Accelerating Climate Change Impacts on Alpine Glacier Forefield Ecosystems in the European

Alps
Author(s): Nicoletta Cannone, Guglielmina Diolaiuti, Mauro Guglielmin, Claudio Smiraglia
Source: Ecological Applications, Vol. 18, No. 3 (Apr., 2008), pp. 637-648
Published by: Ecological Society of America
Stable URL: http://www.jstor.org/stable/40062174 .
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 18(3), 2008, pp. 637-648
EcologicalApplications,
© 2008 by the Ecological Societyof America

ACCELERATING CLIMATE CHANGE IMPACTS ON ALPINE GLACIER

FOREFIELD ECOSYSTEMS IN THE EUROPEAN ALPS
Nicoletta Cannone,1'4 Guglielmina Diolaiuti,2 Mauro Guglielmin,3 and Claudio Smiraglia2
1
2 Universityof Ferrara,Department of Biologyand Evolution,Ferrara,Italy
1 Universityof Milan, "A. Desio" Department of EarthSciences,Milan,Italy
of Insubria,Department
University of Structuraland FunctionalBiology,Varese,Italy

Abstract. In theEuropeanAlps theincreasein air temperature was morethantwicethe

increasein global meantemperature overthelast 50 years.The abiotic(glacial) and thebiotic
components(plantsand vegetation)of themountainenvironment are showingampleevidence
of climatechangeimpacts.In theAlps mostsmallglaciers(80% of totalglacialcoverageand
an importantcontribution to waterresources)could disappearin the nextdecades.
Recentlyclimatechange was demonstratedto affecthigherlevelsof ecological systems,
withvegetationexhibitingsurfacearea changes,indicatingthatalpine and nival vegetation
may be able to respondin a fastand flexibleway in responseto 1-2°C warming.
We analyzed the glacier evolution(terminusfluctuations, mass balances, surfacearea
variations),local climate,and vegetationsuccessionon the forefield of SforzellinaGlacier
(Upper Valtellina,centralItalianAlps) overthepast threedecades.We aimedto quantifythe
impacts of climate change on coupled biotic and abiotic componentsof high alpine
ecosystems,to verifyif an accelerationwas occurringon themduringthe last decade (i.e.,
1996-2006)and to assess whethernew specificstrategies wereadoptedforplantcolonization
and development.
All the glaciologicaldata indicatethata glacial retreatand shrinkageoccurredand was
muchstronger after2002 thanduringthelast 35 years.Vegetationstartedto colonizesurfaces
deglaciatedforonlyone year,witha rateat leastfourtimesgreaterthanthatreportedin the
forthe establishment
literature of scatteredindividualsand about two timesgreaterforthe
well-establisheddiscontinuousearly-successionalcommunity.The colonization strategy
changed:thefirstcolonizersare early-successional, screeslopes,and perennialclonal species
withhighphenotypicplasticityratherthanpioneerand snowbedspecies.
This impressiveaccelerationcoincidedwithonly slightlocal summerwanning(approx-
imately+0.5°C) and a poorly documentedlocal decrease in the snow cover depth and
duration.Are we facingacceleratedecologicalresponsesto climaticchangesand/ordid we go
beyonda thresholdoverwhichmajorecosystemchangesmayoccurin responseto evenminor
climaticvariations?
Key words: alpine glaciers; climate warming;clonal plants; colonizationstrategies;Italian Alps;
mountainecosystems;
SforzellinaGlacier;vegetation.

Introduction addressed.Likewise,as manyglaciersare foundat high

The increasingrecessionof thecryosphere in theAlps
is probablyrelatedto importantchanges occurringin
mid-tropospheric conditions,such as the widelyrecog-
nized rapid increase in temperatureduring recent
decades(IPCC 2001). The worldwideretreatof glaciers,
from alpine areas (Haeberli and Beniston 1998) to
Antarctica(Rott et al. 1996, Cook et al. 2005), in the
courseof the last fewdecades,is frequently mentioned
as a clear and unambiguoussign of global warming
(Oerlemans2005). Comparedwithotherclimateindica-
torslike treerings,glacial systemsreactin a relatively
simplewayto climatechange:thetransfer functiondoes
not change in time and geometriceffectscan be
Manuscriptreceived19 July2007; accepted 16 November
2007. Corresponding Editor:H. P. Schmid.
4 E-mail: nicoletta.cannone@unife.it
637
elevations,a climatesignal reflectedin glacierfluctua-
tionscan be studiedas a functionof height(Oerlemans
2005).
In the Alps atmosphericwarmingwas found to
increasemorethandouble overthesame period(Bohm
et al. 2001), witha significant summerwarming,which
was particularly severesince 1970(Castyet al. 2005). As
a resultof this rapid climate evolution,many small
glaciers(i.e., glacierswithsurfacearea <1 km2)located
at mid-elevationcould disappear in the next few
decades. These small glaciersare commonin the Alps,
wheretheyrepresent80% of total glacial coverageand
make an importantcontributionto water resources
(Oerlemansand Fortuin1992).
The rapid "disintegration"of alpine glaciers has
already been pointed out in an analysis of the Swiss
GlacierInventory 2000 by Paul et al. (2004). Amongthe
638 NICOLETTA CANNONE ET AL. EcologicalApplications
Vol. 18,No. 3

others,smallerglacierswerefoundto showbotha wider provideenvironments subjectonlyto naturaldynamics,

