An Experimental Comparison of Leaf Decomposition Rates in a Wide Range of Temperate

Plant Species and Types

Author(s): J. H. C. Cornelissen
Source: Journal of Ecology, Vol. 84, No. 4 (Aug., 1996), pp. 573-582
Published by: British Ecological Society
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Journalof
Ecology1996,
An experimentalcomparison
ofleafdecompositionrates
84, 573-582 in a widerangeoftemperate
plantspeciesand types
J. H. C. CORNELISSEN
NERC Unitof ComparativePlant Ecology,Department
ofAnimal& Plant Sciences,The University,
Sheffield
S1O 2TN, UK

Summary
1 An experimental multispeciesscreeningofleafdecompositionrateswas undertaken
in order to identifyand quantifygeneralpatternsin leaf decompositionrates in
functionalplanttypesand taxa. Functionalspeciesgroupswerecharacterizedusing
whole-plantand whole-leaffeaturesrelevantto the functioning of plants in their
naturalenvironments.
2 The experimentincludedfreshleaf littersof 125 Britishvascular plant species,
coveringa wide rangeof life-forms, leaf habitsand taxa. Preweighedlittersamples
wereenclosedin two typesof litterbags and exposed to naturalweatherconditions
and soil-bornedecomposersby buryingthem simultaneouslyin an experimental
outdoorleaf-mouldlayer.
3 Relative litterdry weightlosses showed largelysimilarpatternsamong species
betweenboth litterbag types,between8 and 20-weekburialperiodsin winterand
betweenwinterand summerburial.
4 Life-form, deciduousvs. evergreenhabit,autumncolorationof leaf litter,family
and evolutionary advancementsensuSpornecould each explainpartofthevariability
in litterdryweightloss among species.The correlationwithlitterspecificleaf area
appearedconfoundedwithtaxonomy.
5 Some of theseeasy-to-assesspredictorsof species'relativeleafdecompositionrates
may proveusefulformodellingsoil decompositionratesundervegetationsdiffering
in speciescomposition.
Stace (1991)
Nomenclature:
Keywords:colour,deciduous,evergreen,
functional,
herb,life-form,
litter,taxonomy,
weightloss,woody
JournalofEcology(1996) 84, 573-582

surface.The potentialdecompositionrate forleaves

Introduction
ofa givenspeciesis a functionoftheinteractiveeffects
Itiswellestablished thatpotential decomposition rate of all theseand otherfactors,each ofwhichmayvary
ofleaflitter dependsgreatly on thephysico-chemical greatlywithinfunctionalor taxonomicgroups.This,
properties oftheleavesofthespeciesconsidered (e.g. combinedwiththe small numbersof speciesused in
Broadfoot& Pierre1939; Aber et al. 1990; Gillon et moststudies,has made it difficultto identify
patterns
al. 1994;Couiteauxet al. 1995). As reviewedby Swift and testgeneralitiesin potentialleaf decomposition
et al. (1979) and Anderson(1991), severalfeatures ratesamong speciesgroups.Withoutthiswe cannot
ofundecomposed leaflitter
arenegatively associated evaluateor predicttheeffects ofchangingplantspec-
withdecomposition rate.Thesefeatures includelignin ies compositionon soil decompositionratesin differ-
content(Heal et al. 1978; Taylor et al. 1989; Berget ent ecosystems.One new way forwardis to screen
al. 1993; Van Vuurenet al. 1993), contentof other representativenumbersofspeciesor groupsofspecies
phenoliccompounds suchas tannins(Nicolai1988; thatare identifiablein termsof functionalattributes
Kuiters1990),lignin:nitrogen ratio(Melilloet al. or taxonomy.
1982; Buth & Voesenek 1987; Cotrufoet al. 1994), This paper reportsan experimental, simultaneous
c) 1996British physicalleaftoughness (Gallardo& Merino1993) screeningof leafdecompositionratesfor 125 species
Ecological Society and physical barriers
(e.g.hairs,spines,wax)on the that are native,naturalisedor commercially impor-

