An Experimental Comparison of Leaf Decomposition Rates

Transcription

Journalof
Ecology1996,
84, 573-582
An experimental
comparison
ofleafdecomposition
rates
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
multispecies
screeningofleafdecompositionrateswas 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-assess
predictorsof 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
effects
ofa givenspeciesis a functionoftheinteractive
Itiswellestablished
thatpotential
decomposition
rate
of all theseand otherfactors,each ofwhichmayvary
ofleaflitter
on thephysico-chemical greatlywithinfunctionalor taxonomicgroups.This,
dependsgreatly
properties
oftheleavesofthespeciesconsidered
(e.g.
combinedwiththe small numbersof speciesused in
Broadfoot& Pierre1939; Aber et al. 1990; Gillon et
to identify
patterns
moststudies,has made it difficult
al. 1994;Couiteauxet al. 1995). As reviewedby Swift
and testgeneralitiesin potentialleaf decomposition
et al. (1979) and Anderson(1991), severalfeatures
ratesamong speciesgroups.Withoutthiswe cannot
ofundecomposed
arenegatively
associated evaluateor predicttheeffects
ofchangingplantspecleaflitter
withdecomposition
rate.Thesefeatures
include
lignin ies compositionon soil decompositionratesin differcontent(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
representative
numbersofspeciesor groupsofspecies
phenoliccompounds
suchas tannins(Nicolai1988;
thatare identifiable
in termsof functionalattributes
Kuiters1990),lignin:nitrogen
ratio(Melilloet al.
or taxonomy.
1982; Buth & Voesenek 1987; Cotrufoet al. 1994),
This paper reportsan experimental,
simultaneous
physicalleaftoughness
(Gallardo& Merino1993)
screeningof leafdecompositionratesfor 125 species
and physical
barriers
(e.g.hairs,spines,wax)on the
importhat are native,naturalisedor commercially
Introduction
c) 1996British
Ecological Society
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574
Leaf
decomposition
rates
tant in the BritishIsles. It teststhe hypothesisthat
potentialdecompositionrateof leaflittercan be predictedfromwhole-plantfeaturesthatreflect
thefunctioningof the plants in theirnaturalenvironments.
These featuresinclude life-form
and deciduous vs.
evergreenhabit,as well as specificleaf area and autumncolorationofthefreshlitter.Sourcesofvariation
in potentialdecompositionrateare testedusingfunctional analysesof a representative
part of a presentday floraas wellas taxonomicanalyses.
Methods
SELECTING
?
1996 British
EcologicalSociety,
JournalofEcology,
84, 573-582
AND
COLLECTING
LEAVES
Leaf litterwas collectedwithina 25-kmradiusaround
UK (53?23'N, 1?33'W),wherepossiblefrom
Sheffield,
the species' (semi)naturalsites includingparks and
woodlands.For deciduouswoody species,theplants
thatweresampledshowedtypicalautumncoloration
(see below) as seenin theSheffield
area. Some species
thatcould not be foundin thewild locallywerecollectedfromSheffield
University's
ExperimentalGardens and a few fromprivategardens.The original
data can be retrieved(as AppendixI) fromthejournal's archiveon the World Wide Web (see a recent
issuefordetailsoftheJournalofEcologyhomepage).
All but one species had thereforebeen exposed to
similarclimaticconditions,which facilitatedinterspecificcomparisonoflitterweightlosses.OnlyPicea
sitchensisleaves were collectedoutside the area, in
northernScotland.
Throughoutautumn1993,freshly
senesced,undecomposedleaveswerecollectedfromsexuallymature
plants. In virtuallyall woody and some of the herbaceous 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 functionallydisconnectedfromtheplant.Leaves werecollectedwithoutbias towardssize,shape or colour,but
those with symptomsof significant
herbivorywere
avoided. In some summer-shedding
evergreens,the
collectedleaves weresuspendedin the standingvegetationor on top ofthelitterlayer.For theseand a few
othersummer-shedding
evergreenspeciesadditional
collectionsof freshlyshed leaves weremade in summer 1994 and treatedin a controlexperiment(see
below). Petioleswereconsideredpart of the leaf. In
compound leaves the laminae and rhachisweretreated separately.This would permitinterspecific
comparison of whole leaves as well as leaf analogues.