scatterof variationwithrespectto the largerones, and withoutdirectanthropogenic impacts,whereitis possible
to contribute morethanproportionally to thearea they to compare the responses of related environmental
represent. In fact,44% of thearea loss between1973and components (glaciersand vegetation) to thesameclimatic
1998/1999refersto glaciersof lengthsof <1 km and inputsand to quantifytheirvariationsin responseto
coveringonly 18% of thetotal area in 1973 (Paul et al. climatechanges.
2004). Thus smallglaciersare showinghighersensitivity The aim ofour researchwas to quantifythebioticand
than largerones due to theirveryfast reactiontime abiotic impactsof climatechange over the past three
(sensu Haeberli and Hoelzle 1995) and are, hence, decades in an alpine glacier area without direct
suitablesitesforassessmentand monitoring of climate anthropogenicimpacts,to verifyif an accelerationhas
changeimpacts(Dyurgerovand Meier2000). Moreover, been occurringon them duringthe last decade (i.e.,
at highelevationstemperature, moisture,and pressure 1996-2006),and to assesswhethernewspecificstrategies
trendsand anomalies are clearerthan at lower levels, wereadopted forplantcolonizationand development.
where the large-scale climate signals tend to be For thispurposewe analyzedthepatternsand rateof
dampened(Beniston2000, 20Q3). Also soil characteris- glacier evolution (abiotic component)and vegetation
ticsof the glacierforefield wereused as climatechange succession(bioticcomponent)on theforefield of Sforzel-
indicators(i.e., Egli et al. 2006). lina Glacier(UpperValtellina,centralItalianAlps).
Not only the abiotic (glacial) but also the biotic
Study Area
components(plants and vegetation)of the mountain
environmentare showing ample evidence of climate The SforzellinaGlacier is a southwest-facing cirque
changeimpacts(e.g., Grabherret al. 1994,Benistonet glacierextendingfrom2850 to 3100 m above sea level
al. 1997,Beniston2003,Waltheret al. 2005,Cannoneet (a.s.l.; 46°20/55// N, 10°30'50" E), located in Valfurva
al. 2007). Accordingto quantitativeestimatesof the Valley (Upper Valtellina),in the centralItalian Alps.
biologicalimpactsof theclimatechange"fingerprint," a The glacial forefield extendsbetween2850 and 2700 m
greateramplitude is expected at high latitudesand a.s.l., and it is characterizedby surfaceswithdifferent
altitudes(Root et al. 2003). In theAlps,theshiftsin the ages since deglaciation,rangingfromone year to >80
altitudinalrange marginsof plant species and biocli- years,whileundisturbedby periglacialfeaturessuch as
maticzones in thelast 50 years,withupwardmigration sorted or unsortedpolygons. Only a few not well-
of alpineand nival-plantspeciesat a rateof 8-10 m per developedterracettes and solifluctionlobes occuron the
decade (Grabherret al. 1994,Waltheret al. 2005), and northeast-facing slope of themoraineridgedepositedat
changesin community composition(Keller et al. 2000) the beginningof the 20th century(1920-1925) on the
providethefirstevidenceof the sensitivity of mountain southernborderof theglacier.
habitatsto climaticchange. Sforzellina Glacier representsa unique case of
Surfacearea changesof thevegetation in a highalpine glaciologicalstudyin Italy because it has one of the
site of the European Alps between 1953 and 2003 older and more continuousrecordsof terminusfluctu-
demonstrated thatclimatechangeis able to affecthigher ations(1925 to today)and massbalance(1987 to today).
levelsofecologicalsystems and thateven 1-2°C warming It is also one of the few glaciers of which thereis
of air temperature may produce significant changesin relatively good knowledgeofitsthickness and geometry.
vegetation community dynamics(Cannone et al. 2007).As On Sforzellina Glacier different geophysicalsurveys
thesechangesfollowthesuddenwarmingof summerand were applied in order to evaluate ice thicknessand
annualtemperatures after1980,theseresultssuggestthat bedrockmorphology.Geoelectricalsurvey(VES [verti-
alpineand nivalvegetation mayrespondfasterand more cal electricalsounding];Resnati and Smiraglia 1989,
flexibly to climaticchange than previouslybelieved(Pauli Guglielminet al. 1995) and seismicreflection (Pavan et
et al. 1999,Waltheret al. 2005). al. 2000) gave a maximumice thicknessof 42 and 60 m,
Glacierforefields are suitableenvironments to investi- respectively. In 1999 a ground probingradar (GPR)
gate the processesof vegetationcolonizationand devel- surveywas performed to obtain high-resolution topog-
opmentand, because of the age controlof the surfaces, to raphy of the glacier bed. The maximumice thickness
calculate the speed of primary succession and to calculatedwas of -60 m in thecentralarea of theglacier
understandtheir mechanisms(e.g., Whittaker1993, and this value agrees with seismicdata. In addition,
Chapinet al. 1994).In theEuropeanAlps severalauthors geomorphological surveyswerealso performed (Rossi et
have investigated glacialchronosequences and described al. 2003) to map the glacial landforms related to past
different stagesof vegetationsuccessionfrompioneerto glacialevolution.
climaxcommunities in relationto siteage (e.g.,Pirolaand The studyarea is includedin the upper alpine belt
Credaro1993,1994,Burga1999,Caccianigaand Andreis (2600-2800 m a.s.l.) and in the nival belt (>2800 m
2004,Raffland Erschbamer 2004) and to themechanisms a.s.l.). The vegetationis a mosaic of discontinuous
of seedlingestablishment (e.g., Stocklinand Baumler alpine grasslands (includingthe climax communities
1996, Niederfriniger Schlag and Erschbamer2000). dominatedby Carex curvulaand the initialgrasslands
Moreover,in highmountainareas,mostglacialforefields dominatedby Poa alpina), snowbed vegetation(with
April2008 CLIMATE CHANGE AND GLACIER ECOSYSTEMS 639