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574 tant in the BritishIsles. It teststhe hypothesisthat
Leaf potentialdecompositionrateof leaflittercan be pre-
decomposition dictedfromwhole-plantfeaturesthatreflect thefunc-
rates tioningof the plants in theirnaturalenvironments.
These featuresinclude life-form and deciduous vs.
evergreenhabit,as well as specificleaf area and aut-
umncolorationofthefreshlitter.Sourcesofvariation
in potentialdecompositionrateare testedusingfunc-
tional analysesof a representative part of a present-
day floraas wellas taxonomicanalyses.
Methods
SELECTING AND COLLECTING LEAVES
Leaf litterwas collectedwithina 25-kmradiusaround
Sheffield,UK (53?23'N, 1?33'W),wherepossiblefrom
the species' (semi)naturalsites includingparks and
woodlands.For deciduouswoody species,theplants
thatweresampledshowedtypicalautumncoloration
(see below) as seenin theSheffieldarea. Some species
thatcould not be foundin thewild locallywerecol-
lectedfromSheffield University'sExperimentalGar-
dens and a few fromprivategardens.The original
data can be retrieved(as AppendixI) fromthejour-
nal's archiveon the World Wide Web (see a recent
issuefordetailsoftheJournalofEcologyhomepage).
All but one species had thereforebeen exposed to
similarclimaticconditions,which facilitatedinter-
specificcomparisonoflitterweightlosses.OnlyPicea
sitchensisleaves were collectedoutside the area, in
northernScotland.
Throughoutautumn1993,freshly senesced,unde-
composedleaveswerecollectedfromsexuallymature
plants. In virtuallyall woody and some of the her-
baceous species this meant that they were either
picked up fromthe groundwithina fewdays after
falling,or collectedaftershakingtheplantgently.In
many herbaceousspeciesthe entireshoot dies back
and fallsover. Since, in thesecases, leaves were not
shed, we harvestedthose that had lost theirgreen
colour and could be assumed to have been func-
tionallydisconnectedfromtheplant.Leaves werecol-
lectedwithoutbias towardssize,shape or colour,but
those with symptomsof significant herbivorywere
avoided. In some summer-shedding evergreens,the
collectedleaves weresuspendedin the standingveg-
etationor on top ofthelitterlayer.For theseand a few
othersummer-shedding evergreenspeciesadditional
collectionsof freshlyshed leaves weremade in sum-
mer 1994 and treatedin a controlexperiment(see
below). Petioleswereconsideredpart of the leaf. In
compound leaves the laminae and rhachisweretre-
ated separately.This would permitinterspecific com-
parison of whole leaves as well as leaf analogues.
For Cytisusscopariusand Ulex spp. smalltwigswere
? 1996 British
EcologicalSociety, treatedas leaves, because in these species the twigs
JournalofEcology, are the main photosynthetic producers.For Calluna
84, 573-582 vulgarissmall shoots (78% of dryweightbeingleaf)
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were treatedas leaves, since theywere shed in this
form.In the biennialsAnthriscus sylvestris
and Digi-
talispurpureabothfirstand secondyear'sleaveswere
harvestedafterthesecondgrowingseason; in thelat-
ter species both rosetteleaves and leaves on the
flowering stemsweretaken.
Initial mean specificleaf area (SLA, area: dry
weightratio) of the litterwas calculated fromthe
area (estimatedusingpunchedlaminadisksofknown
diameteror measuredusing a Delta-T Area Meter,
Burwell,Cambridge,UK) and subsequently theoven-
dryweightin a minimumsubsampleof eightleaves.
Since surfaceaccess to soil decomposerswas likelyto
be a determinant ofdecompositionrate,thetrueone-
sidedsurfacearea ofnonlaminarleaveswas calculated
bymultiplying theprojectedleafarea bya conversion
factorderivedfromobservationsof leaf cross-sec-
tions.
As autumnleaf colorationmay reflectthe chem-
istryoflivingleavesthatsecondarilyaffects decompo-
sition rate, each deciduous woody species was
assignedto one of ninequalitativeleafcolourclasses
(definedin WWW archive)based on thepredominant
colour(s).Sincecolourcan changerapidlybeforeand
afterleaf fall (Hendryet al. 1987), the leaves were
collectedveryshortlyafterabscission.For virtually
all speciesa typicaloverallcolour class was evident
even when therewas variabilitybetweenor within
plants.Cornussanguineaand Ligustrum vulgareleaves
werecollectedfromshadedshrubsand weretherefore
green,althoughleaves whichhave been exposed to
fullsunlightmayshowpartialor totalpurplishcolor-
ation.Althoughchoosingthecolourclasswas to some
degreesubjective,it was objectivein thesensethatit
was done beforetheexperimental treatment.
PREPARATION OF THE LEAF MATERIAL
The littercollectionswere sortedand cleaned,then
air-driedand stored in open paper bags in a lab-
oratory(20 ?C). Most leavesreachedtheirequilibrium
moisturecontent(7-8%) withinfourdays. Samples
of air-dryleaf material(1.0 + 0.1 g) were weighed,
thensealed into tube-shapedbags, eithermade of a
singlelayer of 0.3mm or a double layerof 5-mm-
meshnylonnet. The double layerappeared effective
in retainingleafparticlesand at thesame timeallow-
ing free movementof macrofauna,includingthe
earthworm Lumbricusterrestris.In narrow-leafedand
needle-leafed specieswheremajor lossesmightoccur
throughthe 5-mm-mesh,only 0.3-mm-meshbags
wereused. Precautionsweretakennot to break air-
driedleaves.Whenflattened, bothtypesof litterbags
had an innercompartment of 10cm x 7 cm. Sam-
ples consistingof 4-cm x 1-cmpieces of Whatman
grade 540 filterpaper, includedfor potentialcom-
parisonwithotherstudiesbecause of its guaranteed
constantquality,were treatedas if theywere litter
samples. For each species,a subsampleof the leaf
575 litterwas weighedair-dryand again after48 h in an
J.H. C. Cornelissen oven at 80 "C, in orderto calculate initialoven-dry
weightsof the materials.All preparedsampleswere
storedair-dryin thesame laboratoryuntiltreatment.
largepetiolesweresuspectedto affect
Since relatively
decompositionrates of whole leaves negatively,the
initialpetioledryweightas a percentageof totalleaf
dryweightwas assessedinat least8 leavesperspecies.
TREATMENTS
The percentagedryweightloss of the littersamples
was assessed in the followingtreatmentsof burial
undernaturalweatherand soil conditions:
W8F = (fine)0.3-mm-mesh bags buriedfor8 winter
weeksfrom24 November1993;
W8C = (coarse) 5-mm-mesh bags buriedfor8 winter
weeksfrom24 November1993;
W20F = 0.3-mm-meshbags buried for 20 winter
weeksfrom24 November1993;
W20C = 5-mm-mesh bags buriedfor20 winterweeks
from24 November1993
S8F = 0.3-mm-mesh bags buriedfor8 summerweeks
from29 July1994.
Replicationper treatment was eightin mostspec-
ies,butfiveto sevenwheretheamountofleafmaterial
was limited.For thesummer(S8F) treatment, stored
materialcollectedin autumn1993was used,as wellas
freshlyshedevergreen leavescollectedduringsummer
1994.This would allow an evaluationof theeffects of
timeoflittercollectingon thepotentialdecomposition
rate.
The samples were buried in a purpose-built
decompositionbed at SheffieldUniversity'sExper-
imentalGardens. The bed measured6 m x 5 m and
was on a 3? slope facing north-northwest, partly
shaded by neighbouring trees.In September1993 all
rootingplantsand thelitterlayerwereremoved.The
top 20cm of the loamy-clayey brownforestsoil, pH
5.93 + 0.34 (measuredin 24 samples,with 1-mdis-
tancesbetweensamples),was mechanicallydug over
and wooden walkwayswere installedlengthwaysat
1--mdistances. On 7 October 1993 leaf-mouldat
differentstagesof decay (mostly1-2 yearsold) was
collectedfroma nearbycompost pile consistingof
litterfromspeciescommonin local cityparks,suchas
Acerpseudoplatanus, Betulapendula,Fagus sylvatica,
Fraxinus excelsior,Pinus sylvestris,Quercus spp.,
Sambucusnigraand Taxus baccata. This leaf-mould
was mixedthoroughly and spreadoverthedecompo-
sitionbed inan 8-10-cm-thick layer.Leaf-mouldsam-
ples contained 1800 + 188 earthworms(Lumbricus
terrestris)m-3 of the length class 3-5 cm and
525 + 125m-3of theclass > 5cm. Twenty-four 1-IM2
plots werelaid out betweenthewalkways,each plot
? 1996British
Ecological Society, assignedat randomto treatment W8 or W20. All13024
JournalofEcology, wintersampleswereremoistened withrainwater,ran-
84, 573-582 domizedwithinW8 or W20, and buriedon thesame
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day (24 November1993),at a depthof 4-5 cm in the
leaf-mould.Additionalcontrolsamplesof 14 species
leaf
(chosen to representa diversityof life-histories,
sizes and leaf shapes) were treatedidentically,but
retrievedimmediately afterburial.Thesewereusedto
assesstheloss ofleafparticlesfromthelitterbags and
the effectsof remoistening (leaching).The relatively
low numberof summer(S8F) sampleswereburiedas
above in one of thecentralplots only.A 3-cm-mesh
nylonnetwas stretched acrossthedecompositionbed
to protectthe samplesfromgreysquirrelsand other
macrofauna.Accumulationsofleaveswereshakenoff
regularly.
Afterretrieval(W8 on 19 January;W20 on 12
April;S8F on 22 September1994),the sampleswere
storedat - 14'C. Afterdefrosting, adheringsoil, soil
faunaor otherextraneousmaterialwas removedfrom
thedecomposedleaflitterbybrushingor swiftly rins-
ing withwater.The macroscopicfauna foundinside
litterbags includedsmallerslugs and snails, centi-
pedes,Collembola,acarid mites,Coleopteranlarvae
and Oligochaetae (includingLumbricusterrestris).
Litter samples were dried for 48 h at 80 'C, then
weighed.Decompositionratewas definedas theper-
centagedryweightloss after8 or 20 weeksof burial.
The effectsof taxonomicrelatednesson the pat-
ternsfoundamongfunctional woodyplanttypeswere
testedfollowingKelly & Beerling(1995), who used
a method derivedfromFelsenstein(1985). First a
phylogenetic treefromspeciesup to class level was
constructedfollowingCronquist(1981). For each set
of two or moretaxa belongingto the same taxon of
the nextlevel up a contrastwas taken (e.g. is litter
weightloss higherin life-form A thanin life-formB?),
afterwhichthe species used were taken out of the
analysisto ensureindependenceof thedifferent con-
trasts.Whereall membersof a taxonbelongedto the
samecategory,themeanvalue forthesememberswas
used fora contrastat thenexttaxonomiclevelup.
Results
WEATHER
Data fortheexperimental period(fromWestonPark
MeteorologicalStation,Sheffield,sited800m north-
northeastof decompositionbed; details not given
here)showedthatprecipitation and temperaturewere
broadlysimilarto thoseforthepast 10years.Rainfall
was substantialin all experimentalmonths,so that
thedecompositionbedwas neversubjectedto drought
duringtreatment. Snow coveredthe bed duringtwo
periodsof about a week.Onlymildfrostoccurred.
DECOMPOSITION RATES
forindi-
Mean percentageweightloss was determined
(Appendix2, WWW
vidual speciesin each treatment
archive).Withineach treatment valuesweregrouped
576 into classes (class 1 = very slow, 2 = slow, 3 = inter- Table 2 Mean (? SE) initialpercentagedryweightloss until
Leaf mediate,4 = fast,5 = veryfast)such thatall classes and includingburial
decomposition containedsimilarnumbersof species. Mean weight
Mesh size
rates loss class foreach specieswas thencalculatedfrom
the values forthe fivetreatments. Variabilityin dry 0.3mm 5mm
weightloss among replicateswas generallylow and
no correctionforpositionwithinthedecomposition Acerpseudoplatanus 4,6 + 0,28 4.7 + 0.46
Alnusglutinosa 4.1 + 0.30 4.6 + 0.24
bed was necessary.Relative dry weight losses of
Betulapendula 1.6 + 0.073 2.1 + 0.31
laminae only was similarto that of whole leaves in Bromuserectus 0.6 + 0.30 1.3 + 0.37
all specieswithcompound leaves (Table 1), both in Buddlejadavidii 1.0 + 0.23 1.7 + 0.17
treatments W8F and W20F. Carpinusbetulus 2.2 + 0.15 2.9 + 0.16
In 0.3-mm-mesh bags relativeweightlossesamong Digitalispurpurea 10.6 + 1.85 9.9 + 1.72
Epilobiumhirsutum 3.6 + 0.87 4.6 + 0.82
121 species after8 winterweeks (W8F) were cor-
Fagus sylvatica 0.4 + 0.11 0.2 + 0.049
relatedstronglywith losses after20 weeks (W20F) Festucaovina 0.8 + 0.072
(r = 0.96, n = 121, P < 0.001). In 5-mm-meshbags Fraxinusexcelsior 1.0 + 0.23 1.8 + 0.11
(W8C vs. W20C) thecorrelationbetweentreatments Larix decidua 0.03 + 0.028
was less strong(r = 0.82,n = 75, P < 0.001),because Juniperus communis 0.2 + 0.23
Plantagolanceolata 2.5 + 0.68 3.6 + 0.50
in severalspeciesmaximumweightloss was reached
before20 weeks.Relativedryweightlossesin thetwo Resultsoftwo-wayANOVA: mesh-sizeeffect F, = 3.42,NS;
types of bags after8 weeks (W8F vs. W8C) were specieseffectF13= 38.9, P < 0.001; interactionFlo = 0.36,
closelycorrelated(r = 0.93, n = 75, P < 0.001), but NS.
after20 weeks(W8C vs. W20C) the correlationwas
again less (r = 0.86, n = 75, P < 0.001), owing to
near-maximum weightlosses in 5-mm-mesh bags by suggeststhatmacrofaunahad notconsumedanyfilter
20 weeks of burial. Ten species had verylow and paper and thatenvironmental conditions(e.g. moist-
similarrelativeweightlosses in 0.3- and 5-mm-mesh. urecontent)had been similarformicro-organisms in
Species-burial-periodand species-mesh-sizeinter- bothmeshtypes.
actions were evidentbetweenindividualspecies but Relativeweightlossesin S8F correlatedwiththose
did not obscurethegeneralpatterns. in W8F (r = 0.91, P < 0.001, not shown)and W20F
(r = 0.95, P < 0.001, Fig. 1). These data indicatethat
theeffects of burialseason,althoughpossiblyimpor-
CONTROL TREATMENTS AND COMPARISONS
tantwhereindividualspeciesare compared,did not
Initialrelativedryweightloss untiland includingthe affectthe rankingof specieswithregardto relative
momentof burialvariedsignificantly withina subset weightloss.
of 14species(Table 2). Therewas no overalldifference In a subset of threeevergreenspecies (Table 3),
between0.3 and 5-mm-mesh, which indicatesthat only one (Hypericum calycinum) showed a significant
initialloss from5-mm-mesh was negligible(as it was difference in relativeweightloss betweenleafmaterial
assumedto be from0.3-mm-mesh) and initialweight collectedin autumn 1993 againstsummer1994,but
loss was due presumablyto initialleachingof water- this was not enough to take it into anotherweight
solubleorganicmaterial(cf.Nykvist1963). loss class. In addition,relativeweightlosses of leaves
There was no significantdifferencein relative of Prunus laurocerasus, Taxus baccata and Vaccinium
weightloss of Whatmanfilter paperbetween0.3- and vitis-idaea,collectedin autumn 1993 and buriedin
5-mm-meshin eitherW8 or W20 treatments.This winteras againstthosecollectedand buriedin sum-

