin the immediate reasons for the decline of Ruff are listed below and then dissurroundings of smallponds,puddlesor undrainedsurface cussedbriefly. wateronpermafrostor peatgrounds(Ztckler in press).High waterlevelsthereforeappearto be essential, thoughprobably not the only factor, for maintaining wet grassland bird comLocal, site-related factors: munities, such as Ruff, Black-tailed Godwit Limosa limosa [] [] [] [] Drainage Agriculturalintensification Abandonment of land-usepracticesandsuccession Predation(by foxes,weasels,crowsandbirdsof prey) andotherwet grasslandbirdsat healthypopulationlevels. By itself, however,drainageno longerexplainsthe continueddeclineof wet grasslandwadersfor they have also declinedin well-managed,wet and deliberatelyfloodedreserves, such as the WQmmewiesen reserve near Bremen, Global factors: Germany(Fig. 3), theAlte Sorgeschleife reservein Schleswig- [] Changesalongflyways [] Globalchange,includingclimatechange Holstein, Germany, and the Biebrza marshes in Poland. Ruffshaveevendeclinedat extremelywet coastalsites,such asthe Matsa!ureservein Estonia(Fig. 4) wheredunlinhave beenincreasing(M•igi & Kaisel 1999).This demonstrates the Drainage The declinein wet grasslandbirds, particularlyRuff, has mostfrequentlybeenconnectedwith drainageandimprovement of land for agriculture.Withoutdoubtthiswasresponsiblefor thelarge-scaledeclineoverthepastcenturyin many areasof Western and Central Europe, and also in parts of Eastern Europe (e.g. Glutz et al. 1975, Beintema 1986, need to look for other reasons. The decline of Ruff in Finn- ish peatlandreservesis alsonot necessarilyconnectedwith drainageor wet conditions.Certainlythe lossof peatlandin the 1950s and 1960s (J•irvinen& Sammalisto1976) caused a hugelossof habitatfor Ruff and otherpeatlandspecies. This cannot,however,explainthecontinuation of thedecline in the 1980sand 1990s,afterlarge-scalepeatdestructionhad ceased. Buffetin 97 April 2002 22 Wader Study Group Bulletin 1000 ........................................................................................ 100 10 1968 1973 1978 1983 1988 1993 1998 Fig. 1. Populationtrendof Ruffsat the Lauwersmeerpoldernaturereserve,the Netherlands(basedon van Dijk et al. 2000 and pers. comm.). Agricultural intensification Poland(Table 1, Figs 1 & 2)). Secondlyand more significantly,the declineof Ruff in EasternEuropestartedbefore The useof land for agriculturehasintensifiedfor centuries thelarge-scaleintensification of agriculturebegan,asshown andat an increasing rateduringthe last50 years,especially for examplein EstoniaandPoland(Table 1). For thesereain Western and Central Europe, with a generally adverse sonsit is importantto look for other explanationsfor the effect on birdlife (Donald et al. 2001). Grasslandbirdshave declinewithoutdenyingthesevereimpactthatintensiveland declinedas fertiliseruse,grazingandthe frequencyof me- usehashad on wet grasslandbird communities. chanical treatment have increased. Two observations,how- ever, indicatethat additionalexplanationsare needed. First,thereverseprocess,theextensification andde-nitri- Abandonment of land-use practices and succession ficationby zero-application of fertilisers,hasnotalwaysled WhenRuff breedonfarmland,outsideprimarynaturalhabitat, moderategrazingor mowingis requiredfor it to remain to a re-colonisationby grasslandbird communities(e.g. Nehls 2001). Moreover, the few sitesthat never faced the intensificationprocessand remainedmostlyuntouchedexperienceda declinein wet grassland birds,particularlyRuff (see examples in North Germany, the Netherlands and suitable.If thisstops,it is destroyed asbreedinghabitatby successional changes thatleadto a higheranddenservegetation structure.Severaltimesthe declineor disappearance of Ruff and other waders has been attributed to the abandonment ;••i; tkr•.•l• IredrilKøl•og ! •• _•!.