For Cytisusscopariusand Ulex spp. smalltwigswere
treatedas leaves, because in these species the twigs
are the main photosynthetic
producers.For Calluna
vulgarissmall shoots (78% of dryweightbeingleaf)
were treatedas leaves, since theywere shed in this
form.In the biennialsAnthriscus
sylvestris
and Digitalispurpureabothfirstand secondyear'sleaveswere
harvestedafterthesecondgrowingseason; in thelatter 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
theovendryweightin a minimumsubsampleof eightleaves.
Since surfaceaccess to soil decomposerswas likelyto
be a determinant
ofdecompositionrate,thetrueonesidedsurfacearea ofnonlaminarleaveswas calculated
bymultiplying
theprojectedleafarea bya conversion
factorderivedfromobservationsof leaf cross-sections.
As autumnleaf colorationmay reflectthe chemistryoflivingleavesthatsecondarilyaffects
decomposition 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 totalpurplishcoloration.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 laboratory(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-mmmeshnylonnet. The double layerappeared effective
in retainingleafparticlesand at thesame timeallowing free movementof macrofauna,includingthe
earthworm
In narrow-leafed
Lumbricusterrestris.
and
needle-leafed
specieswheremajor lossesmightoccur
throughthe 5-mm-mesh,only 0.3-mm-meshbags
wereused. Precautionsweretakennot to break airdriedleaves.Whenflattened,
bothtypesof litterbags
had an innercompartment
of 10cm x 7 cm. Samples consistingof 4-cm x 1-cmpieces of Whatman
grade 540 filterpaper, includedfor potentialcomparisonwithotherstudiesbecause of its guaranteed
constantquality,were treatedas if theywere litter
samples. For each species,a subsampleof the leaf
litterwas weighedair-dryand again after48 h in an
575
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;
bags buriedfor8 winter
W8C = (coarse) 5-mm-mesh
W20F = 0.3-mm-meshbags buried for 20 winter
bags buriedfor20 winterweeks
W20C = 5-mm-mesh
from24 November1993
S8F = 0.3-mm-mesh
bags buriedfor8 summerweeks
from29 July1994.
?
1996British
Ecological Society,
JournalofEcology,
84, 573-582
was eightin mostspecReplicationper treatment
ies,butfiveto sevenwheretheamountofleafmaterial
stored
was limited.For thesummer(S8F) treatment,
materialcollectedin autumn1993was used,as wellas
leavescollectedduringsummer
shedevergreen
freshly
of
1994.This would allow an evaluationof theeffects
timeoflittercollectingon thepotentialdecomposition
rate.
The samples were buried in a purpose-built
decompositionbed at SheffieldUniversity'sExperimentalGardens. The bed measured6 m x 5 m and
partly
was on a 3? slope facing north-northwest,
trees.In September1993 all
shaded by neighbouring
rootingplantsand thelitterlayerwereremoved.The
brownforestsoil, pH
top 20cm of the loamy-clayey
5.93 + 0.34 (measuredin 24 samples,with 1-mdistancesbetweensamples),was mechanicallydug over
and wooden walkwayswere installedlengthwaysat
1--mdistances. On 7 October 1993 leaf-mouldat
different
stagesof decay (mostly1-2 yearsold) was
collectedfroma nearbycompost pile consistingof
litterfromspeciescommonin local cityparks,suchas
Betulapendula,Fagus sylvatica,
Acerpseudoplatanus,
Fraxinus excelsior,Pinus sylvestris,Quercus spp.,
Sambucusnigraand Taxus baccata. This leaf-mould
and spreadoverthedecompowas mixedthoroughly
sitionbed inan 8-10-cm-thick
layer.Leaf-mouldsamples 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
W8 or W20. All13024
assignedat randomto treatment
withrainwater,ranwintersampleswereremoistened
domizedwithinW8 or W20, and buriedon thesame
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
afterburial.Thesewereusedto
immediately
retrieved
assesstheloss ofleafparticlesfromthelitterbags and
(leaching).The relatively
the effectsof remoistening
low numberof summer(S8F) sampleswereburiedas
above in one of thecentralplots only.A 3-cm-mesh
acrossthedecompositionbed
nylonnetwas stretched
to protectthe samplesfromgreysquirrelsand other
macrofauna.Accumulationsofleaveswereshakenoff
regularly.