Table 1 Mean percentagelitterweightloss in compoundleaves: laminaevs. whole leaves withineach of two treatments.
All
meanvalues are based on at leastfivereplicates

W8F W20F

petioles laminae leaves petioles laminae leaves

Aesculushippocastanum 17.5 14.7 15.2 33.1 23.9 25.4

Fraxinusexcelsior 32.8 33.0 32.9 51.3 55.0 54.5
Laburnumanagyroides 30.5 45.8 43.4 52.0 63.8 62.0
Rosa arvensis 18.5 23.1 22.6 38.4 37.0 37.2
Rosa canina 21.8 25.8 25.3 45.3 47.0 46.7
Rubusfruticosus 22.9 16.8 17.8 42.3 36.4 37.4
? 1996British Rubusidaeus 33.5 29.4 29.9 61.2 61.8 61.7
Ecological Society, Sambucusnigra 47.4 48.6 48.4 63.6 76.0 73.9
JournalofEcology, Sorbusaucuparia 24.7 26.3 25.9 48.2 44.7 45.4
84, 573-582

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 70
577 PLANT TYPES AND DECOMPOSITION RATES
J.H. C. Cornelissen 60 r 0.95
Unless otherwisespecified,resultsindicate weight
50
lossesin0.3-mm-mesh, becauseoftheessentially simi-
40 *
lar patternsin 0.3- and 5-mm-meshand the larger
numberof speciestreatedin 0.3-mm-mesh.
Therewas significant overallheterogeneity inmean
20
litterweightloss amongninegroupsof species differ-
10
ing in life-form or leaf habit (Fig. 2, Table 5). The
patternwas the same afterburialfor8 or 20 weeks,
0 10 20 30 40 50 60 70 80 as shownbytheabsenceofan interaction effect(Table
W22F % weightloss in
5). Withintheherbaceous(nonwoody)species,dicots
Fig.w1Relative dry weightlosses in W20F vs. S8F. Open
decomposedon averagefasterthangraminoidmon-
circlesindicatethatthesame leafcollectionwas used in both
treatments. Solid squaresindicatethatleavesin W20F were ocots. Withinthe woody species,deciduous species
collectedin autumn1993and leavesin S8F in summer1994, decomposed twice as fast as evergreens(28.4 + 1.5
bothfromsimilarsites. vs. 12.9 + 1.6% weightloss in W8F; 48.2 + 2.3 vs.
24.7 + 1.0% in W20F). These resultswereconfirmed
whentaxonomicrelatednesswas takeninto account
(Table 6). Life-form was a significant sourceof vari-
Table 3 Mean (? SE) percentagedryweightloss in S8F in
ation in decomposition rate in deciduous woody
evergreenscollectedfromsimilarsitesin autumn 1993 vs.
summer1994.Quercusilexleaveswerecollectedfromunder plants, but not in evergreens (Table 5). Within the
thesame treein bothperiods deciduous woody species, climbers and scramblers
decomposed faster than self-supportingspecies
Collected
(48.0 + 7.2 vs. 27.4 + 1.4% weight loss in W8F;
autumn1993 summer1994 76.1 + 2.3 vs. 46.8 + 2.2% in W20F). Owing to the
small numberof comparisons,thiscould not yetbe
Quercusilex 12.64 + 0.40 11.46 + 0.69 supportedbya taxonomicrelatednessanalysis(Table
Hedera helix 38.68 + 0.74 36.74 + 1.70 6). The latterdid demonstratethatleaf litterof sub-
Hypericumcalycinum 10.62 + 1.11 ** 15.01 + 0.70
shrubsgenerallydecomposedmore slowlythan that
**P < 0.01 (t-test).
of shrubsand trees.
Relativeweightloss in the entirespecies set was
correlated with initial SLA of the litter (W8F:
r = 0.45, P < 0.001, W20F: r = 0.44, P < 0.001).
mer 1994, were close to the fittedregressionline in This was due mostlyto the woody species (W8F:
Fig. 1. These data indicatethatrelativeweightlosses
of leaves of the summer-shedding evergreensin this 100
*W8F
studyweregenerallysimilarwhethercollectedfreshly
90 - W20F
fallenin summeror,withsome delay,in autumn.
In a subsetof threedeciduouswoody species,col- 80

lectedfromcontrastedsites,two revealedsignificant 70
intraspecificvariabilityin relativeweightloss (Table
4), unrelatedto the degreeof exposureof the site.
0 60 T
Althoughdata on only threespecies do not justify *F3 50

conclusions,this intraspecific variabilitywas much 40

smallerthanthevariabilityamongdeciduouswoody
30T
species(cf.Appendix2).
20

10 o~~~~~~~
Table4 Mean (? SE) percentagedry weightloss in S8F
sites
withindeciduouswoodyspeciescollectedincontrasting u ~ 0 u 0 u ~ U

in autumn1993 .
(D
0 -o C.) 0
0~~E
- o -C E
C0
0
Site o

exposed sheltered
in
dryweightlossesin 0.3-mm-mesh
Fig.2 Mean relative
Cornussanguinea 62.79 + 1.41 69.63 + 4.59 habit.Woody
groups of specieswithsimilarlife-form/leaf
G 1996 British Fraxinusexcelsior 47.42 + 1.25 ** 53.87 ? 0.86 climbers= woody andscramblers.
climbers D = deciduous,
Ecological Society, Viburnum opulus 55.77 + 1.43 * 50.86 + 1.00 from
Meanvalueswerecalculated
E = evergreen. themean
JournalofEcology, percentage species.Standard
weightlossesof individual
84, 573-582 *P < 0.05, **P < 0.001 (t-tests). errorbarsareshownone-sided.

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578 Table 5 Summaryoftwo-factor ANOVAS withfactorsfunctional
group(life-form/leaf
habit,see Fig. 2) and burialperiod(W8F
Leaf vs. W20F). All data werenormalizedthroughln-transformation
priorto analyses
decomposition p
rates
Functional Burial Total
Functionalgroup group period Interaction d.f.