•11c••• wHa•• rK;•irv o rla ! •'Katlnger• •• Fig. 2. Populationchanges of Ruff at selected breedingsites in recentlyenclosedpolders(K6ge) along the North Sea coast of Germany during the past 20-30 years (based on H6tker et al. 2001). Bulletin 97 April 2002 Z0ckler: DecliningRuff populations 23 2O • 12 • 8 4 Fig. 3. Populationchangesof Ruffat an inlandsite:the naturereserveat WOmmewiesen,on a floodplainnear Bremen,Germany(based on Eikhorst & Mauruschat 2000 and the author's own observations). of land-usepractices.Soikkeli & Salo (1979), for example, demonstrated that a declinein populationsalongthe south coastof Finlandwasdueto the abandonment of lessproductive farmland. Moreover, the decline in the Netherlands has beenattributedlargelyto the succession of vegetation(van Eerdenet al. 1979,Altenburget al. 1985,vanDijk et al. 1999). of the decline. Moreover, the decline has also been observed on Dutch islandswheremostof thesepredatorsare absent (van Dijk pers.comm.).The well-beingof somespeciesat particularsitesandthedeclineof others,aswith theDunlins andRuffs at Matsalu (Fig. 4), suggests that predationis not an importantreasonfor the large-scaledeclinein wet grassland birds. Predation Global Predatorsplay an importantrole in wet grassland bird communities(Lugert 1994, Schoppenhorst 1996, Hase & Ryslavy 1998 & Seitz 2001). Most commonlyrecordedin Europe are Red Fox Vulpesvulpes,Weasel Mustela nivalis, Mink Lutreola lutreola and domesticdog. Among avian predatorsare crows Corvus spp., Marsh Harriers Circus aeruginosus andPeregrines Falco peregrinus.However,it seemsunlikelythatconditionsfor predatorshavesimultaneouslychangedin their favourall over the Ruff's breeding range,to the extentthatthattheyhavebeenthemajorcause factors Global factorsare thosethat impacton a speciesat the global or flyway level andflyway factorsarethosethat impact alongan entiremigrationroute.Global factorsincludeclimatechangeandpollutionand alsothe assemblage of multiple humanactivitiesthatimpacttheEarthat a globalscale. Severalrecentstudieshaveindicatedtheimportanceof such mattersin explainingdistributionchangesand population trendsin a varietyof taxa (e.g.butterflies(Parmesan1996)). Thesefactorsmay be workingtogether.For example,wader 1000 100 10 1962 i i 1972 1982 1992 2002 Fig. 4. Populationtrend of Ruff in the Matsalu nature reserve, Estonia (based on E. M•.gi pers. comm.). Bulletin 97 April 2002 24 Wader Study Group Bulletin Flyway Aewa boundary Countries/Russian oblast I1[111I! [__.____j Extinct • Stable I Declining No data Fig. 5. Trends in the Ruff populationon the breedingand winteringgroundsin the contextof three recognisedflywaysI-III; based on Boere (1991 ), Syroechkovski& Rogacheva (1996), Rogacheva (1992), Underhillet al. (1999) (sourcesof trend data: Tables 1 & 2, Triplet & Yesou (1998) and unpublisheddata from the IWC for Africa, courtesy Wetlands International). habitatsare likely to be changingnot only becausecarbon The flyway hypothesis dioxide and other greenhousegasesare leading to global climatechange,but alsobecauseof the impactof increasing This assumes thatthedeclineis particularlyprominentin one amounts of air- and waterborne nutrients. flyway populationcomparedwith othershaving different that the causelies somewhere Two broadhypotheses areproposed astheexplanation for migratingroutes,suggesting thewidespread declineof theRuff andmanyotherwet grass- alongthat particularflyway. There appearsto be someeviland waders: dence for different flyway trends in the case of the Ruff (Fig. 5), but winter count data are too sparsefor any firm 1) The flyway hypothesis:Somethinghashappenedto the conclusion.In particular,datafor Africa are insufficientto birdsalongtheflywaysof thepopulationin theirwinter- determinelong-termtrends(Trolllet & Girard 2001). However, the fact that thereis alsoa decliningtrendin the rather ing or staginggrounds 2) The globalchangehypothesis: The gradualalterationof separateSouthAsian populationsuggestsa more general wet grasslandhabitats,due to direct and indirecthuman declineoverall flyways.Similarlythe declineof mostwader activities, has caused the decline. populationsin theAmericas(Morrisonet al. 2001) notonly Bulletin 97 April 2002 ZOckler:Declining Ruff populations 25 indicatesthat the problemis moreglobalthanlocal, but also ing species,suchas Curlew SandpiperCalidrisferruginea, suggests that it is largerthansomethingthathasoccurredin Temminck's Stint C. temminickii, Little Ringed Plover just oneor two flyways.However,a varietyof localchanges Charadrius dubius, Lesser Sand Plover Ch. mongolus, havedevelopeda globalcharacter,which include: PacificGoldenPloverPluvialisfulva,GreyPloverPluvialis squatarola,Black-tailedGodwit, Bar-tailedGodwit Limosa [] Desertification,droughtsandthe lossof wetlands, lapponica,Whimbrel Numeniusphaeopus,EurasianCurlew [] Eutrophicationof stagingandwinteringwetlands, N. arquata,CommonRedshank,CommonSandpiperActitis [] Generalchangesin land-useandapplicationof chemicals hypoleucos,Terek SandpiperXenus cinereus and Blackand nutrientson winteringgrounds,leadingto changes winged Stilt Himantopushimantopus(Lopez & Mundkur in vegetationandchangesin growthrate andbiomass. 