Salix herbacea, Veronica alpina, Sagina saginoides, data with the krigingalgorithm.In addition,several
Cerastiumcerastioides),pioneerand early-successional nodes withknowncoordinateswerepositionednear the
communities(withRanunculusglacialis,Geumreptans, glaciersto improvethe accuracyof the measurements
Cerastium uniflorum,Oxyriadigyna,Saxifragabryoides), takenby differential GPS withshortbases accordingto
and, at the higheraltitudes,scatteredindividualsof Diolaiuti et al. (2004).
vascular plants and cryptogams(see Plate 1). At the The glacierlimitsdatingback to 2002,2003,and 2006
elevationsof our site,theeffectsof anthropogenic land were mapped on the fieldby DGPS surveys,and the
use changeon vegetationare mostlynegligible(Kelleret 1920and 1980limitswerereconstructed by themoraine
al. 2005). Previousstudieson theglacierforefield of the ridgeboundarymarkedon the fieldby DGPS as well.
SforzellinaGlacierwerecarriedout by Caccianiga and The glacierlimitsfrom1991to 2001 werereconstructed
Andreis(2004) reportingthe occurrenceof a pioneer by a geographicinformationsystem(GIS) using the
community (Sieversio-Oxyrietum digynae)withCerasti- measuredfluctuation data and thepositionof the 1991
umuniflorum, Geumreptans,Oxyriadigyna,Ranunculus benchmarkused for measuringthe distancefromthe
glacialis,and Poa laxa on the 1980smorainicridgeand glacier snout. Orthophotosdating back to 1999 and
of the initial grassland dominatedby Poa laxa and 2003 (Regione Lombardia2004) werealso used to map
Saxifragabryoideson the LittleIce Age moraine. the glacierboundaries.GIS mappingwas applied not
Material and Methods only to past glacierlimitsbut also to assign different
ages to theland surfaceson the glacierforefield (1991-
Glaciermonitoring 2001). All the vegetationplots were located by DGPS
Recent (last three decades) glacial changes were and included in the GIS mapping, allowing the
reconstructed using all the available sources of data. vegetationcoveringof these areas to also be dated.
Terminusfluctuations were analyzed from1971 up to Surfacearea changesweredeterminedby GIS through
now, in order to evaluate the glacier's behavior(i.e., thecomparisonof thehistoricalglacierboundariesthus
retreatvs. advance),and to calculatethe ratesof these obtained. The final planar accuracy value was then
changes.These data have been collectedfrombench- evaluatedaccordingto Vogtleand Schilling(1999).
markslocatedon theglacierforefield withoutinterrup-
Vegetation survey
tions,providinga data set on theglacier'sevolutionfor
the last 35 years(CGI 1971-1977,1978-2006).In 1991 The vegetation of the glacierforefield
was described,
the benchmarkused for the terminusvariationmea- analyzinga totalof 23 samplingplotsof 1 X 1 m. Their
surements changedand the new one was set at -80 m positions were acquired through DGPS surveysas
(withan azimuthof 145°N) fromthe present(summer describedpreviouslyand then mapped by GIS. The
2006) glacierlimit. ages of the surfaceswherethe plots weresampledwere
Mass balances were evaluated from the authors obtained fromthe glacieranalysis.Withineveryplot,
(Catasta and Smiraglia1993)from1987up to now using total vegetationcover and the cover of each vascular
thestandardglaciologicalmethod(0stremand Brugman specieswas estimatedvisuallyusing indicesto express
1989,Kaser et al. 2003) based on a networkof ablation therelativeabundanceofeach species(+, <1%; 1, 1-5%;
stakesat different altitudes.Several differentialglobal 2, 5-10%; 3, 10-15%; 4, 15-20%; 5, 20-25%; 6, 25-50%;
positioningsystem (DGPS) campaigns were performed 7, 50-75%; 8, >75%). Vascularplantspeciesnomencla-
with kinematictechniquesaimed at obtainingdigital ture is in accordance with Pignatti (1982). Cover
elevation models (DEM) of the glacier surface and estimateswere also givenfor cryptogams, whichwere
altimetry (in 1999,2000,2002) as wellas to map theGPR not identifiedas species but grouped into three broad
profiles.The fast static techniquewas employed on categories:mosses,epilithiclichens,and groundlichens.
DGPS surveysthat focused on delimitingthe glacier Slope and aspectwererecordedforeach plot.
boundaries or on mapping moraine ridge positions
Climate
(requiringfewerpoints,in 2002, 2003, 2006). DGPS
surveyswere also used to acquire the position of the Local meteorologicaldata were collectedand pro-
benchmark used to measure the glacier terminus cessedto analyzerecentclimatebehaviorand evolution
fluctuations from1991up to now and map themoraines in the studyarea. The closest and highestautomatic
of the 1920sand 1980s,whichservedto reconstruct the weatherstation(AWS), whichwas foundto be running
older and recentglacier advances, as well as to map during the last three decades without significant
supraglacialdebriscoverand thevegetationsurveyplots. interruptions, is located at Forni dam at 2180 m a.s.l.,
The DGPS surveyswerealwayscarriedout at theend ~5 km northwardfrom SforzellinaGlacier. Other
of thesummerseason,whentheablationzone is largest AWSs, whichhave been workingoverthelast 35 years,
and no snow is presenton the glaciersnout.To obtain are located at Santa CaterinaValfurva(1730 m a.s.l.)
DEMs of the glaciersurfaceand bedrockmorphology and Uzza (1250 m a.s.l.), respectively. At thesestations
(fromGPR data), the fieldsurveyswere carried out air temperatureand liquid precipitationdata were
using high densityglobal positioningsystem(GPS) measuredeveryhour and thenrecordedon the general
pointdata (i.e., 8000 points/km2) and interpolating the databaseoftheGeologicalMonitoringServiceofARPA
640 NICOLETTACANNONEET AL. ^^^dT^o °3

Fig. 1. (A) Sforzellina

Glacierinsummer G. Diolaiuti)withdebris
2006(Photocredit coverexceeding
40%ofthewholeglacier
area.(B) Map showingthelocationofthe2006glacierandmoraine ridgesofthe1920sandearly1980sinrelation
tothemountain
ridgesandpeaks(height giveninm a.s.l.).

Lombardia (LombardyRegional Environmental Agen- occurrenceof debris cover that now exceeds 40% of
annual,and seasonal
cy,Sondrio,Italy).Daily,monthly, the whole glacierarea (Figs. 1 and 2). Two ice-contact
averages of hourly air temperaturesand monthly, lakes developedduringthelast 10 years,one of themis
annual, and seasonal cumulated liquid precipitation presently active(at thewestside of theglacier;see map
werecomputedfortheclimateanalysis. in Figs. 1 and 2) and calvingphenomenaoccurredat the
Results glacierwater-contactarea. Glacier volume was deter-
mined fromthe comparisonof the DEMs processed
Glacier fromgeophysical(bedrock topographyby GPR) and
During the last centurySforzellinaGlacier suffered topographical(glaciersurfaceby GPS) data collectedin
substantialreductionsin its lengthand surface,as well 1999. The measuredvolumewas -8.1 x 106m3 of ice
as changes in its supraglacial conditions, with the (i.e., 7.8 X 106m3 waterequivalent,w.e.).
April2008 CLIMATE CHANGE AND GLACIER ECOSYSTEMS 641

between1994 and 2006, the moraineridgesof the early 1920sand

Fig. 2. Map showingthe glacierboundaryfluctuations
1980s,and the locationof thevegetationplots.

The analysis of the mass balance data collectedon w.e.). The wholeloss by ablationprocessesthusexceeds
Sforzellinafrom the hydrologicalyear 1986-1987 to 8 X 106m3(morethantheremainingvolumeestimated
present(Fig. 3) underlinesthat (withexceptionof the in 1999). The rate of the glaciervolumeloss increased
from0.30 X 106 m3/yr
hydrologicalyear2000-2001) the glacieralways lost in significantly w.e. in the period
mass (yearlyaverage of net specificbalance = -1.1 m 1986/1987-1999/2000 to 0.50 X 106 m3/yrw.e. in the

Fig. 3. SforzellinaGlaciernetspecificbalance data (graycolumns,values in m waterequivalent,w.e.) and trends(thickgray

line),and the ELA (equilibriumline altitude)yearlyvalue (curve,in m a.s.l.) and trends(thinblack line).
642 NICOLETTA CANNONE ET AL. Ecological
Applications
Vol.18,No. 3