Amongall life-form and leafhabitgroups < 0.001 < 0.001 < 0.99 237
Graminoidmonocotsvs. herbaceousdicots < 0.001 < 0.001 < 0.62 71
Amongall deciduouswoodylife-forms < 0.001 < 0.001 < 0.97 121
Amongall evergreen woodylife-forms < 0.030 < 0.001 < 1.00 43
Woody species:deciduousvs. evergreen < 0.001 < 0.001 < 0.33 167
Deciduous woodyspecies:climbers/scramblers < 0.001 < 0.01 < 0.86 121
vs. self-supporting
plants

Table 6 Analysisof relationships

withinthewoodyspeciesbetweenlitterweightloss and functionalgrouptakingintoaccount
taxonomicrelatedness.In each case thenullhypothesisstatesthat,across all contrasts,mean litterweightloss class is higher
in thefirstfunctionalgroupno moreoftenthanit is in the second group.In thecase of SLA thenull hypothesisstatesthat
therelationshipwithlitterweightloss shows,across all contrasts,a positivetrendno moreoftenthana negativeone

No. ofcontrasts/
Functionalgroups No. of disagreements x2, p

Deciduous climbers/scramblersvs. otherdeciduouswoodyspecies 3/3 2.00 NS

Subshrubsvs. shrubs/trees(deciduousand evergreen habitsin separatecontrasts) 7/7 4.67 < 0.05
Deciduous vs. evergreenhabit 15/15 10.0 < 0.01
High SLA vs. low SLA 27/17 0.92 NS
Autumnleafcolorationgreenor yellow-green vs. less thanhalfgreen 11/10 7.71 < 0.01

r = 0.61, P < 0.001; W20F: r = 0.54, P < 0.001). indicatedby thelack of interactioneffects (Table 7).
This relationshipwas, however,not confirmedby a The most strikingdifference in relativeweightloss
taxonomicrelatednessanalysis(Table 6). Therewas was betweenleaflitterwithvs. without(partial)green
no significant correlationbetweenSLA and weight coloration, a differenceconfirmedby taxonomic
loss withintheherbaceousdicots(W8F: r = 0.41,NS, relatednessanalysis(Table 6). Multicolouredleaves
W20F: r = 0.33, NS) or graminoidmonocots(W8F: (i.e. withmixturesof at least red,yellowand green)
r = 0.22, NS, W20F: r = 0.25, NS). wereintermediate in relativeweightloss. The results
Withinthe deciduous woody species, significant for brown leaves should be interpreted with some
heterogeneity in relativeweightloss could be explai- caution,sincethisgroupconsistsof threespeciesof
ned byleafcolorationat thetimeofshedding(Fig. 3). Fagaceae only.
The patternswerenot significantly affectedbyperiod
of burial (8 vs. 20 winterweeks) or mesh size, as
PLANT FAMILIES AND DECOMPOSITION RATES

Mean weight loss class was significantly

hetero-
80 geneous among the 10 main plant familiesin this
70 T
0 60 EW8F

50 W8C Table 7 Summary

of three-way
ANOVA withfactors
leaf
coloration(see Fig. 3), burial period (8 vs. 20 weeks) and
40T
meshsize(0.3vs.5mm).Thedatawerenormalized through
30
ln-transformation Totald.f.= 215
priortoanalysis.
E 20

10
Sourceofvariability P
0

Colour < 0.001

Burialperiod < 0.001
Meshsize < 0.001
Fig.3 Mean relativedryweightlossesafter8 weeksingroups Colourx burialperiod < 0.36
? 1996 British ofspecieswithsimilarleafcolorationat thetimeofshedding. Colourx meshsize < 0.21
Ecological Society, Mean values were calculated from the mean percentage Burialperiodx meshsize < 0.22
Journalof Ecology, weightlosses of individualspecies.Standarderrorbars are Colour x burialperiodx meshsize < 0.44
84, 573-582 shownone-sided.

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All use subject to JSTOR Terms and Conditions
579 study (Fig. 4, Kruskal-Wallis nonparametrictest, and tanninsare known to act against folivoresin
J.H.C. Cornelissen %2 = 37.7, P < 0.001). The variabilitydoes not just livingplants(Coley 1983;Kuiters1990). The present
reflectdifferencesbetweenwoody and herbaceous data, whenlinkedto initiallignin:N ratiosand lignin
taxa; slowly decomposingFagaceae and Ericaceae contentsof leaf littersfound by previous authors,
and fastSalicaceae and Caprifoliaceaeare all woody, provideadditionalevidenceoftheimportanceoflitter
whereasall-herbaceoustaxa includemoderatelyslow qualityacross a rangeof woody species.Despite the
Poaceae as wellas fastAsteraceae. diversedata sources and analyticalmethodsused,
Mean weightloss classesofthedicotplantfamilies lignin:N ratio (r = -0.78, n = 12, P < 0.01, Fig. 6)
in the experiment werelooselyassociatedwiththese and lignincontent(r = -0.66, n = 14,P < 0.01) were
families'evolutionaryadvancement(ranging from bothnegatively correlatedwithlittermeanweightloss
primitiveto advanced)as quantifiedin thePercentage class in thisstudy.Notwithstanding theemphasison
AdvancementIndex (Sporne 1980; Sporne 1982) litterquality,it is recognizedherethatthereare sig-
(Fig. 5, r = 0.40, P < 0.05). nificantinteractions of litterqualitywithdurationof
decomposition(cf. Mommaerts-Billiet 1971; Wieder
Discussion & Lang 1982; Berg & Ekbohm 1991) and with
environmental factors(Swiftet al. 1979). The latter
EFFECTS OF LITTER QUALITY AND
includefeaturesof thelitterlayerand itsdecomposer
ENVIRONMENT
community (Bocock et al. 1960;Heal & Ineson 1984;
A broad body of literaturehas shownthatvariation Buth & De Wolf 1985; Elliot et al. 1993), mac-
in leaf decompositionrates among species depends roclimate (Meentemeyer1978) and microclimate
greatly on litter (resource) quality. Lignin, for (Escuderoet al. 1987;Taylor& Parkinson1988).The
instance,may enhancephysicaltoughnessof leaves, methodologyadopted, for instancelitterbag type,
may affectsome of the above environmental factors
5 (Anderson1975; Louisier& Parkinson1976). All of
theseinteractions may be relativelyimportantas far
as relativeweightlossesofindividualspeciesare con-
cerned.The presentresultshave shown,however,that
n)3
interactive effectsof litterqualityand burialperiod,
season or litterbag type,althoughconsiderable,were
not so greatthat theyalteredthe broad rankingof
E T T
125 specieswithregardto litterweightloss. Onlythe
near-maximum weightloss in a large proportionof
thespeciesin 5-mm-mesh after20 weekscaused pro-
E
a)
t
a)
$1
a)
1r
a)
0
a)
l
a) a)
|
a)
|0
a) a)
Loi nounced deviationsfromlinearitybetweenrelative
weightlosses in different treatments,butevenin that
U) 0
U-~~~~U
case did not fundamentally altertherankingof spec-
Fig.4 Mean weightloss class (averagedover thefivetreat- ies. Possible interactionsbetweenqualityof the leaf
ments)forgroupsof speciesbelongingto the same family. samplesand featuresof the surrounding leaf-mould
Onlyfamilieswithat least two generaand fourspeciesrep- layerhave not been assessedin thisstudy.However,
resentedin thisstudyare used. Mean classeswerecalculated
themixedleaf-mouldin thedecompositionbed had a
fromthemean weightloss classes of the individualspecies
in each family.Standarderrorbars are shownone-sided.

40

r=-O78, P'0 01
30
u' 4 *.

(n
20
n* E20-
o 3

E
10

C * * r=O.40, P<O.05

E
0 1 2 3 4 5
lIgninN ratio
0
30 40 50 60 70 80
Fig.6 Relationship
between initiallignin N ratioof leaf
litter
(fromtheliterature)andmeanlitter weight lossclass
Sporne's percentage advancement index (thisstudy)in 12 woodyspecies.Data sources:King&
Fig.5 Mean weightloss class of dicotyledonousfamilies Heath(1967);Healetal. (1978);Chauvet (1987);Hendriksen
? 1996British againsttheirSporne'sPercentage Advancement Index.All (1990);Berg& Ekbohm (1991);Slapokas& Granhall (1991);
Ecological Society, dicotfamiliesincludedin thisstudyarerepresented.
Mean Van Vuuren(1992);Tietema(1993);Cotrufo et al. (1994);
JournalofEcology, classeswerecalculatedfromthemeanweight lossclassesof Domenachet al. (1994).Themeanlignin:N ratiowascal-
84, 573-582 theindividualspeciesineachfamily. culatedwhereonespeciesappearedintwopapers.