1997). The Ruff is declining, whereas Common Snipe, PintailSnipeGallinagostenuraandWoodSandpiperTringa The impactof habitatlossanddesertificationon winter- glareola, recognisedwidely over Siberia as frequent coing wadershas been particularlynoticedin West Africa breederswith Ruff (Z6ckler in prep.),showa stableor only whereRuff andBlack-tailedGodwit arehighly nomadicand slightlydecliningtrend.Only Little StintC. minuta,the speoften changetheir wintering site from year to year (Altencieswith the secondlargestwaderpopulationin the region, burg& van der Kamp 1986, Yesou& Triplet 1996, Triplet alsoshowsa cleardecliningtrendsimilarto the Ruff. There & Yesou 1998). Thesetwo specieshave becomemore and is little information available either on the use of fertilisers more dependanton inundatedrice fields after the loss of or on land-usechangesor the impactof huntingin the winmany naturalwetlands(Altenburg& van der Kamp 1986). tering areas. Dependinglargely on rainfall, rice productionvariesfrom year to year, so bird countsfrom one areamight not reveal The global change hypothesis the full picture.Only aerialsurveyscoveringseveralregions reveal a picture that is closeto reality (Trolliet & Girard Global change comprises all natural and human-driven 2001). Even so, countsfrom differentregionsoften derive changeson our planet, including climate changeby the from differentyears,sotrendscannotreadilybe identified. accumulationof greenhousegases,and also eutrophication In someregions,however,numbersseemedto havedeclined throughfertilisersand nutrientreleasethroughfossilburnconsiderably,particularlyin Senegaland NorthernNigeria ing. The impactof thesechangesis complexandfew factors (Glutz et al. 1975, OAG MQnster 1991, Triplett & Yesou are traceable and identifiable as responsiblefor what we 1998, Trolliet & Girard 2001). observein wet grasslandandits birds.The main featuresof The nominatesubspeciesof the Black-tailed Godwit in globalchangerelevantto wet grasslandandtundrabirdscan Europe,L. I. limosa,moreor lesssharesthe sameflyway and be summarised as: winteringgroundsasthe Ruff (Glutz et al. 1975, Crampet al. 1983) andmostlikely facesthe samethreats.Both are de- [] Changesin climate, clining stronglyand persistentlyall over Europe, whereas ß generally more rain in temperate regions, warmer other species with different flyways, such as Common annualmean temperature RedshankTringa totanus and Common Snipe Gallinago ß drier andhotterclimatein winteringareaswith losses of wetlands gallinago,haveonly declinedin thoseareaswheretherehave beenhabitatlosses.Similarly,theIcelandicsubspecies of the [] Extendedgrowingseasonandearly startof vegetation Black-tailed Godwit, L. I. islandica, which winters in the growth BritishIslesandsouthwestEurope,hasan increasingpopu- [] Increase of biomass lation (Gill et al. 2001). [] Changeof vegetationheightanddensity When each Ruff flyway is consideredseparately,it ap- [] Impoverishment of vegetationin structureanddiversity pearsthatwesternpopulationshavedeclinedmostandthose [] Increaseof naturalforestationin themainbreedingarea breedingand wintering further easthave remainedmore in the tundraregion. stable.Thereforethe declinein Europe,includingthe western partsof Russia,is matchedby similarfalls in the main Climatechangeis a complexprocessthatmay affectwet winteringgroundsof the samepopulationsin West Africa grasslandbirds, and Ruff in particular, in three different (Fig. 5). In Siberia, Tomkovich (1992) consideredthat the areas.First,theymaybe affectedin theirprimeArcticbreedRuff had extendedits rangeinto Chukotka,Kamtchatkaand