Afterretrieval(W8 on 19 January;W20 on 12
April;S8F on 22 September1994),the sampleswere
adheringsoil, soil
storedat - 14'C. Afterdefrosting,
faunaor otherextraneousmaterialwas removedfrom
rinsthedecomposedleaflitterbybrushingor swiftly
ing withwater.The macroscopicfauna foundinside
litterbags includedsmallerslugs and snails, centipedes,Collembola,acarid mites,Coleopteranlarvae
and Oligochaetae (includingLumbricusterrestris).
Litter samples were dried for 48 h at 80 'C, then
weighed.Decompositionratewas definedas thepercentagedryweightloss after8 or 20 weeksof burial.
The effectsof taxonomicrelatednesson the patwoodyplanttypeswere
ternsfoundamongfunctional
testedfollowingKelly & Beerling(1995), who used
a method derivedfromFelsenstein(1985). First a
treefromspeciesup to class level was
phylogenetic
constructedfollowingCronquist(1981). For each set
of two or moretaxa belongingto the same taxon of
the nextlevel up a contrastwas taken (e.g. is litter
A thanin life-form
B?),
weightloss higherin life-form
afterwhichthe species used were taken out of the
conanalysisto ensureindependenceof thedifferent
trasts.Whereall membersof a taxonbelongedto the
samecategory,themeanvalue forthesememberswas
used fora contrastat thenexttaxonomiclevelup.
Results
WEATHER
Data fortheexperimental
period(fromWestonPark
sited800m northMeteorologicalStation,Sheffield,
northeastof decompositionbed; details not given
were
and temperature
here)showedthatprecipitation
broadlysimilarto thoseforthepast 10years.Rainfall
months,so that
was substantialin all experimental
thedecompositionbedwas neversubjectedto drought
Snow coveredthe bed duringtwo
duringtreatment.
periodsof about a week.Onlymildfrostoccurred.
DECOMPOSITION
RATES
forindiMean percentageweightloss was determined
(Appendix2, WWW
vidual speciesin each treatment
valuesweregrouped
archive).Withineach treatment
576
Leaf
decomposition
rates
into classes (class 1 = very slow, 2 = slow, 3 = intermediate,4 = fast,5 = veryfast)such thatall classes
containedsimilarnumbersof species. Mean weight
loss class foreach specieswas thencalculatedfrom
the values forthe fivetreatments.
Variabilityin dry
weightloss among replicateswas generallylow and
no correctionforpositionwithinthedecomposition
bed was necessary.Relative dry weight losses of
laminae only was similarto that of whole leaves in
all specieswithcompound leaves (Table 1), both in
treatments
W8F and W20F.
In 0.3-mm-mesh
bags relativeweightlossesamong
121 species after8 winterweeks (W8F) were correlatedstronglywith losses after20 weeks (W20F)
(r = 0.96, n = 121, P < 0.001). In 5-mm-meshbags
(W8C vs. W20C) thecorrelationbetweentreatments
was less strong(r = 0.82,n = 75, P < 0.001),because
in severalspeciesmaximumweightloss was reached
before20 weeks.Relativedryweightlossesin thetwo
types of bags after8 weeks (W8F vs. W8C) were
closelycorrelated(r = 0.93, n = 75, P < 0.001), but
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
20 weeks of burial. Ten species had verylow and
similarrelativeweightlosses in 0.3- and 5-mm-mesh.
Species-burial-periodand species-mesh-sizeinteractions were evidentbetweenindividualspecies but
did not obscurethegeneralpatterns.
CONTROL
TREATMENTS
AND
COMPARISONS
Initialrelativedryweightloss untiland includingthe
momentof burialvariedsignificantly
withina subset
of 14species(Table 2). Therewas no overalldifference
between0.3 and 5-mm-mesh,
which indicatesthat
initialloss from5-mm-mesh
was negligible(as it was
assumedto be from0.3-mm-mesh)
and initialweight
loss was due presumablyto initialleachingof watersolubleorganicmaterial(cf.Nykvist1963).