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All use subject to JSTOR Terms and Conditions
580 good representation of both acidic substrates(e.g.
Leaf beech,oak, pine,yew),as favouredby fungiand cer-
decomposition tain invertebrategroups, and more base-richsub-
rates strates(e.g. ash, alder,elder),supporting bacteriaand
otherinvertebrate groups(cf.Swiftet al. 1979).It can
therefore be assumedthatthedecomposercommunity
in thisleaf-mouldis to some degreerepresentative of
thatinthenaturalhabitatsofmostspeciesin thisstudy.
In orderto explainthevariationin leafdecompo-
sitionratesfoundamongspecies,manypreviousstud-
ies have scaled down fromthe leaf level to its indi-
vidual structuraland biochemicalcomponents:the
mechanisticapproach. In contrast,the presentmul-
tispeciesscreeningis an experimental attemptto scale
up fromtheleaflevelto thelevelof wholeplantsand
plant types:the functionalapproach. In supportof
thehypothesis, theresultsindicatethatrelativeweight
losses of leaf litterdo indeedreflect, to some extent,
featuresthatenhancethefunctioning of livingplants
and theirleaves in theirnaturalenvironments. For
instance,ecological strategymay explain the fast
decompositionof deciduous woody climbersand
scramblers in thisstudy.Comparedto self-supporting
woodyplants,theyare seentypically inheterogeneous
lightenvironments, whichpromotea highleaf turn-
over as determinedby thechangeableavailabilityof
lightpatches(Castellanos 1992). In suchplantsnatu-
ral selectionmay have favouredshort-livedleaves
equippedforefficient photo-assimilation witha mini-
mumof defencechemistry (cf. Chapin 1980; Chabot
& Hicks 1982; Coley 1988). Such leaves would be
palatable to soil-bornedecomposers(Grime& And-
erson 1986). Subshrubs, on the other hand, are
exposed to herbivoryby mammals (e.g. rabbits,
sheep) and leaves of such plantscan be expectedto
have developed antibrowserdefences,which could
also resultin less palatable litter.Indeed, leaf litter
weightlosses were consistentlylower in subshrubs
thanin shrubsand trees,plantsin whichmostof the
foliageis out of reachof mammals.
The negativerelationshipbetweenleaf life-span
and decompositionrate, implied above, was more
directlydemonstratedby theconsistently smalllitter
weightlosses of woody evergreensas compared to
deciduous species. The latterrelationshiphas been
supportedby some earlierevidence,as reviewedby
Aerts (1995), but has never previouslybeen dem-
onstratedacrossa representative rangeofwoodylife-
formsand (bothbroad-leafedand needle-leafed) taxa.
The relationshipheld for a big slice of the woody
Sheffield flora,whetheror not taxonomicrelatedness
was accountedfor.
Long-livedleaves,withrelatively largeinvestments
in compoundsnot directlyinvolvedin photo-assimi-
lation,generallyexhibita low specificleafarea (Reich
et al. 1992). At least among thewoody species,SLA
? 1996British
Ecological Society, of litterwas closely correlatedwith that of living
JournalofEcology, leaves (J. H. C. Cornelissen,unpublisheddata). The
84, 573-582 positivecorrelationbetweenlitterSLA and weight
This content downloaded from 2.103.241.231 on Mon, 16 Jun 2014 21:21:19 PM
All use subject to JSTOR Terms and Conditions
loss in thewoody speciesin thisstudytherefore sug-
gestsa link betweenleaf functioning and decompo-
sition. Mechanistically,this relationshipmay be
explainedbyvariationin structural chemistry.Leaves
witha smallspecificarea can be expectedto be physi-
callytough(in termsof resistanceto penetration), as
was demonstratedin tropicalAsian (Choong et al.
1992)and Amazoniantreespecies(Reich et al. 1991).
Indeed,leaftoughnesswas a good negativeindicator
of decompositionratein nineMediterraneanwoody
species (Gallardo & Merino 1993). However, the
relationshipbetweenSLA and decompositionrate
can, at thisstage,not be extrapolatedto otherfloras,
since it was confounded with taxonomy. It also
remainsto be studiedto whatextenttheSLA ofliving
herbaceousleavestranslatesintothatofthelitter, and
why the relationshipbetween litterSLA and de-
composition ratewasnotseenamongherbaceous species.
The autumnleafcoloursin deciduouswoodyspec-
ies may also in partreflectfunctionalfeaturesof the
living leaves. Brown coloration,for instance,rep-
resentsmostlyphenoliccompounds such as lignins
and tanninsthatbecomeapparentin senescedleaves
once thegreenpigmentshave been transformed into
colourlesscompounds.Indeed, in this study,leaves
that were brown or yellow-brownupon shedding
were broken down comparativelyslowly. In most
deciduous species the greenchlorophyllsand, to a
lesserextent,yellowcarotenoidsthatoccurin assimi-
latingleaves, are substantiallydegradedbeforeleaf
fall(Hendryet al. 1987). Severalspeciesin thisstudy,
however,shedtheirleaveswhilestill(partially)green.
WiththeexceptionofHippophae(which,interestingly
in this context,is a nitrogenfixer),all eightnative
species that shed theirleaves predominantly green
occurin theSheffield area on (moderately)productive
soilsofmediumto highpH. On suchsoils,fastgrowth
and competitive vigourmaybe ofmoreadaptivevalue
than stress-tolerance(sensuGrime 1974),and leaves
can be expectedto be photosynthetically productive
withlow contentsof protectivesecondaryand struc-
tural compounds. In such leaves it mightpay off,
ratherthangraduallytransforming and withdrawing
theirphotosynthetic chemistry - a processmarkedby
colour change- insteadcontinuingto produce new
photosynthates up to themomentof leafabscission.
Such green,poorlyprotectedleaves,presumably with
high contents of photosynthesis-related nutrients,
would be palatable to decomposers. This would
explainin functionaltermswhygreenautumncolor-
ationis an indicatorof fastleafweightloss in decidu-
ous woody species,independentof taxonomy.Fur-
ther study is needed to test whether the fast
decompositionof shed greenleaves is a consequence
of thelack ofinvestment in compoundsthatpromote
stress-toleranceor of highernutritional value. It also
remains to be determinedwhethercoloration of
senescedleavescan be used to predictdecomposition
ratein herbaceousplants.Leaf colorationcan change
581 rapidlyduringsenescence(Sanger 1971)and thismay
J.H.C. Cornelissen make it difficult to standardizethecolour recording
in themanyherbaceousspeciesthatdo not shedtheir
senescedleaves.
Whilstmost of the functionalgroups of species
in thisstudycompriseda varietyof taxa, significant
variabilityin leaf decompositionrate was also cor-
relatedwithtaxonomy.Planttaxa are probablyto an
important extenttheoutcomeofmultipleadaptations
to the environment. Some primitivetraitsthat have
remainedin higherplanttaxa are probablystilladapt-
ive in the species' presentenvironmentsand may
thereforepartiallycoincide with the traitsof func-
tional typesin this study(cf. Westobyet al. 1995).
EvolutionaryadvancementsensuSporne (1980), for
example, has been shown previouslyto correlate
withecological strategy.Families witha highmean
Sporne's PercentageAdvancementIndex are com-
parativelysuccessfulin disturbedand productive
habitats(Hodgson 1986;Grime1986).It can therefore
be expectedthat'primitive'familieswitha low index
are moresuccessfulin environmentally stressedhabi-
tats. Since the leaves of plants in such habitatscan
be expectedto be well protectedand theirlitterto
decompose relativelyslowly (Grime & Anderson
1986),thepositivecorrelationbetweenSporne'sPer-
centageAdvancementIndex and relativeweightloss
in dicotfamiliesmaybe an evolutionary as wellas an
ecologicalrelationship.
Life-form, leaf habit,autumnfoliagecoloration,
taxonomyand evolutionaryadvancementindexcon-
stitutespeciesfeatureswhichareeithereasyto observe
in thefieldor whichcan be obtainedfromthelitera-
ture.In viewof theircorrelationswithleafdecompo-
sitionrate,thesefeaturesmay proveusefultools for
predictingdecompositionratesin vegetationsdiffer-
ingin speciescomposition.The resultsmayalso have
implications for ecosystem modelling. Anderson
(1991) predictedthat in temperateareas a future
amelioration of the climate would enhance the
importanceof the biologicalcontrolover decompo-
sitionprocesses.He arguedthatshiftsin speciescom-
positionin existingbiomeswould have major effects
on decompositionin thenearfuture,and shiftsin the
distributionof the biomes themselvesin the more
distantfuture.The presentresultsmay be a firststep
towards predictingincreases or decreases in litter
decompositionrates (and carbon turn-over)under
differentvegetations fromthepotentialleafdecompo-
sitionratesoftheircomponentspecies.It mayalso be
possible to inferpotentialdecompositionrates for
speciesnot includedin this studyfromsome of the
functionaland taxonomicfeaturesshownhereto be
correlatedwithdecompositionrate.
c 1996British Acknowledgements
Ecological Society,
JournalofEcology, I thankGeorgeHendryforhisacademicand physical
84, 573-582 contributionsduringvarious phases of the experi-
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All use subject to JSTOR Terms and Conditions
ment;JohnHodgson,Ken Thompsonand Sue Hillier
forprovidingherbaceousleafmaterial;StuartBand,
Suzanne Hubbard,Debbie Cornelissenforassistance
in thelab; JonathanKieltyforidentifyingsoil invert-
ebrates. Alastair Fitter, Philip Grime, Bill Heal,
GeorgeHendry,Ken Thompsonand twoanonymous
refereesgave valuable commentson an earlierdraft
of themanuscript.The authorwas sponsoredby the
European Union throughits Human Capital and
MobilityProgramme,whilsthe also benefitedfrom
the long term support of UCPE by the Natural
Environment ResearchCouncil.
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Received24 August1995
revisedversionaccepted15 April1996
J. H. C. Cornelissen(1996). Leaf decompositionratesappendix I

Characteristics ofthestudyspeciesandtheirleaflitter.The status(N = native,Nd = naturalised,

P = planted.)is
according to Stace(1991). * Leaves collectedinUniversity's
Experimental Gardenorprivategardenin
Sheffield.Life-form/leafhabit(following Stace 1991):T = tree,S = shrub(max.heightbetweenIm and 10 m),
SS = subshrub (max.heightlessthan1 m),C = woodyclimberor scrambler, HM = herbaceous(graminoid)
monocot, HD = herbaceousdicot.d = deciduous,se = semi-evergreen, e = evergreen.
Mean relativepetioledry
weightis expressedas % ofwhole-leaf dryweight.n.m.= notmeasured.Classesofpredominant initialleaf
coloration(deciduouswoodyspeciesonly):B = brown,BB = blackishbrown,BY = brown-yellow orB & Y
mixture,Y = yellow,YR = orangeorY & R mixture, YG = yellow-green orY & G mixture,G = green,MC =
multi-colour(mixture ofat leastyellow,redandgreen).