ing range.Habitat conditionswill changedue to warming. otherpartsof CentralYakutia.However,it is quitepossible Most of theopentundrahabitatscurrentlyoccupiedby Ruff, that the specieshad previouslybeen overlookedin those for example,will changeinto foresttundraandforesttundra areas.Indeed, in Chukotka, ancientRuff skeletonshave been may becomedenselyforested, a processthat has already foundin Chukchisettlementsindicatingthe species'pres- started, as observed in Alaska (Scott et al. 1996, see also encetherein historicaltimes (P. Tomkovichpers.comm.). Z6ckler & Lysenko 2000). Stableor increasingwinteringpopulationsin East Africa Second,climatechangealsoaffectsthetemperateregion, may reflect more healthy populationsin Eastern Russia thoughnot so fundamentallyasin the Arctic, andits effects (Fig. 5). The Central Asian flyway may be similar, though arealreadyevidentin the growingseasonof Europeantrees therearenotenoughdatafromthebreedinggroundsto prove (Menzel & Fabian 1999). The combinationof global warmthis conclusively.Broadly thereforeit seemsthat there is ing and eutrophicationfrom both airborne nutrients and someevidenceto supportthe flyway hypothesis,but there fertiliserrun-offhasbeenidentifiedasresponsible for boostaretoo manygapsin the datafor confidence. ing vegetationgrowthin mostpartsof Europe.It is estimated In Southern Asia, the Ruff is the seventhmost numerous that about50 kg of nitrogenper hectareis appliedannually wintering wader species.The highest count in 1995 was by rainfall alone.In additionnutrientsarrive via floodwater 11,400 with a decliningtrend comparedto severalincreas- on wet grassland.In combinationwith globalwarming,this Bulletin 97 April 2002 26 Wader Study Group Bulletin Table 3. Populationtrends of Ruff at various breedinglocationsin the Arctic(DEC = decliningor low numberscomparedto year before, INC= increasing,S = southern tundra, N = Northerntundra). Location Trend Years, comments Fenno-ScandinaviaS DEC Last 20 years Northern Murmansk coast S DEC 1989, 1990, 1994, 1998, 1999, 2000 MalazemelskayaTundra $ DEC 1998 Kanin Peninsula S DEC 1996 Russki Zavarot, Pechora S DEC 1997, 1998, 1999 Russki Zavarot, Pechora S INC 1994 Central Yamal N DEC 1993 INC 1982-1991 Southern Yamal $ DEC 1993 Southern Yamal DEC 1998 Lower Ob valley S EasternTaimyr N DEC INC 1999 1995-2001 Yana Delta N INC 1998 Western Taimyr, LowerKolymaN DEC 1988 compared to yearbefore Southern Yamal S S Sources V•iis•inen et al. (1998) Tatarinkova(1998) and in Tomkovich & Lebedeva(1996), Soloviev & Tomkovich (1999, 2000, 2001) Mineev cit. in Soloviev & Tomkovich (1999) Filchagov in Tomkovich & Zharikov 1997 Shchadlikov& Belousovain Tomkovich & Zharikov (1998), Soloviev & Tomkovich (1999, 2000) Shchadlikov& Belousovain Tomkovich & Lebedeva (1996), Shtro in Tomkovich (1998b) Ryabitsev& Alekseeva(1998) Paskhalnycit. in Tomkovich (1998b) Morozov in Soloviev & Tomkovich (1999) Paskhalnyin Soloviev& Tomkovich(2000) Soloviev et al. (2001) Keremyasov& Turakhovin Soloviev& Tomkovich(1999) Tomkovich (1998a) hasresultedin the growing seasonstartingfour weeksear- befound inthetundra andpeat•bogs where future changes in lier, and in an increaseof biomasswith a more lush and dense vegetationareunlikelyto affectRuff. In fact,there,changes aremorelikely to be beneficialto Ruff thanmostotherspecies (ZiSckler& Lysenko2000) becauseRuff canbe found at moderatelyhighdensitiesin foresttundra.It hasalsobeen observedthat Ruff will toleratea certainamountof change in the vegetationstructure,includingbushesand scattered trees,as long as the basevegetationlayer doesnot get too denseandthe waterconditionsremainunchanged andsuitableto provideenoughfoodresources (Z/3cklerin prep.). IndicationsthatRuff havedeclinedin southern regionsof vegetation comparedto the 1950s in northwestEurope (Beintemaet al. 1995). Of all thewet grassland birdsof central Europe,theRuff hasbeenrecognised asthe speciesthat is the mostsensitiveto thesechangesandoneof the first to reactto drainageand eutrophication(Beintema& MQskens 1987). The Ruff, however, is not the only one affected. Otherslike CommonSnipe,CommonRedshankandBlacktailed Godwit are following. Vegetationsamplestakenat onesitewithoutanyfertiliser