There was no significantdifferencein relative
weightloss of Whatmanfilter
paperbetween0.3- and
5-mm-meshin eitherW8 or W20 treatments.This
Table 2 Mean (? SE) initialpercentagedryweightloss until
and includingburial
Mesh size
Acerpseudoplatanus
Alnusglutinosa
Betulapendula
Bromuserectus
Buddlejadavidii
Carpinusbetulus
Digitalispurpurea
Epilobiumhirsutum
Fagus sylvatica
Festucaovina
Fraxinusexcelsior
Larix decidua
Juniperus
communis
Plantagolanceolata
0.3mm
5mm
4,6 +
4.1 +
1.6 +
0.6 +
1.0 +
2.2 +
10.6 +
3.6 +
0.4 +
0.8 +
1.0 +
0.03 +
0.2 +
2.5 +
4.7 +
4.6 +
2.1 +
1.3 +
1.7 +
2.9 +
9.9 +
4.6 +
0.2 +
0,28
0.30
0.073
0.30
0.23
0.15
1.85
0.87
0.11
0.072
0.23
0.028
0.23
0.68
0.46
0.24
0.31
0.37
0.17
0.16
1.72
0.82
0.049
1.8 + 0.11
3.6 + 0.50
Resultsoftwo-wayANOVA: mesh-sizeeffect
F, = 3.42,NS;
specieseffectF13= 38.9, P < 0.001; interactionFlo = 0.36,
NS.
suggeststhatmacrofaunahad notconsumedanyfilter
paper and thatenvironmental
conditions(e.g. moisturecontent)had been similarformicro-organisms
in
bothmeshtypes.
Relativeweightlossesin S8F correlatedwiththose
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,althoughpossiblyimportantwhereindividualspeciesare compared,did not
affectthe rankingof specieswithregardto relative
weightloss.
In a subset of threeevergreenspecies (Table 3),
only one (Hypericum calycinum) showed a significant
difference
in relativeweightloss betweenleafmaterial
collectedin autumn 1993 againstsummer1994,but
this was not enough to take it into anotherweight
loss class. In addition,relativeweightlosses of leaves
of Prunus laurocerasus, Taxus baccata and Vaccinium
vitis-idaea,collectedin autumn 1993 and buriedin
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
? 1996British
Ecological Society,
JournalofEcology,
84, 573-582
Aesculushippocastanum
Fraxinusexcelsior
Laburnumanagyroides
Rosa arvensis
Rosa canina
Rubusfruticosus
Rubusidaeus
Sambucusnigra
Sorbusaucuparia
W20F
petioles
laminae
leaves
petioles
laminae
leaves
17.5
32.8
30.5
18.5
21.8
22.9
33.5
47.4
24.7
14.7
33.0
45.8
23.1
25.8
16.8
29.4
48.6
26.3
15.2
32.9
43.4
22.6
25.3
17.8
29.9
48.4
25.9
33.1
51.3
52.0
38.4
45.3
42.3
61.2
63.6
48.2
23.9
55.0
63.8
37.0
47.0
36.4
61.8
76.0
44.7
25.4
54.5
62.0
37.2
46.7
37.4
61.7
73.9
45.4
577
J.H. C. Cornelissen
70
PLANT
r 0.95
60
50
40
*
20
10
0
W22F
10
40
30
% weightloss in
20
50
60
70
80
Fig.w1Relative dry weightlosses in W20F vs. S8F. Open
circlesindicatethatthesame leafcollectionwas used in both
treatments.
Solid squaresindicatethatleavesin W20F were
collectedin autumn1993and leavesin S8F in summer1994,
bothfromsimilarsites.
Table 3 Mean (? SE) percentagedryweightloss in S8F in
evergreenscollectedfromsimilarsitesin autumn 1993 vs.
summer1994.Quercusilexleaveswerecollectedfromunder
thesame treein bothperiods
Collected
Quercusilex
Hedera helix
Hypericumcalycinum
autumn1993
summer1994
12.64 + 0.40
38.68 + 0.74
10.62 + 1.11 **
11.46 + 0.69
36.74 + 1.70
15.01 + 0.70
**P < 0.01 (t-test).
mer 1994, were close to the fittedregressionline in
Fig. 1. These data indicatethatrelativeweightlosses
evergreensin this
of leaves of the summer-shedding
studyweregenerallysimilarwhethercollectedfreshly
fallenin summeror,withsome delay,in autumn.