Life- Litter Litter

form %petiole
seiila nta
Woodyspecies Family and
Native/naturalised/planted dry speii leafinta
leaf weight (mm/mg) colour
habit
it
Acercampestrej[Aceraceae [N ](Td [9 ]22.8 IIYR
platanoides[Aceraceae
Acer [Nd ][Td 15 ]j18.8 ][MC
Acer d 16
[Aceraceae [Nd 25.9 YG
Aseudoplatanus [ 1
Aesculus P/Nd Td 17 135.3 BY
Hippocastanaceae
hippocastanum 11 I 11 __1 11_1 _

Alnusglutinosa I[Betulaceae |N ]|Td J[6 21.3 JIG

Amelanchier [Rosaceae [*Nd Td 8 21.1 [YR
lam arckii__ _ _ _ _ _ _ I.] d__ _ _ _ __ _ _ _ _ _ _ _ _

Andromeda [Ericaceae |*N SSe 2 7.9

polifolia _ _ _ _ _ _ _ _ [] Se ~ _ _ _ _ _ _ _ _

unedo
Arbutus ricaceae II *N jISe jI3 ZI7.3 ______
Aucubajaponia !Comnaceae lP ISe 114 1118.6 1!
vulgarisBerberidaceae IN?
Berberis 6Sd 6 2.3 MC
etula
nana Betulaceae *N SSd 4 16.7 IBY
Betulapendula |Betulaceae || |d 19 ||16.1 1Y 1
|uddlejadavidi |Buddlejaceae ||d |Sd |1 |11.4 IMC
3uxus i|Buxaceae ]S*N S3 15.7 ||.
sempervirens .1 11 H
Calluna1vulg icaceae SSe 10.8
betulusBetulaceae
Carpinus IN IIId I [ ] 21 G
Castaneasativa ||agaceae |P(Nd) ||d 15 |28.3 IB

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Corylusavellana ||Betulaceae JjN [[Td 114 1130.8 IIY_ I
Cotoneaster Rosaceae *Nd FSSd F6 14 YR
horizontalis 1111111111
Crataegus 1Rosaceae N [Sd [9 20.4 IMC
monogyna 11 I. [1 11 =
Cytisusscoparius ]|Fabaceae J|N [Se J[0 ||n.m. I___
DryasoctopetalaJ[Rosaceae J[*N |SSe 16 13.4 ||
Empetrum
n_i_gT_____um
||Empetraceae |N SS 6 18.4 1_
Ericacinerea j[Ericaceae S[N I[Sse J16 112.8 _____

Buonymus
europaeus
[Celastraceae
11
N
1
Sd 6 20.4 MC
I_ _ 11 11 11
Fagussylvatica |[Fagaceae J[N I[Td J[5 [9.1 ][BY
Ficuscarica J[Moraceae I*Nd j[Td I1 ][24.6 _IG

rxcelsior [Oleaceae [N [Td |14 ]|16.6 __G_

_

FFuchsia
chsia[ [Onagraceae [*P/Nd 1111
[Sd 7 ][27.1 1
]{MC
magellanica 1111111111
Hebex 0
Scrophulariaceae*Nd [Se 111.3
franciscana 1111 11 11
Hederahelix [Araliaceae I|N |Ce (14 ]|14.4
[elianthemum Cistaceae [N ]|SSe 7 120.7 ||
nummularium 111IL 1111
11
Hippophae
Ihppophae
rhamnoydes
'[Ele
ir
Elaeagnaceae [*N i
][Sd 3 ]1
17.9 IIG
Hypericum 11
calycinum
Hypericaceae 1*P/Nd
11- 11111111 J[SSe [o 112.7 11._
Ilex aquifolium ][Aquifoliaceae j[N ]Te _
__5 ]15.7 _____

Juniperus j[Cupressaceae [*N ISe [o 113 1___

Laburnum IFabaceae I*P/Nd ][Td [15 127.1 IIYG
anagyroides 111111111
Larixdecidua j[Pinaceae IIP |]Td I[0 1115.5 ]IYR
Larixkaempferij[Pinaceae j[*p ][Td 1[o ]120.4 ]IYR
Ligustrum
ovalifolium
OlOeaceae
11111
IIP __Se 13 ]4.7 ||.
.1111

Lgustrum |Oleaceae ||N Sd 1331.3

1R
G__

'LOniCera Caprifoliacac NIM ld j4S

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MalussylvestrisRosaceae jN Td 9 [ |23.3 MC
Piceasitchensis Pinaceae p Te Z.71
Pinussylvestris|Pinaceae |N|Te 11 15.61
Populusigra Salicaceae |N/P d 12 13.4 Y
Populustremula|Salicaceae |N|Td |13 |16 |YG I
Prunusavium |Rosaceae | Id 18 120.1 iYR
'runus
laurocerasus 11
jRosaceae
IK!Z
*Nd Se 18.7 1|. __
Prunuslusitanica|Rosaceae |*P(Nd) |Te |6 ||.8 .
runuspadus |Rosaceae |N|Td |12 |43.2IYI
'runusspinos Rosaceae Sd | Z6 | U7.7 IIY I
yruspyraste Rosaceae Nd Td I0 14.5 IYI
ilex
Quercus !Fagaceae 1*P/Nd IIe 17 10.O8 -
u1erCUs
Petraea FagaCeaeN Td 6 149 B
|| |d 14 115.7 |IBI
]______
robur |Fagaceae
|Quercus
Rhamnaceae N Sd 1 38.5
cahamnius
tharticus IN]s [1]____ 1IG ___

;hododendron j[Ericaceae [*Nd ][Se J59 [8 || _

I
ponticum 11 11 1111111
Ribesalpinum
|Grossulariaceae|N Sd 181121[2
LIIMc I
Ribesfnigrum Grossulariaceae*Nd? iSd 19 1Z3
|R.besrubrum |Grossulariaceae||d |Sd |13 |28 l
Ribesuva-crispaIGrossulariaceaeI*N? {Sd 118 120.2 !I
Rosaarvensis !|osaceae 1 |Sd |12 |29.3IY
||osa canina |Rosaceae |N|Sd 113 1|5.9 IMC I
|Rubusfruticosus
||osaceae |NISCse 18 123.1 1
Rubusidaeus Rosaceae IN Sd 11 30.4 y
|Salixalba |Salicaceae || |d 14 114.9 |GI
Saixcaprea |Salicaceae |N|Td 17 ||15.9 |M
Salix
fragilis Salicaceae |N lTd E15ZIiIY
111 I
Sambucusnigra CaprifoliaceaelN ]ISd 117 125.8 I

dulcamara [Solanaceae j[N SCd ji 1 [43.6 IxY

Sorbus
aria Rosaceae *N I|Td Z7 15.4 By
orbusaucupa |saceae Td II22 [
X17.4 ||YR

Carifoliaceae
ISymphoricarpos *Nd Sd 3 I

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 gallica Tamaricaceae *P(Nd)
Tamarix n.m.IYG
axusbaccata Taxaceae Te 0 6.5 ____

)olytrichus Lamiaceae!N SSe 12 !3.