Fig.4. Glacierterminus
fluctuations
from1971to 2006.Thebarsindicate values,thelinethecumulative
yearly value.When
theglacierterminus (change= 0), no barsarereported.
wasfoundto be stationary

followingsix years.The accumulationarea decreasedas Corresponding to theterminus theglaciated

fluctuation,
the equilibriumline altitude (ELA), calculated from area decreasedfrom0.383 ± 0.005 km2(mean ± SE) in
yearlymass balance profileanalysis,shiftedat higher 1981to 0.2381 ± 0.002 km2in 2006,witha reductionof
elevations(from3029 m a.s.l. in the hydrologicalyear -38%. The yearlyrate of surfacearea loss (Table 1)
1986-1987,to 3189 m a.s.l. in 2005-2006; Fig. 3), and remained almost constant (around -0.005 km2/yr)
presentlyalmost the entireglacierarea lies below the between1981 and 2002, while it doubled (to -0.0097
ELA. The terminusfluctuations (Fig. 4) of Sforzellina km2/yr) in theperiod2002-2006.
Glaciershoweda generalretreatoverthepast 35 years.
During the 1971-2006period,the snout of Sforzellina Vegetation
Glacier retreatedby -75 m. The reductionin length
Vegetationoccursin all theproglacialarea (Tables 2
over the 35 years of surveysequals 9% of the 1971 and 3), showingdifferent patternsin relationto the
glacierlength.The glacierretreated25 yearsout of the surfaceage (Fig. 1; fourthcolumn of Table 2), which
35 yearsanalyzed(equal to 71% ofthetime);theaverage was assigned on the basis of the GIS analysis,and
retreatratewas approximately -2 m/yrover the whole
rangingfrom terrainsdeglaciated after one year to
period(1971-2006),was -2.6 m/yrconsideringonlythe morainedepositsof the 1920s.
shortertimespan 1985-1995,and acceleratedup to -5
Colonizationstartsone year afterdeglaciationwith
m/yrin thelast decade (1996-2006).
In spite of this reduction trend, the terminus generallyverylow coverage(althoughit reaches14% in
fluctuationsalso showeda small glacieradvance in the
Table 1. Yearlyrate of surfacearea loss of Sforzellina
period1975-1984(+14.5 m equal to a rateof-hi.5 m/yr). central
Glacier, ItalianAlps.
It was followedby a transitionphase duringwhichthe
glacier alternatedretreatsand small advances, then, area
Yearlysurface
startingfrom1992,theglaciershrinkageproceeded.The Timeframe change(km2/yr) ± Accuracy
(km2)
advance led to the formationof small moraineridges 1920-1981 -0.0029 0.009
between1977and theearly1990s(Rossi et al. 2003). The 1981-1985 -0.0053 0.008
1985-2002 -0.0050 0.006
glacierlimitsfrom1991 to 2006 and themoraineridges 2002-2006 -0.0097 0.005
of 1920 and 1980 are reportedin Figs. 1 and 2.
April2008 CLIMATE CHANGE AND GLACIER ECOSYSTEMS 643

Table 2. Featuresof investigated

vegetationplots. to persistduringsuccessionas well as to carryout the
' firststagesof colonization)(Jochmisen1970,Matthews
Plot Total No. Age since Slope 1992,
no. cover(%) deglaciation(yr) (°)
Burga 1999). Moreover,these species belong to
species
differentstages of vegetation developmentand to
10 0 0
2 16 10
different vegetationseries.In particularGeumreptans,
3 12 10 Cerastiumuniflorum, and Poa laxa are characteristic
4 14 7 1 0 species of the scree slopes vegetation,Poa alpina and
5 0.1 6 6 are typicalspeciesof theinitialalpine
6 3 5 8 2 Saxifraga bryoides
7 2 1 7 grassland,and Sagina saginoidesis a snowbedspecies.
8 35 6 7 4 On surfacesdeglaciated6-1 1 yearsago, thevegetation
9 7 5 11 2 is still scatteredwith low coverage (2-3% as average
10 22 7 25 2
11 45 8 25 althoughup to 35% in morefavorablesites),composed
12 65 8 25 18 of mosses and 12 species of vascular plants, and
13 18 5 25 20 dominatedby Geumreptansand Cerastiumuniflorum.
14 4 3 25 2
15 20 5 25
Here we observeda changein speciescompositionwith
16 10 4 25 respectto theyoungersurface,withthedisappearanceof
17 18 4 25 Ranunculusglacialis, and a slightlydifferent pool of
18 38 4 25 theingressionof Oxyria
19 0.1 2 25 early-successional species (with
20 15 2 25 digyna)and snowbedspecies{Cerastiumcerastioides).
21 60 9 >80 Vegetationcoverage and species richnessincreases
22 65 12 >80 on the 25-year-oldsurfaces,whereclosed
23 90 12 >80 significantly
patchesofvegetationoccur.The relatively highcoverage
of Oxyriadigyna,similarto thevalues of Geumreptans
some sheltered sites). Vegetation is composed of and Cerastiumuniflorum, indicatesa furtherevolutionof
scatteredindividualsof eightspeciesof vascularplants thevegetationdevelopment.
and by mosses. It is remarkable that, instead of The initialgrasslandand the morematuregrassland
exclusivelypioneerspecies (i.e., species that appear in of theLuzuletumspadiceaeoccuron surfacesolderthan
thefirststagesofcolonizationbutare notable to persist 80 years, where it is possible to observe a shiftin
during succession), the firstcolonizers are mainly communitycomposition,with the dominanceof Poa
composedof early-successional species(i.e., speciesable alpina and/orLuzula spicata, the ingressionof Salix

Table 3. Relativeabundanceof speciesin vegetationplots.