In a subsetof threedeciduouswoody species,collectedfromcontrastedsites,two revealedsignificant
variabilityin relativeweightloss (Table
intraspecific
4), unrelatedto the degreeof exposureof the site.
Althoughdata on only threespecies do not justify
variabilitywas much
conclusions,this intraspecific
smallerthanthevariabilityamongdeciduouswoody
species(cf.Appendix2).
AND
TYPES
DECOMPOSITION
RATES
Unless otherwisespecified,resultsindicate weight
simibecauseoftheessentially
lossesin0.3-mm-mesh,
lar patternsin 0.3- and 5-mm-meshand the larger
numberof speciestreatedin 0.3-mm-mesh.
inmean
overallheterogeneity
Therewas significant
differspecies
litterweightloss amongninegroupsof
5).
The
Table
2,
(Fig.
or leaf habit
ing in life-form
20
weeks,
or
patternwas the same afterburialfor8
(Table
effect
as shownbytheabsenceofan interaction
5). Withintheherbaceous(nonwoody)species,dicots
decomposedon averagefasterthangraminoidmonocots. Withinthe woody species,deciduous species
decomposed twice as fast as evergreens(28.4 + 1.5
vs. 12.9 + 1.6% weightloss in W8F; 48.2 + 2.3 vs.
24.7 + 1.0% in W20F). These resultswereconfirmed
whentaxonomicrelatednesswas takeninto account
sourceof variwas a significant
(Table 6). Life-form
woody
in
deciduous
rate
in
ation
decomposition
the
Within
5).
(Table
in
evergreens
but
not
plants,
and
scramblers
climbers
species,
deciduous woody
decomposed faster than self-supportingspecies
(48.0 + 7.2 vs. 27.4 + 1.4% weight loss in W8F;
76.1 + 2.3 vs. 46.8 + 2.2% in W20F). Owing to the
small numberof comparisons,thiscould not yetbe
supportedbya taxonomicrelatednessanalysis(Table
6). The latterdid demonstratethatleaf litterof subshrubsgenerallydecomposedmore slowlythan that
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).
This was due mostlyto the woody species (W8F:
100
*W8F
90 -
W20F
80
70
0
60
*F3
50
T
40
30T
20
o~~~~~~~
10
Table4 Mean (? SE) percentagedry weightloss in S8F
sites
withindeciduouswoodyspeciescollectedincontrasting
in autumn1993
Site
G 1996 British
Ecological Society,
JournalofEcology,
84, 573-582
Cornussanguinea
Fraxinusexcelsior
opulus
Viburnum
*P <
exposed
sheltered
62.79 + 1.41
47.42 + 1.25 **
55.77 + 1.43 *
69.63 + 4.59
53.87 ? 0.86
50.86 + 1.00
0.05, **P < 0.001 (t-tests).
u
.
~
0
0
u
0
C.) 0
-o
(D
- o
0
o
-C
u
~
U
0~~E
E
C0
in
dryweightlossesin 0.3-mm-mesh
Fig.2 Mean relative
habit.Woody
groups of specieswithsimilarlife-form/leaf
D = deciduous,
andscramblers.
climbers
= woody
climbers
themean
from
Meanvalueswerecalculated
E = evergreen.
species.Standard
weightlossesof individual
percentage
errorbarsareshownone-sided.
578
Leaf
decomposition
rates
Table 5 Summaryoftwo-factor
ANOVAS withfactorsfunctional
group(life-form/leaf
habit,see Fig. 2) and burialperiod(W8F
vs. W20F). All data werenormalizedthroughln-transformation
priorto analyses
p
Functionalgroup
Amongall life-form
and leafhabitgroups
Graminoidmonocotsvs. herbaceousdicots
Amongall deciduouswoodylife-forms
Amongall evergreen
woodylife-forms
Woody species:deciduousvs. evergreen
Deciduous woodyspecies:climbers/scramblers
vs. self-supporting
plants
Functional Burial
group
period
Interaction
Total
d.f.