Tiliacordata iliaceaeN Td 10 24.3 BY
Tiliaceae |(N)/P
Tiliaxvulgaris lTd 112 IEmI BY
Ulexeuropaeus abaceae |N Se || Z XM.I
Ulexgallii Fabaceae N Se 1EinImI1ZIZ
Ulmusglabra Ulmaceae ___ |Td ||2 ||29.8 |IY
_

accinium Ericaceae ] SSd 4_1 23.4 YR

vitis- Ericaceae
Vaccinium SSe 3 7.6
idaea i 11 1I
ICaprifoliaceae ]
Viburnum_lantana ]ISd 1]7 19 __G_ 11
ViburnumopulusIlCaprifoliaceae ]uN ISd 1__
_9 7.9
Herbaceous
species
capi11arisPoaceae
Agrostis ]uN ]IHM ]1o ]131.2
1
Anthoxanthum Poaceae iN HM 0 31.7
odoratum _______ ___________]___ ____ _____ ___

|Alvsthriscs Apiaceae ][N ] HD ][n.m. 33.4

Aylhenaterum 11
Arrhenaterum Poaceae
elatius
][N ]HM
1
0|o36.1
11----- -------- -

pinnatum |Poaceae ]1 ]
i ]_39 ___

rizamedia IPoaceae ][N ]uHM 10 1[22 11 _

I
BromopsiserectaIPoaceae ][N ]uHM [0 1|20.9 11 _
1
Carexflacca ]Ccyperaceae ][N ]IHM [0 1120.3 1! _ [
jentaurea Asteraceae ]N HD n.m. 19.3 J_ _

nerastium [N
[Caryophyllaceae [HD 0 26.3
fontanum111 !
Chamerion
[1
f.

Onagraceae [N ][HD 23.3

angustifolium 111 1 1nl

al1um Chenopodiaceae N HD [Ii 21.4

Conyza Asteraceae Nd |HD 29.3

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Dactylis Poaceae N HM 0 26.3
glomerata
Deschampsia Poaceae N HM ][o_____
[21
flexuosaI
Digitalis J
purpUrea][Scrophulariaceae ]IHD ] I[29.5
Epilobium N HD l
hirsutum
Onagraceae
_ _ [ FI ][ 23.9
23 ___ 1
Eriophorum Cyperaceae IN ][HM 11.9
]_01_____
vaginatum I 11 11 111
Festucaovina j|Poaceae N ][HM [0 1[28.4 |1I
estucarubra IPoaceae I ][HM l 1130.2 1! _
I
Galiumaparine ][Rubiaceae [N ]|HD [O 5I3
][35.3
Helianthus
Atrca pH 02.
annuus
Helictotrichon Poaceae [N ]HM ]22.2 1
Holusan]us Poaceae [N]HM J[ ]39.9____
|Koeleria PaeeliEi
[N ]HM [ ]16.9 ___

macrnt]aPoaceae
Leontodon
Asteraceae IN [HD i 25.1
_ _

Loliumperenne ]|Poaceae I|N ][HM JIo 122.3 ]1_ I

Lotus
comniculatus
[Fabaceae [N
111
[HD 3 125.6 11 __
|o___latus 11l
Origanum
Pilosella
IffiCinarUm
!
vulgare|[Lamiaceae
IAsteraceae
[N
[N
1
HD
IHD
I1
k
J[4 ][23.4
][21.1
1I
11
11
_

__
I

Plantago
lanceolata iPlantaginaceae [N 1111111 rHD 15
1_
1_8.4 _ _

Poa annua ||Poaceae J[N I[HM 1[? ][n.m. 11

Rubus [NRosaceae N HD 620
chamaemorus I I
RumexacetosellaPolygonaceae[N j[HD
.r
26 20.1 Zf
rtc doca Utiaea. HD 1.9

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J. H. C. Cornelissen (1996). Leaf decompositionratesappendix2

Dryweightloss ofleaflitter as % ofinitialdryweightin fivetreatments(see text).In thespeciesmarkedwith

*, samplesweighing 0.5 g wereused,and% dryweightloss was multiplied bya factorof 1.11,as derivedfrom
a controlexperiment withBetulapendula(1.16 timesmoreweightloss in 0.5 vs. 1 g samplesin W8, 1.13times
morein W20) andFraxinusexcelsior(1.07 timesmorein 0.5 vs. 1 g in W8, 1.08timesmorein W20) using0.3
mmmeshbags.Forcompoundleaves(see Table 1) onlyresultsforlaminaearegivenhere.Boundariesof
weightloss classes(see text)werein eachtreatment:W8F: (1) 0-12(2) 12-24(3) 24-36(4) 36-48(5) 48-60%
W8C: (1) 0-16(2) 16-34(3) 34-54(4) 54-76(5) 76-100% W20F: (1) 0-17 (2) 17-34(3) 34-51(4) 51-68(5) 68-
85 % W20C: (1) 0-30 (2) 30-55(3) 55-75(4) 75-90(5) 90-100% S8F: (1) 0-16 (2) 16-32(3) 32-48(4) 48-64
(5) 64-80% weightloss. Meanweightloss class is themeanofthevaluesinthesefivetreatments.

---
|WTreatment ILI
*IiiW20FIL iZ ||W20C|i I S8F *LIiIZ n
l St.error
classMean MeanSt.errorMean St.errorMean St.errorMeanSt.errorl
class
Woodyspecies Liii LI]Z 1][ * L .11._I .
Acercampestre 0.77 j|1.74
251190.72 45.7 0.85 41.1 J ][2.5
A,cerplatanoides 18
18 .81 15.2 0.96 1136.3 |1.7 II42 4i I I I ] I2Ii I
Acerpseudoplatanus40.3_1.65 2.5 3.13 553 111.54 J[81.3_4.13 ][. ] _ 4
Aesculus i139
14.7 1.6 3 H4O117
15.7 1.9 23.9 1[3737.7 1.0
hippocastanum I
1.i.8
. 11

A.lnus
glutinosa 139.5 |1.35 |6615.72 177.3 |1.85 ||46 01.49 C I14
Amelanchierlamar 5.20.4 116.6 10.54 1124.1 j73 j37.9 i7.49 2II |IiiI
Andromeda1polifo5 056 10.64E IZ ] |1Z 5
rbutusunedoEfi EZi EZZZLjEZE 111Z
Aucubajaponica : I i1l 1 1 1.4
13 .0-8
|Berberis
vulgarils 35.2|0.52 471 2.07 56.5 1.4 |86.1 |2.88 3.5
1
Betula1nana 17.20 jE6 =lZ 94 1I49 ]LZ i1I I] IIZi
etulapendula 2 .60.42 122.711.6 129.911.0 I 62iI.29 J[ II21 I
Buddlej 2 142.7282 183.28.1
adavidii 22 1.39 137.218 I3
B uxussempervirens*
115.7 0.?95 1 11127.7 11.4 11 1 1 11 2 1

Callunavulgaris .210.57 IC I1E|.05 | |IL

C'arpinusbetulus 17.90.5 1122. 0.79 1134.2111.07 I49I7 33 IZIII 12.3I I

Castanea
sativa 6_
I _1.35 15.3 08 31.3 1.3 ||35.6 |ZI
I|6I73 LI1||1.81 I
ClematisIvitalba 371.35 8.1 117.62 73512.74 97.31121 4.5
Cornus
sanguinea 13.71.35 7E6.54 |71.6 |2.8 1p89|0.41 6
34.5S9 14-6
avellana
ICorylus 125.5 095 26.3 860 401 1.56 154 19.1 2.51g
|Cotoneaster
horizontalis 20.2 :l
1.17 r L 11 11.4
134 il m iL 1.r 2.5 m
1j26 fj31 117.31[4Z IIiL IZr 188.811j.6
rCategusmongyna j1 yy 13y

octopetala* j5S.3 [67.3

|Dryas 1[| 1|~ 8||30.48 I. 1. [1 7] 11

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 10.31 I.
nigrum* 117.2
Empetrum I 127.411.11 2jJ . f J
Erica
cinerea 1131 0.53 20.4 l |l1 1.5
Buonymuseuropaeus [391 1.03 51.9683 165.5 ||168 |94.2 113.53 4LII
Fagussylvatica E18.30.31 18.2 0.52 122.7 0 ||21 26.3 10.48 I.8
Ficuscarica 56.8 1.71 86.7 4.03 83.32.86 |98.6 |0.22 ] l
Fraxinusexcelsior 33 1.24 5.318.57 1155 IEEiIIii|90.2 113.93 1147.410.86 1
Fuchsiamagellanica 120.8079 21.94.33 |46.2 |0.8 |89.2 |5.87 z .1 3
HebexZfranciscana ||0.47 ZTI Z
H ederahelix 28.4 1.29 25.81.8 47.9 1.45 |62.9 |4.41 36.70.6 2.8
Helianthemum 123
22
2 [ 36
1 116 ir 11 [ 11
nummularium* 12 2I

|Hippophaerhamnoides 4][10.64 __2 3 1.1 ||2|| 1[. iz 2z |

calycinum iI3.1I0.41 114.7
Hypericum 10.34 17.3 099 8.1 [069 1[I0. 1
| 1xaquifolium 14150.65 1 54 130 1.84 i45.5 116.79 33.8 0.8 1
communis5EZ 0.22
Juniperus J 3 j 13 EZZIZ
0.47 Z
|Labumnumanagyroide5.8 0.91 71.4 .74 638 1.8 |98.1 10.47 4.3. 114.
Larix decidua 15j490.32 IEZZIj 22.9 10Z34 Z2IiZ IiUI
Larixkaempferi 14.4 .6 1z.118.9 10.71 21 1 G
ovalifolium
L igustrum 461|| | | | 1|.8 6
|Ligustrumvulgare* 131.3 | 1.68 |36.7 4.09 ||53.8 13.57 |63.2 6p.06 1 13.3
Lonicera
p]riclymenum
5.9 2.5 80.6 ]2.1 ][ ]_ . 1_ ____