Relativeabundanceby plot

Species 12 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 .21 22 23
Ranunculus glacialis + + + + + + 12
Cerastiumuniflorum + 1 + 1 2 2171 1112 14
Mosses + + 1 1 1 1 11 2 12
Poa laxa + 1 +1 2
Geumreptans +1 + 1121162111 + 34 + 3 5
Leucanthemopsis alpina +1 + 1 2+ 21 33
Saxifragabryoides 1 1 111
Poa alpina +1 6 2 1
Sagina saginoides + + 1 + 1 +
Poa alpina vivipara + 1 1
Oxyriadigyna + 11111 12 3 1
Cerastiumcerastioides + 3 1
Cerastium pedunculatum 1
Arabiscaerulea + + + 1 + 1
Veronicaalpina 2 1 1
Linariaalpina 1
Epilobiumanagallidifolium
Taraxacumalpinum +
Sedumalpestre 1 +
Salix herbacea 2 + 1
Saxifragaoppositifolia 1
Armeriaalpina +
Luzula spicata 7
Cardamineresedifolia +
Groundlichens 1
Epilithiclichens 1
Note: Indicesexpresstherelativeabundanceof each species(+, <1%; 1, 1-5%; 2, 5-10%; 3, 10-15%; 4, 15-20%; 5, 20-25%; 6,
25-50%; 7, 50-75%; 8, >75%).
644 NICOLETTA CANNONE ET AL.
Fig. 5. Mean summerair temperatures in theperiod 1988-2005recordedat Uzza (solid dots) and Forni(solid
(June-August)
triangles).
herbacea and Sedum alpestre (snowbed species also
occurringfrequently in the climax grassland),and the
persistenceof the early-successionalGeum reptans,
Cerastiumuniflorum, and Leucanthemopsis alpina, and
average total coverageup 90%. Only at this stage do
lichensmaketheiringressionas epilithiclichensgrowing
on bouldersas well as groundfolioselichens.
Climate
Despitethegeneralstrongwarmingfoundin theAlps
(Beniston2000), in ValfurvaValley duringthe period
1988-2006 the mean annual air temperatures show a
veryslightdecrease(-0.2°C at 2180 m a.s.l.), whilethe
mean summer (June-August) air temperaturesare
clearly increasing(+0.5°C; Fig. 5). The precipitation
patternwe evaluatedonlyreferred to liquidprecipitation
(as theclosestAWS [Forni,2180 m a.s.l.] onlyrecorded
water equivalentdata); in the areas surroundingthe
glacierdata for snow precipitationwere not available,
making it difficult
to evaluateanychangein snowdepth
and duration.In anycase, a remarkabledecreasein total
precipitation (around -10% at 2180 m a.s.l.) has been
recordedsince 1988.
Discussion
At a global scale,glaciersare verysensitiveto climate
change. In particular,the smaller ones seem to be
consistentclimatechangeindicators,giventhattheyare
showingfasterreactiontimes (Dyurgerovand Meier
2000, Paul et al. 2004). On the otherhand, previous
studiessuggestit would be inappropriateto use length
changes of a single glacier as being representative of
climatechange(Chinn 1999).
Our data show the trendof terminusfluctuations of
SforzellinaGlacier to be in agreementwiththe general
patternof glaciers spread all over the Alps, which
resultedin retreating fromtheend of the LittleIce Age
Ecological
Applications
Vol.18,No. 3
(LIA) up to now (Zemp et al. 2006) with a short
interruption that occurredbetweenthe 1970s and the
1980s(Patzelt1985,Wood 1988).In addition,therecord
of glaciermass balances,whichare generallyconsidered
an unambiguousmarkerof climatechange(Cogleyand
Adams 1998,Haeberliet al. 1999,Dyurgerovand Meier
2000, Oerlemans2005), underlinesa strongervolume
reductionin thelast decade (theaverageyearlyvalue of
the last 10 years was 66% largerwith respectto the
1987-1996averageyearlyvalue).
Moreover,the representativeness of SforzellinaGla-
cier as witnessof the ongoingalpine glacierchangesis
furthersupportedby the strikingrelationshipbetween
theSforzellinamass balance (1986/1987-2002/2003) and
thoseof thealpineglaciersreportedin theGlacierMass
Balance Bulletins (IAHS (ICSI)-UNEP-UNESCO,
1988-2005;Bravais-Pearsoncorrelationcoefficient, r=
0.84). Furthermore,the ELAs of SforzellinaGlacier
(1986/1987-2002/2003) comparedwiththe mean alpine
ELA (IAHS (ICSI)-UNEP-UNESCO, 1988-2005)also
show a strongcorrelation(r = 0.82).
An acceleratingsurfacearea loss was also foundfor
the period 2002-2006, and the glaciersurfacein 2006
resultedin a decreased of approximately-14% with
respectto the2002 glacierarea.
The glacial reductiondemonstrated by thesefindings
(i.e., terminusfluctuations, mass balances,and surface
area changes) is interpretedas a truthfulimpact of
climatechange.Indeed,as the transfer functionamong
climatechangesand glaciervariationsdoes not change
in time (Oerlemans 2005), an accelerationin glacial
changesis suggested.
Nevertheless,it is less easy to identifythe climatic
elementsdrivingthe glacier changes and, despite the
many investigations on this topic (e.g., Hoelzle et al.
2003,Oerlemans2005), thescientific debateis stillopen.
Air temperature and precipitation,thetwo factorsmost
April2008 CLIMATE CHANGE AND GLACIER ECOSYSTEMS
Plate 1. Androsacealpinais a pioneerspeciescolonizingthealpineand nivalbeltsof theEuropeanAlps and is representative
of the speciesactuallysuffering
the highestimpactsof climatechangeon high-elevation
commonlycorrelatedwithglacierfluctuations, are only
two elementsof the complexchain of processeslinking
climateand glacierfluctuations(Haeberli 1995, Chinn
1999). Therefore,the glacier fluctuationsindicate a
complexcombinationof mass and energyexchangeat
the Earth's surface. The air temperatureincrease
occurringin the alpine areas since the end of the LIA
activated a positive feedback, with the consequent
increaseof both the downwardsensibleheat fluxand
thelong-wave radiationbalance (Oerlemanset al. 1998).
Furthermore, duringthe last two decades, Sforzellina
Glacierexperienceda strongdecreaseof surfacealbedo
(due to increasingdebriscoverage),whichsurelyplayed
a key role in increasingthe glacier absorption of
incomingenergyfluxes,thusmakinglargerthequantity
of energyavailable forglaciermelting.
To quantifythe possibleinfluenceof temperature on
the recentevolutionof SforzellinaGlacier, neglecting
thecontribution due to changingprecipitation, a simple
approach was followed,according to Chinn (1999) and
Oerlemans(2001). Analyzingall theEL As of theperiod
1987-2006 (Fig. 3), the averagedifference betweenthe
estimatedsteady-stateELA and the calculated annual
ELAs, givesa mean increaseof -150 m. This upward
shift, obtained using a standard lapse rate of
- 0.0065°C/m,represents a generalwarmingof approx-
imately + 0.6°C since 1987. Therefore,consideringthat
thesummerwarmingthatoccurredin thesame periodis
645
ecosystems.Photo credit:N. Cannone.
+0.5°C, the difference indicatesthatchangesin precip-
itationalso have to be considered.
In the context of vegetationsuccession,our data
supportrecentevidence(Cannoneet al. 2007) suggesting
that the significant changes in vegetationcommunity
dynamicswe foundare consequencesof an actual air
temperature warmingcombinedwiththe reductionof
precipitationand the shorteningof the snow season.
Like the glacier reduction,all processes seem to be
accelerated,includingthevegetationdynamics.
The comparisonof our data withglacial chronose-
quences describedfor the European Alps allows an
assessmentof theaccelerationrateof theseprocesses.In
the alpine belt (>2200 m a.s.l.) the colonizationof the
recentlydeglaciatedterrainsis reportedto startwithin
4-8 yearsafterdeglaciation(Stocklinand Baumler1996,
Burga 1999,Tscherkoet al. 2005), withscatteredearly
pioneerspecies.At least 10-25 yearsare requiredforthe
ingressionof the early-successionalspecies and the
developmentof a well-establishedearly-successional
community(Pirola and Credaro 1993, Niederfriniger
Schlag and Erschbamer2000, Caccianiga and Andreis
2004, Raffland Erschbamer2004,Tscherkoet al. 2005).
Our data provideevidencethattheseearlystagesof
colonization suffereda dramatic acceleration:plant
species are able to colonize the glacier forefieldonly
one year after the glacier retreat.Consideringthe
differences of colonizationtimebetweenour data and
the existingliterature,the colonizationspeed increased
646 NICOLETTA CANNONE ET AL.
Ecologica^AppHcations