<
<
<
<
<
<
<
<
<
<
<
<
237
71
121
43
167
121
0.001
0.001
0.001
0.030
0.001
0.001
<
<
<
<
<
<
0.001
0.001
0.001
0.001
0.001
0.01
0.99
0.62
0.97
1.00
0.33
0.86
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
Functionalgroups
No. ofcontrasts/
No. of disagreements x2,
p
Deciduous climbers/scramblers
vs. otherdeciduouswoodyspecies
Subshrubsvs. shrubs/trees
(deciduousand evergreen
habitsin separatecontrasts)
Deciduous vs. evergreenhabit
High SLA vs. low SLA
Autumnleafcolorationgreenor yellow-green
vs. less thanhalfgreen
3/3
7/7
15/15
27/17
11/10
NS
< 0.05
< 0.01
NS
< 0.01
r = 0.61, P < 0.001; W20F: r = 0.54, P < 0.001).
This relationshipwas, however,not confirmedby a
taxonomicrelatednessanalysis(Table 6). Therewas
no significant
correlationbetweenSLA and weight
loss withintheherbaceousdicots(W8F: r = 0.41,NS,
W20F: r = 0.33, NS) or graminoidmonocots(W8F:
r = 0.22, NS, W20F: r = 0.25, NS).
Withinthe deciduous woody species, significant
heterogeneity
in relativeweightloss could be explained byleafcolorationat thetimeofshedding(Fig. 3).
The patternswerenot significantly
affected
byperiod
of burial (8 vs. 20 winterweeks) or mesh size, as
60
EW8F
50
AND
DECOMPOSITION
RATES
coloration(see Fig. 3), burial period (8 vs. 20 weeks) and
meshsize(0.3vs.5mm).Thedatawerenormalized
through
ln-transformation
priortoanalysis.
Totald.f.= 215
30
20
10
0
1996 British
Ecological Society,
Journalof Ecology,
84, 573-582
FAMILIES
Table 7 Summary
of three-way
ANOVA withfactors
leaf
W8C
40T
?
PLANT
T
70
E
indicatedby thelack of interactioneffects
(Table 7).
The most strikingdifference
in relativeweightloss
was betweenleaflitterwithvs. without(partial)green
coloration, a differenceconfirmedby taxonomic
relatednessanalysis(Table 6). Multicolouredleaves
(i.e. withmixturesof at least red,yellowand green)
wereintermediate
in relativeweightloss. The results
for brown leaves should be interpreted
with some
caution,sincethisgroupconsistsof threespeciesof
Fagaceae only.
Mean weight loss class was significantly
heterogeneous among the 10 main plant familiesin this
80
0
2.00
4.67
10.0
0.92
7.71
Fig.3 Mean relativedryweightlossesafter8 weeksingroups
ofspecieswithsimilarleafcolorationat thetimeofshedding.
Mean values were calculated from the mean percentage
weightlosses of individualspecies.Standarderrorbars are
shownone-sided.
Sourceofvariability
P
Colour
Burialperiod
Meshsize
Colourx burialperiod
Colourx meshsize
Burialperiodx meshsize
<
<
<
<
<
<
Colour x burialperiodx meshsize
0.001
0.001
0.001
0.36
0.21
0.22
< 0.44
579
study (Fig. 4, Kruskal-Wallis nonparametrictest,
J.H.C. Cornelissen %2 = 37.7, P < 0.001). The variabilitydoes not just
reflectdifferences
betweenwoody and herbaceous
taxa; slowly decomposingFagaceae and Ericaceae
and fastSalicaceae and Caprifoliaceaeare all woody,
whereasall-herbaceoustaxa includemoderatelyslow
Poaceae as wellas fastAsteraceae.
Mean weightloss classesofthedicotplantfamilies
in the experiment
werelooselyassociatedwiththese
families'evolutionaryadvancement(ranging from
primitive
to advanced)as quantifiedin thePercentage
AdvancementIndex (Sporne 1980; Sporne 1982)
(Fig. 5, r = 0.40, P < 0.05).