applicationin the WQmmewiesenreserve,Germany,betweentheearly1980sand1995showedanincrease in biomass after a shortperiod of decrease(Warnken 1994, Rosenthal in prep). Anotherindicationof earlier growth with more biomassis the dateon whichthe first cutof harvestedgrass the tundra and increased in the north and east (Table 3, Tomkovich& Sorokin 1983, Tulp et al. 1997) supportthe ideathatthedistributionhasshiftedbecause of thewarming climate.Similarly,in Europethe declinestartedfirst in the south,in AustriaandBavaria,earlyin the20thcentury,then is taken. In all areas without conservation restrictions, this in FranceandBelgiumin the 1980sandin the UK, southern hasbeentakenearlierandearliereveryyear andit hasnow RussiaandKaskhstan in the 1990s(Glutzetal. 1975,Ogilvie movedto earlyMay (Nehlset al. 2001). In 2000, the first cut et al. 2001, Morozov pers.comm.).Recently,it hascontinuedin westEuropeandin north-centralEurope,asfar north in the vicinity of WQmmewiesenwas as early as 1st May (Warnken,pers.comm.)comparedwith the endof May in asArcticFinland.Today,thedeclinemayevenextendto the the early 1980s. Drainagetendsto enhancebiomasseven southernpartsof the Russiantundra. furtherbecausedrainedmeadowswarmupfasterandwarmth The emergingpictureis thereforethat the Ruff is being forcedto retreatto its corenorthernhabitatsthroughglobal increasesgrowth. Third, thewarmingclimatewill alsochangeconditionsin climatechangeandthe effectof thaton thequalityof its wet the winteringareas.Wetlandswill decreaseasa directresult grasslandhabitats.The declineis steepestin the southof the of the increasingdrynessof the climate.Moreover,second- breedingrange, particularly in Finland (V•iis•inenet al. ary processes,suchas the increasingdemandfor water to 1998), but alsoover the last ten yearsin Denmark(50%), irrigatepreviouslyrain-fedcrops,will reinforcethis effect. Germany(75%) and The Netherlands(80%) (Table 1). As Accordingto a studyby Universityof Wisconsin-Madison thedeclinehasoccurredall overEurope,includingEuropean researchers usingsatelliteimages,Lake Chadis now only a Russiaandin all habitats,includingsometundraareas(but twentiethof the sizeit was 35 yearsago.The regionhassuf- not stronglyif at all in Siberia),it canbe concludedthatthe fered from an increasinglydry climate, with a significant impactof globalchangeon Ruff breedingin wet grassland declinein rainfallsincetheearly 1960s(Coe& Foley2001). habitatsis greaterthanon thosebreedingin tundrahabitats. In addition, the lake has become the source of water for This patternwould seemto be consistentwith changes massiveirrigation projects. In the winter of 1999/2000, apparentfrom the fossilrecordwhich showssequences of 339,000 Ruff were countedin the Lake Chad area (Trolliet colonisation, disappearance andre-colonisation in cooling & Girard 2001). However, becauseof changesin survey and warmingperiodsbetweenthe ice ages.Evidencefrom methodsandthe areacovered,it is notpossibleto determine Pleistocene fossilRuffsin Denmark,Finland,Azerbaijanand any populationtrend. Hungary(van Rhijn 1991) might indicatebreedingfurther In the Arctic, plenty of suitablebreedinghabitatcan still southduringcolderperiods,but it is equallypossiblethatthe Bulletin 97 April 2002 ZOckler: Declining Ruff populations fossilsare from migrants.The tundrabiome during recent glaciationswasmuchfurthersouththanit is todayandcould well haveenabledRuff to breedin thoseareas.Undoubtedly Ruffs musthave accompaniedthe cyclic shift of the tundra and its adjacentbiomesthroughthe ice-agesand it is quite possiblethat this processcontinuesto this day with the currentretreat northwardsas the globe warms. This coincideswith therecentnorthernexpansionof other wet grasslandwaders,suchas Common Snipe in the Bolshemelzkayatundra(Morozov 1998), Black-tailedGodwit andNorthernLapwing Vanellusvanellusin NorthernRussia concomitantwith a northwardexpansionof agriculture includingsownmeadows(Morozov 1987, Lebedeva1998). Several other bird specieshave recentlyrecordedin more northernlocationsin the Arctic (Z6ckler et al. 1997) and this confirmsa generaltrendof speciesshiftingtheirdistributions in responseto changingclimate. There seemto be a variety of factorsresponsiblefor the increaseof vegetationbiomass.The mostobviousare linked with the generalintensificationof landuse.But evenin those naturereserveswhere no fertilisershave been applied and water quality protected,the Ruff and severalother species havedeclined.In theseplaces,subtlechangesassociated with global climate change, particularly earlier and warmer springs,have boostedvegetationbiomass,often aided by drainageas well as by airborneand waterbornenutrients. 