at least four timesfor the establishment of scattered clonal plantson theone-year-old surfacesdemonstrates
individuals(one yearin theSforzellinaarea vs. 4-8 years the correctnessof this hypothesisand explains how
reportedbyliterature) and about two timesforthewell- colonization could occur with such fast rates at an
establisheddiscontinuousearly-successional community elevation>2800 m a.s.l.
(6-1 1 yearsat Sforzellinavs. 10-25yearsreportedbythe Accelerationof dynamicsis evidentonly on recent
literature). vegetation;the factthat older stages of successionare
On theotherhand,thefollowingstagesof vegetation quite similar to those described by other glacial
development(>25 years) do not show remarkable chronosequencesindicatesthatclimatechangeimpacts
differences with respect to the chronosequencesde- on these stages are not severe enough to induce
scribedin theliterature fortheAlps (such as Pirola anc significant changesand thatolder and moredeveloped
Credaro 1993, Niederfriniger Schlag and Erschbamer stages are probably more resistantto, and buffered
2000, Caccianiga and Andreis2004, Raffland Ersch- from,climaticperturbation.
bamer2004,Tscherkoet al. 2005). Withinthe well-documented increaseof air tempera-
The ecologicalrequirements of the speciesoccurring ture duringthe 20th centuryover the planet (IPCC
on the one-year-oldsurfacemay explainthe abilityof 2001), the warmingwas significantly higher in the
thesespeciesto colonize such youngsubstrata.In fact, European Alps (Beniston 2000, Bohm et al. 2001).
mostof themare speciestypicalof thescreeslopesor of More recentlyCasty et al. (2005) in theirtemperature
rocks, with only one snowbed species (Sagina sagi- reconstruction fortheAlps since 1500,foundthat1994,
noides). The absence of snowbed species in a glacier 2000, 2002, and 2003 werethewarmestyearssince1500.
forefield appearsto be a paradox ifwe do not take into Moreover, they found that summer warming was
accountthe strongreductionof snow cover abundance particularlysevere after 1970, reaching,in 2003, the
and permanencedocumentedfora siteveryclose to the highestpeak of summertemperaturesince 1500. Our
studyarea (Cannone et al. 2007). The shortening of the data confirmthe generalsummerwarmingof the last
snowcoverlengthsupportsthehypothesis byGalen and two decades, even if, at this site,verywarm summers
Stanton(1995) that climatechange may induce inter- wererecordedalso in 1991 and 1998.
specificdifferences in growthphenologyof coexisting The temperatureincrease we found (-K).5°C in
species and promote shifts in snowbed plant communi- summer)has to be consideredin the frame of the
ties.Moreover,Sagina saginoidesis a short-lived species generalpost-1985 climatewarming,whichwas accom-
withabundantseed production,and it is possible that panied by reducedprecipitationon the Alps. Whereas
the successof thisspeciesmay be relatedto its efficient the numberof days withsnow on the groundshowed
dispersalstrategies. littleevolution,the durationof continuoussnow cover
The functional types of most of thespecies involved in was clearly declining at all elevations. Although
the earlystagesof colonizationmay also providesome continuoussnow cover tendedto startearlier,it also
explanation for the accelerated rates of vegetation meltedmuch earlier(Benistonet al. 2003, Martinand
colonization.All thesespeciesare long-livedperennials Durand 2006). This generaltrendseems relatedto a
adapted to the harshenvironmental conditionsof the positiveNorth AtlanticOscillation(NAO) index (Up-
alpineand nival beltsand of the glacierforefield. Most penbrink 1999) and to the summer warming.Our
of theearly-successional speciesthatare able to colonize precipitationdata indicatea generaldecrease (-10%)
the one-year-oldterrainsare clonal species,including in agreementwith the general trend (Brunettiet al.
clonal plants with widely spaced ramets(e.g., Geum 2000). It is unfortunatethat thereare not any snow
reptans,Saxifragabryoides, Cerastiumuniflorum) as well depth and durationdata available for the Sforzellina
as clonal plantswitha clumpedgrowthform(e.g., Poa Glacier forefield, but if we considerother local data
alpina). Clonal growthis one of the most important recordedin another localityof the Upper Valtellina
adaptationsto the severeclimaticconditionsand the since 1978,we findtheyagreewiththegeneralpatterns
nutrientshortagecharacteristic of the alpine environ- of the snow coverdecrease(Cannone et al. 2007).
ments(Stocklinand Baumler1996,Pluess and Stocklin
Conclusions
2005). Thus clonal reproductionincreaseswithaltitude
both in closed grasslandsand in pioneercommunities The strongerice mass loss affectingthe Sforzellina
(Stocklin1992). Clonal growthis a key factorfor the Glacierduringthelast decade (1996-2006) may be due
successful establishmentof the primarysuccession to the local summerwarmingcombinedwithreduced
becauseit provideslong-livedperennialswiththeability precipitationand the shorteningof the snow season
to persistduringsuccession(e.g., Geumreptans)witha (Cannone et al. 2007). This abiotic evidenceof climate
large amount of phenotypicplasticity(Stocklin and change is correlatedwithchangesin the vegetationof
Baumler1996). Phenotypicplasticityis one of themost the glacierforefield, whichindicatedan accelerationof
importantmechanismshypothesizedallowingplantsto colonizationratesin thesameperiod.Herewe showthat
persistin the environment modifiedby climaticchange thereis a strongcorrespondence betweentheabioticand
(Theurillat and Guisan 2001), thus avoidingmigration biotic of
components high altitudeecosystemsand that,
and/or extinction.Therefore the preponderanceof overthelast decade (1996-2006) and, in particularsince