Discussion
EFFECTS
OF LITTER
QUALITY
AND
ENVIRONMENT
A broad body of literaturehas shownthatvariation
in leaf decompositionrates among species depends
greatly on litter (resource) quality. Lignin, for
instance,may enhancephysicaltoughnessof leaves,
5
n)3
T
T
E
a)
t
E
a)
$1
a)
1r
a)
a)
0
l
U)
a)
a)
0
U-~~~~U
|
a)
|0
a)
a)
Loi
Fig.4 Mean weightloss class (averagedover thefivetreatments)forgroupsof speciesbelongingto the same family.
Onlyfamilieswithat least two generaand fourspeciesrepresentedin thisstudyare used. Mean classeswerecalculated
fromthemean weightloss classes of the individualspecies
in each family.Standarderrorbars are shownone-sided.
and tanninsare known to act against folivoresin
livingplants(Coley 1983;Kuiters1990). The present
data, whenlinkedto initiallignin:N ratiosand lignin
contentsof leaf littersfound by previous authors,
provideadditionalevidenceoftheimportanceoflitter
qualityacross a rangeof woody species.Despite the
diversedata sources and analyticalmethodsused,
lignin:N ratio (r = -0.78, n = 12, P < 0.01, Fig. 6)
and lignincontent(r = -0.66, n = 14,P < 0.01) were
bothnegatively
correlatedwithlittermeanweightloss
class in thisstudy.Notwithstanding
theemphasison
litterquality,it is recognizedherethatthereare significantinteractions
of litterqualitywithdurationof
decomposition(cf. Mommaerts-Billiet
1971; Wieder
& Lang 1982; Berg & Ekbohm 1991) and with
environmental
factors(Swiftet al. 1979). The latter
includefeaturesof thelitterlayerand itsdecomposer
community
(Bocock et al. 1960;Heal & Ineson 1984;
Buth & De Wolf 1985; Elliot et al. 1993), macroclimate (Meentemeyer1978) and microclimate
(Escuderoet al. 1987;Taylor& Parkinson1988).The
methodologyadopted, for instancelitterbag type,
may affectsome of the above environmental
factors
(Anderson1975; Louisier& Parkinson1976). All of
theseinteractions
may be relativelyimportantas far
as relativeweightlossesofindividualspeciesare concerned.The presentresultshave shown,however,that
interactive
effectsof litterqualityand burialperiod,
season or litterbag type,althoughconsiderable,were
not so greatthat theyalteredthe broad rankingof
125 specieswithregardto litterweightloss. Onlythe
near-maximum
weightloss in a large proportionof
thespeciesin 5-mm-mesh
after20 weekscaused pronounced deviationsfromlinearitybetweenrelative
weightlosses in different
butevenin that
treatments,
case did not fundamentally
altertherankingof species. Possible interactionsbetweenqualityof the leaf
samplesand featuresof the surrounding
leaf-mould
layerhave not been assessedin thisstudy.However,
themixedleaf-mouldin thedecompositionbed had a
40
r=-O78, P'0 01
30
4
u'
*.
(n
n*
o
20
E20-
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
Sporne's percentage advancement index
? 1996British
Ecological Society,
JournalofEcology,
84, 573-582
Fig.5 Mean weightloss class of dicotyledonousfamilies
againsttheirSporne'sPercentage
Advancement
Index.All
dicotfamilies
included
in thisstudyarerepresented.
Mean
classeswerecalculated
fromthemeanweight
lossclassesof
theindividual
speciesineachfamily.
between
initiallignin N ratioof leaf
Fig.6 Relationship
litter
andmeanlitter
(fromtheliterature)
lossclass
weight
(thisstudy)in 12 woodyspecies.Data sources:King&
Heath(1967);Healetal. (1978);Chauvet
(1987);Hendriksen
(1990);Berg& Ekbohm
(1991);Slapokas& Granhall
(1991);
Van Vuuren(1992);Tietema(1993);Cotrufo
et al. (1994);
Domenachet al. (1994).Themeanlignin:N ratiowascalculatedwhereonespeciesappearedintwopapers.