Malussylvestris 27034 195 4014.87 56.812.23 95.2 ||2.19 ]L3I8II

Pi.ceasitchensis 51E 0.24 |E l|E|0.32 | || le H

|Pinussylvestris
1E20.6 1Z31 1125 E
0.67 |Z LJZ 1.5|

Po:pulusnigra 33.81.34 3m0.70.0-

3 5.5 |1.46 ||6.2 |5.25 |=|
1pulustremula 31.5 1.31 41.5114.06 1154.5I 1.48
LI 76.2 115.19 3.5IlI
P|runusavium 20j7i0.36 25.2 4.22 1135.2||1.07 ||88.4 114.48 |= II .I
laurocerasus 17.4 0.69
runus 15.210.43 112.6 2.31 44.6 118.47 33 1.24 I
1runuslusitanica 271421.36 30 2.05 1142.7 1i0.94 |J75.8115.61 II II Z
'runus
padus 4?3.61.61 72|.4 3 ||7222.13 195.3 |1.46 |11
Prunus
spinosa 32fi6 0.72 60.49.52 151.5 11.87 1957 13.94 4z1 1

Pyruspyraster 28.9 1.43 361.7 5 |52 |1.9 185./ 155 I=ll3.5

Quercusilex 7.4 0.52 .9 1.56 15.1 1.07 150.9 lE . 0.69

Quercuspetraea 13.20.35 13.90.44 |26.3 10.74 |25 |1.83 |z.|1.5

Quercusrobur 8.4 0.63 ---9.60.36 116.3 11.13 1116.5 11.36 11 111

Rhamnnus catharticus 4821.79 1z .I80.3 12.11 11 1 511
1hododendron 11.
6.9 1.07 11652 61 116.2
936.0778 6 1661 1
I~ 0.78cu
~ ~ ~~. ]j111 111071 16. 1.1

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Ribes alpinum 17170.82 |1.25 27.1 2.25 60.2 | J9.07 L.2.3
Ribes nigrum 11591.65 34.8 7.85 37.2 1.88 1185.4 |I3.36 3IW I
Ribes
rubrum 2 0.63 2.4 I1.27 3.5ZLII.
17.69 148.512761194.2
Ribesuva-crispa 831.3 892 2.77 I15792.86 !90.1 1.192 4.5
1
REosaarvensis I3ZZIZ
12310.62 37 1.48 K 1 12.5
|.osacanina jO
12.91 E29.7 1.26 1 .88 1189.2 I 15.35 LiL iI
Rubusfruticosus 16.8|0.88 L19.3 |36.4 ||2.3 ||48.7 |6.21 | |
Rubusidaeus 129.4 1.66 41.7 16.1 1161.8||2.5 199.3 |0.28 38
Salixalba 390.67 6 I7.49 161.9 1.91 1I9.5 | I2.15 ]I II
Salixcaprea I 120.84 I3 16.19 138.3 I 90.9 J3.74 ][. ][ 3.3
fragilis
|Salix 28. 094 4IE=I.
I[ Z .87 111118
3 11IIE
Sa4bucusInigra 861.12 I2.2 7.78 62.519 11900 Ii 11 11LZ
Solanum dulcamara IE 11.09 1238 1.76 7412 I2I2 199.7 ]015 j[__5__
Sorbus3ara 1738 1.83 1.85 37163.03
Ii1897 ]26 J[5 IZ L
aucuparia
lorbus 2 3Ill.25 5IZ . 6.04 1147 11.64 8343 7.61 33
to yrinalis ]I 711.16 1i 1 1.08
iorbus 1375 111.42 37.6 3Z55 2I 1
albus
Viymphoricarpos 01 1.44 1717.2 133 76.5 11.96 1113.9 0 1I124 I1. 1. 4.8 |
gallica* 34.31.47 5EC:E
Tamarix 4.1 1237 31IL.5I1
baccata
Vaxus ]4II41.14 lEI=I1 I iI I10.71Z
139 . 14 IIIE 1
][ ZEI199 I4.I 1][ _18 21 11 IEI ] 1
polytrichus
Hhymus
Tiliacordata 1ir 1.38
]73.76 |128.8
4|1.1 169.7 12 1
iliarxavulgaris 18.4
12.04 5[. ] 07 .9 ]0.91 II.
118 3.29 J4__ _

Brlexyeuropaeus [10 0.84

4 1191 10. 1 11
Ulexgallii l1.01
1l0.2 23.8
11]0.94 11. 1 E I1.5
Ulmusglabra 3 .2 1.07 44.3 .16 57 ]3.44 194.5 13.w3 4EI L
pacciniummyrtillu 1 1.03 _ I ]33 ii.49 11 1 1 2
1.5 [0.59
Bracciniumvitis-idaea 51.4 0.62 114.4 110.75 113.3 .65 16 0.41 1.
Viburnumlantana 28.62.13 lm 9.18 l6-96 IL
52.9 3.37 1l79.1 115i
Viburnumopulus I3 1l.64 6.4 5.6 I603.8 58 l86 l6.27 5. .
H erbaceousspecie
Agrostis
capillaris l 14.810.29 21 . 10.87 21 [
-9 1L] <
Antrhoantarthum
lE0< EH:g
sylvestris 4
Anthriscus 1.85 14il.! 1
.m159
71.7 E
Affhenaterum
elatius 126 IE I147.4
- 11.321 1
1124 IZ 1
BrachvpodiumE-. i11.41. F IL". 1119.
I"0.-11. 11. 11. 11 I1

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Carexflacca 16.5 0[24 5.8 0.81 13.9 0.93 I!
1113.8 I
Centaureascabiosa
jj34 0.67 I5.8 17.84 1164.2 4.5 92.6 14.77
erastiumfontanum*I2L0.83 I 44.1 1.14 _ 3
IChamerion
angustifolium Ill4.6
_
0.8
_
23.3 13.81 140.4 2.13 ]92.1 16.76
_ _ _ _ _ _ _ __I1___
1.___3_1 j.
ChenopodiumalbumI56.3.01 i92 17.67 1182 112.15 1199.31[0.22LiiL..si
onyzacanadensis 42.6 E 1Z45 1 i66.4 11.84 4LIZ IZ ZIIZZ
glomerata 21.2 EIZIZ1
actylis L38.5 110.52 LIIZZIZ ZII2.I
Deschampsiaflexuosa I
2E0 LZ.48 3ZI 8.3 11.72 Z1 IZ iZZE
igitalispurpurea 37.73.86 72.4 16.74 I616I4.12 1193.3 I1L.3I I lZI
pilobiumEhirsutum6.69 41. 45.9 I.49 LL90.6 L5.25 IEI 1
Eriophorumvaginu0.39 6.
2]E0.4 ZI0.74 I2.4 0.5I I 1
FestucaIovina 140.46 1511. ]Z0.8E i
I128.3 1 IZE Z 1.
estuca1rubra I.21E I39 11i1.58
I[ii I .7I1.22
]1 I5__ I2.L3 IE 11LII3Ii
aliumaparine J7351.49 [ 4J j 2.1933 1 1 L1 11E 11141
elianthusannuus 1i.32 115 146 1 3
elictotrichon9pratense14.41.21 1.9 3.5 2.02 135. 091 2II1
Holcuslanatus 2Ij 0.52E 21 L2.23 Z 5 IE.08 Z64.5 2.551
K;oeleriaImacrantha
2II52.2 II 111 12 45
1183 1.9 1I1
Leontodonhispidus E37.5222 IE167.9 12.19 11 1 EII4
Loliumperenne E34.71.39 60.11.El33 3111.5I 1
Lotus
corniculatus 5 s 1I.58 1 1I75.1 112.09 11 113 5
riganumvulgare 0.84 1I69.41 115.1 12.63 1164.3 IE36 II3IE
ilosella1officinarum 110.95 48 1.2 1189
9311I I 11 I
Plantagolanceolata 25.3 2.09 42E314.35 l48.7 1l.52 l88.7 l2.68- 13.3
Poaannua 4031?077 IEI I64.3 1l.27 11 1 EI 14 F
Rubuschamaemorus I9.3 0 67 12.31 .0v I24.9 11.42 122.1 13.17 1.I i31
Rumexacetosella !3io053 1.Z l20.90l.9 1 11 IZ 1.5
Urtica
dioica 52.1l2.01 [98.41 29 I83 4.47 1100 10.03 15.v
Zeamays 25.9 2.88 I32.4 458 I53.1 .93 l64.3 1l5.68 IEIg3

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