April2008 CLIMATE CHANGE AND GLACIER ECOSYSTEMS
2002, the impactsof climatechangehave undergonea
dramaticacceleration,much strongerthan any that
occurredduring the last 35 years. This acceleration
occurredin tandemwitha documentedslightsummer
warmingand a possibledecreasein snowcover.Further
in situ investigations
are requiredto directlymeasure
some of the influential
parametersinvolved(e.g., snow
cover, avalanche frequency)and to analyze species-
specificresponsesto factorslike snow coverreduction.
ACKNOWLEDGM ENTS
We thank Christian Korner and Renato Gerdol for
reviewingthe paper beforesubmission,WilfriedHaeberli and
one anonymous refereewhose commentsand suggestions
allowed us to improve the paper, and the Geological
MonitoringServiceof ARPA Lombardia(LombardyRegional
Environmental Agency)forthe climaticdata. This paper was
funded by the project "Increasing rate of climate change
impactson high mountainareas: cryosphereshrinkageand
environmentaleffects"(PRIN-MIUR 2005).
Literature Cited
Beniston,M. 2000. Environmental change in mountainsand
uplands.Arnold,London, UK.
Beniston,M. 2003. Climaticchange in mountainregions:a
reviewof possibleimpacts.ClimaticChange 59:5-31.
Beniston,M., H. F. Diaz, and R. S. Bradley.1997. Climatic
changeat highelevationsites:a review.ClimaticChange 36:
233-251.
Beniston,M., F. Keller,and A. Goyette.2003. Estimatesof
snow accumulationand volume in the Swiss Alps under
changing climatic conditions. Theoretical and Applied
Climatology76:125-140.
Bohm,R., I. Auer,M. Brunetti, M. Maugeri,T. Nanni,and W.
Schoner. 2001. Regional temperaturevariabilityin the
European Alps: 1760-1998fromhomogenizedinstrumental
timeseries.InternationalJournalof Climatology21:1779-
1801.
Brunetti,M., M. Maugeri,and T. Nanni. 2000. Variationsof
temperature and precipitationin Italy from 1866 to 1995.
Theoreticaland AppliedClimatology65:165-174.
Burga, C. A. 1999. Vegetationdevelopmenton the glacier
forefieldMorteratsch(Switzerland). Applied Vegetation
Science2:17-24.
Caccianiga, M., and C. Andreis. 2004. Pioneer herbaceous
vegetationon glacierforefields in the Italian Alps. Phyto-
coenologia34(l):55-89.
Cannone, N., S. Sgorbati,and M. Guglielmin.2007. Unex-
pected impacts of climaticchange on alpine vegetation.
Frontiersin Ecologyand the Environment 7:360-364.
Casty, C, H. Wanner,J. L. Luterbacher,J. Esper, and R.
Bohm.2005. Temperature and precipitation in the
variability
European Alps since 1500. InternationalJournalof Clima-
tolology25:1855-1880.
Catasta, G., and C. Smiraglia.1993. The mass balance of a
cirqueglacierin theItalianAlps (Ghiacciaiodella Sforzellina,
Ortles-Cevedale Group). Journalof Glaciology39:87-90.
CGI (ComitatoGlaciologico Italiano). 1971-1977.Campagne
Glaciologiche(Glaciological surveys).Bollettinodel Com-
itatoGlaciologicoItaliano SerieI and 11:18-25.
CGI (ComitatoGlaciologico Italiano). 1978-2006.Campagne
Glaciologiche (Glaciological surveys).Geografia Fisica e
Dinamica Quaternaria:l-25.
Chapin,F. S., L. R. Walker,C. L. Fastie,and L. C. Sharman.
1994. Mechanismsof primarysuccessionfollowingdeglaci-
ation at Glacier Bay, Alaska. Ecological Monographs64:
149-175.
647
Chinn, T. J. 1999. New Zealand glacierresponseto climate
changeof the past 2 decades. Global and PlanetaryChange
22:155-168.
Cogley,J.G., and W. P. Adams. 1998.Mass balanceofglaciers
otherthantheice sheets.Journalof Glaciology44(147):315-
325.
Cook, A. J., A. J. Fox, D. G. Vaughan, and J. G. Ferrigno.
2005. Retreatingglacierfrontson the AntarcticPeninsula
over thepast half-century. Science308:541-544.
Diolaiuti,G., C. Smiraglia,G. Vassena, and M. Motta. 2004.
Dry calvingprocessesat the ice cliffof StrandlineGlacier,
NorthernVictoriaLand, Antarctica.Annals of Glaciology
39:201-208.
Dyurgerov,M. B., and M. F. Meier. 2000. Twentiethcentury
climatechange:evidencefromsmallglaciers.Proceedingsof
theNational Academyof Sciences(USA) 97:1407-1411.
Egh, M., M. Wernh,C. Kneisel, and W. Haeberli. 2006.
Meltingglaciersand soil developmentin the proglacialarea
Morteratsch(Swiss Alps): I. Soil type chronosequence.
Arctic,Antarctic,and Alpine Research38:499-509.
Galen, C, and M. N. Stanton. 1995. Responsesof snowbed
plant-speciesto changesin growingseason length.Ecology
76:1546-1557.
Grabherr,G., M. Gottfried, and H. Pauli. 1994.Climateeffects
on mountainplants.Nature369:448.
Guglielmin,M., A. Nardo, and C. Smiraglia.1995.Lo spessore
dei ghiacciaidella Valfurva.MisurazionitramiteSondaggi
ElettriciVerticali.Neve e Valanehe 24:58-67.
Haeberli, W. 1995. Glacier fluctuationsand climatechange
detection - operationalelementsof a worldwidemonitoring
strategy. WorldMeteorologicalOrganisationBulletin44:23-
31.
Haeberli,W., and M. Beniston.1998. Climatechangeand its
impactson glaciersand permafrost in the Alps. Ambio 27:
258-265.
Haeberli,W., R. Frauenfelder, M. Hoelzle, and M. Maisch.
1999.On ratesand accelerationtrendsof global glaciermass
changes.GeografiskaAnnaler81A:585-591.
Haeberli,W., and M. Hoelzle. 1995. Applicationof inventory
data for estimatingcharacteristics of and regionalclimate-
changeeffectson mountainglaciers:a pilot studywiththe
EuropeanAlps. Annalsof Glaciology21:206-212.
Hoelzle, M., W. Haeberli,M. Dischl, and W. Peschke.2003.
Secularglaciermassbalancesderivedfromcumulativeglacier
lengthchanges.Global and PlanetaryChange 36:295-306.
IAHS (ICSI)-UNEP-UNESCO. 1988-2005. Glacier Mass
Balance Bulletins(GMBBs). Volumes 1-8. World Glacier
MonitoringService,Zurich,Switzerland.
IPCC. 2001. ThirdAssessmentReportof the Intergovernmen-
tal Panel on ClimateChange. CambridgeUniversityPress,
Cambridge,UK.