27 from the background"noise" of natural variation. The observedtrendsalong the flyways (Fig. 5) are not basedon systematicmonitoring.This needsto be remediedas a matter of urgency.Futuremonitoringshouldtake accountof all thesefactorsand be designedto coverall flyways from the breeding groundsto the wintering groundsas well as all stopoversitesin between. The plightof the Ruff in Europeis alarming.Althoughit is only understandable in the contextof changesthat are of global application,otherfactorshave undoubtedlyexacerbated the situation. These include large-scale land-use changes, drainage and the general increase of nutrients. Theseaddto the eutrophication of the landscapeandglobal warmingwith a synergeticeffect on the species.Similarly, althoughthe impactof predatorsis usuallyquite local, this may be quite devastatingwhere it causesthe final extinction of a vulnerablepopulationat a degradedsite. This studydemonstrates thatthe declineof the Ruff needs to be considered in the contextof entireflywaysandthe global situation. All conservation action should take this into account.However, despitethe well-documenteddeclineof the breedingpopulationacrosstemperateEurasia,evennow we cannotbe sureexactlywhathashappened.Has the populationmerelyshiftedor hasit declinedglobally?To answer thisquestion,moresystematicmonitoringin the wintering groundsof Africa andsouthernAsia is vital. Also, although we may have confidencein the connectionbetweenthe decline of the Ruff and globalchange,we haveyet to cometo CONCLUSIONS gripswith a real understanding of the proximatefactorsinThe Ruff is still a commonspeciesin the Eurasiantundranor volved. For example, vegetationbiomassin the breeding is it globallythreatened.The recentdeclinein wet grassland habitatsmayhaveincreased, buthowdoesthisimpactonthe habitats,however,hasbeensowidespreadthat it shouldbe ability of Ruffs to surviveand/orreproduce? consideredin a global context. Most other wet grassland wadershave declining populations.Sometimesthis is the ACKNOWLEDG EM ENTS resultof local changesin land-useor the impactof predators, but for certain species,especially Ruff and Black-tailed Among thosewho have supportedthis study,I would like to Godwit, it seems that the causes must be of a nature that is thank many members of the International Wader Study global.Unlike speciesthat are largely confinedto wet grass- Group.Without their enthusiasticcontributionsthe extensive land,Ruff differ in that they maintaina strongpopulationin coveragethat thisprojecthasachievedwould not havebeen tundra habitats. Available data show a shift in distribution, possible. I wouldalsolike to thankGillianBunting,Hermann but it is not possibleto determineto what extent lossesin H6tker and Hartmut Andretzke for comments on an earlier southernpartsof the rangeare compensated by gainsin the draft, CorinnaRaviliousand Igor Lysenkofor GIS support north and east. Similar changesmay apply in the Icelandic and particularlyPeter Boye for many encouragingdiscusandNorwegianpopulationsof the Black-tailedGodwit. It is sionsand support.The GermanFederalAgency for Nature thereforeimportantthat thesebe monitoredclosely(which Conservationsupportedpart of the work financially. is currentlynot being done). The Ruff appearsto be very sensitiveto changingclimate REFERENCES as well as to changingwater tables and faster vegetation growth.Thereforeit might be an ideal indicator-species for Altenburg, W., N. Beemster, K. Van Dijk, P. Esselink, D. Prop & H. monitoringglobal change.In wet grasslandareas,however, Visser 1985. De ontwikkeling van de broedvogelbevolkingvan het Lauwersmeer in 1978-1983. Limosa 58: 149-161. it may be already too scarce to provide adequate data. 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