Jochmisen,M. 1970. Die Vegetationsentwicklung auf Morae-
nenboden in Abhaengigkeitvon einigen Umweltfaktoren.
VerlagUniversitatInnsbruck46:1-22.
Kaser, G., A. Fountain,and P. Jansson.2003. A manual for
monitoringthe mass balance of mountain glaciers. IHP
(InternationalHydrologicalProgramme),Technical Docu-
mentsin Hydrology,59. UNESCO, Paris,France.
Keller, F., S. Goyette,and M. Beniston. 2005. Sensitivity
analysisof snowcoverto climatechangescenariosand their
impacton plant habitatsin alpine terrain.ClimaticChange
72:299-319.
Keller, F., F. Kienast, and M. Beniston.2000. Evidence of
responseof vegetationto environmental change on high-
elevationsitesin the SwissAlps. Regionaland Environmen-
tal Change l(2):70-77.
Martin,E., and Y. Durand. 2006. Precipitation and snowcover
variabilityin the French Alps. Lecture Notes in Earth
Sciences.Volume74. Springer,Berlin/Heidelberg, Germany.
648 NICOLETTA CANNONE ET AL.
Matthews,J. A. 1992. The ecology of recentlydeglaciated
terrain.CambridgeUniversity Press,Cambridge,UK.
Niederfriniger Schlag,R., and B. Erschbamer.2000. Germina-
tion and establishment of seedlingson a glacierforelandin
the Central Alps, Austria. Arctic,Antarctic,and Alpine
Research32:270-277.
Oerlemans,J. 2001. Glaciers and climatechange. Balkema,
Rotterdam,Netherlands.
Oerlemans,J.2005. Extracting a climatesignalfrom169glacier
records.Science308:675-677.
Oerlemans,J., B. Anderson,A. Hubbard,Ph. Huybrechts, T.
Johannesson,W. H. Knap, M. Schmeits,A. P. Stroeven,
R. S. W. van de Wai, J. Wallinga, and Z. Zuo. 1998.
Modelling the response of glaciers to climate warming.
ClimateDynamics14:267-274.
Oerlemans,J.,and J.P. F. Fortuin.1992.Sensitivity of glaciers
and smallice caps to greenhousewarming.Science258:115-
117.
0strem, G., and M. Brugman. 1989. Glacier mass-balance
measurements.National Hydrology Research Institute,
Saskatoon,Saskatchewan,Canada.
Patzelt,G. 1985. The period of glacieradvances in the Alps,
1965to 1980.Zeitschrift furGletscherkunde und Glazialgeo-
logie 21:403^07.
Paul, F., A. Kaab, M. Maisch, T. Kellenberger,and W.
Haeberli. 2004. Rapid disintegrationof Alpine glaciers
observedwith satellitedata. GeophysicalResearch Letters
31:L21402[doi: 10.1029/2004GL0208161.
Pauli, H., M. Gottfried,
and G. Grabherr.1999.Vascularplant
distribution patternsat the low temperature limitsof plant
life- theAlpine-nivalecotoneof Mount Schrankogel(Tyrol,
Austria).Phytocoenologia29:297-325.
Pavan, M., G. Diolaiuti, V. Maggi, C. Smiraglia,and C.
D'Agata. 2000. Prospezionigeofisichesul Ghiacciaio della
Sforzellina(Alpi Lombarde,Gruppo Ortles-Cevedale).Ri-
sultatipreliminari.Neve e Valanghe41:6-13.
Pignatti,S. 1982. Flora d'ltalia. Three volumes. Edagricole,
Bologna,Italy.
Pirola,A., and V. Credaro.1993.Changesin thevegetationofa
recent glacial moraine in the Bernina group. Annali di
BotanicaLI:145-153.
Pirola,A., and V. Credaro. 1994. Osservazionisul dinamismo
della vegetazionedi morenarecentenel Gruppodel Bernina.
Fitosociologia27:139-149.
Pluess, A. R., and J. Stocklin. 2005. The importanceof
population origin and environmenton clonal and sexual
reproductionin the alpine plant Geum reptans.Functional
Ecology 19:228-237.
EcologicalApplications
Vol. 18,No. 3
Raffl,C, and B. Erschbamer.2004. Comparativevegetation
analyses of two transectscrossinga characteristic glacier
valleyin theCentralAlps. Phytocoenologia32:225-240.
RegioneLombardia.2004. LombardyGlacierdata base. (www.
cartografia.regione.lombardia.it)
Resnati,C, and C. Smiraglia.1989.Ghiacciaiodella Sforzelli-
na. Misuredello spessoredel ghiacciaioattraversosondaggi
geoelettrici.RivistaClub Alpino Italiano 110(6):70-75.
Root, T. L., J. T. Price, K. R. Hall, S. H. Scheider,C.
Rosenzweig,and A. J. Pounds. 2003. Fingerprints of global
warmingon wildanimalsand plants.Nature421:57-60.
Rossi, S., G. Diolaiuti,E. Forasacco, M. Pelfini,C. Smiraglia,
and P. Spreafico.2003. Evidenzegeomorfologiche della piu
recenteespansione glaciale correlabileall'episodio freddo
deglianni '50-70 del XX secolo sulleAlpi Lombarde.Pages
377-395 in M. Motta and A. Biancotti,editors.Rispostadei
processi geomorfologicialle variazioniambientali.Glauco
Brigati,Genoa, Italy.
Rott, H., P. Skvarca,and T. Nagler. 1996. Rapid collapse of
northernLarsen Ice Shelf,Antarctica.Science271:788-792.
Stocklin,J. 1992.Umwelt,MorphologieundWachstumsmuster
klonalerPflanzen - eineUbersicht.BotanicaHelvetica102:3-
21.
Stocklin, J., and E. Baumler. 1996. Seed rain, seedling
establishmentand clonal growthstrategieson a glacier
foreland.Journalof VegetationScience7:45-56.
Theurillat,J. P., and A. Guisan. 2001. Potentialimpact of
climatechangeon vegetationin theEuropeanAlps: a review.
ClimaticChange 50:77-109.
Tscherko,D., U. Hammesfahr, G. Zeltner,E. Kandeler,and R.
Bocker. 2005. Plant successionand rhizospheremicrobial
communitiesin a recentlydeglaciatedalpine terrain.Basic
and AppliedEcology6:367-383.
Uppenbrink,J. 1999. The NorthAtlanticOscillation.Science
283:948-949.
Vogtle,T., and K. J.Schilling.1999.Digitizingmaps.Pages201-
216 inH.-P. Bahrand T. Vogtle,editors.GIS forenvironmen-
tal monitoring. Schweizerbart, Germany.
Stuttgart,
Walther,G. R., S. BeiBner,and C. A. Burga.2005. Trendsin
the upward shiftof alpine plants. Journalof Vegetation
Science 16:541-548.
Whittaker,R. J. 1993. Plant populationpatternsin a glacier
foreland succession: pioneer herbs and later-colonizing
shrubs.Ecography16:117-136.
Wood, F. 1988. Global alpine glaciertrends1960s to 1980s.
Arctic,Antarctic,and AlpineResearch20:404-413.
Zemp, M., W. Haeberli,M. Hoelzle,and F. Paul. 2006. Alpine
glaciersto disappearwithindecades?GeophysicalResearch
Letters33:L13504. [doi: 10.1029/2006GL026319]