580
Leaf
decomposition
rates
? 1996British
Ecological Society,
JournalofEcology,
84, 573-582
of both acidic substrates(e.g.
good representation
beech,oak, pine,yew),as favouredby fungiand certain invertebrategroups, and more base-richsubbacteriaand
strates(e.g. ash, alder,elder),supporting
groups(cf.Swiftet al. 1979).It can
otherinvertebrate
therefore
be assumedthatthedecomposercommunity
of
in thisleaf-mouldis to some degreerepresentative
thatinthenaturalhabitatsofmostspeciesin thisstudy.
In orderto explainthevariationin leafdecompositionratesfoundamongspecies,manypreviousstudies have scaled down fromthe leaf level to its individual structuraland biochemicalcomponents:the
mechanisticapproach. In contrast,the presentmulattemptto scale
tispeciesscreeningis an experimental
up fromtheleaflevelto thelevelof wholeplantsand
plant types:the functionalapproach. In supportof
thehypothesis,
theresultsindicatethatrelativeweight
to some extent,
losses of leaf litterdo indeedreflect,
of livingplants
featuresthatenhancethefunctioning
For
and theirleaves in theirnaturalenvironments.
instance,ecological strategymay explain the fast
decompositionof deciduous woody climbersand
in thisstudy.Comparedto self-supporting
scramblers
inheterogeneous
woodyplants,theyare seentypically
whichpromotea highleaf turnlightenvironments,
over as determinedby thechangeableavailabilityof
lightpatches(Castellanos 1992). In suchplantsnatural selectionmay have favouredshort-livedleaves
witha miniphoto-assimilation
equippedforefficient
(cf. Chapin 1980; Chabot
mumof defencechemistry
& Hicks 1982; Coley 1988). Such leaves would be
palatable to soil-bornedecomposers(Grime& Anderson 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
smalllitter
directlydemonstratedby theconsistently
weightlosses of woody evergreensas compared to
deciduous species. The latterrelationshiphas been
supportedby some earlierevidence,as reviewedby
Aerts (1995), but has never previouslybeen demonstratedacrossa representative
rangeofwoodylifetaxa.
formsand (bothbroad-leafedand needle-leafed)
The relationshipheld for a big slice of the woody
Sheffield
flora,whetheror not taxonomicrelatedness
was accountedfor.
largeinvestments
Long-livedleaves,withrelatively
in compoundsnot directlyinvolvedin photo-assimilation,generallyexhibita low specificleafarea (Reich
et al. 1992). At least among thewoody species,SLA
of litterwas closely correlatedwith that of living
leaves (J. H. C. Cornelissen,unpublisheddata). The
positivecorrelationbetweenlitterSLA and weight
sugloss in thewoody speciesin thisstudytherefore
and decompogestsa link betweenleaf functioning
sition. Mechanistically,this relationshipmay be
Leaves
chemistry.
explainedbyvariationin structural
witha smallspecificarea can be expectedto be physias
callytough(in termsof resistanceto penetration),
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
and
herbaceousleavestranslatesintothatofthelitter,
why the relationshipbetween litterSLA and despecies.
ratewasnotseenamongherbaceous
composition
The autumnleafcoloursin deciduouswoodyspecies may also in partreflectfunctionalfeaturesof the
living leaves. Brown coloration,for instance,representsmostlyphenoliccompounds such as lignins
and tanninsthatbecomeapparentin senescedleaves
into
once thegreenpigmentshave been transformed
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 assimilatingleaves, are substantiallydegradedbeforeleaf
fall(Hendryet al. 1987). Severalspeciesin thisstudy,
however,shedtheirleaveswhilestill(partially)green.
WiththeexceptionofHippophae(which,interestingly
in this context,is a nitrogenfixer),all eightnative
green
species that shed theirleaves predominantly
area on (moderately)productive
occurin theSheffield
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 structural compounds. In such leaves it mightpay off,
and withdrawing
ratherthangraduallytransforming
- a processmarkedby
theirphotosynthetic
chemistry
colour change- insteadcontinuingto produce new
up to themomentof leafabscission.
photosynthates
with
Such green,poorlyprotectedleaves,presumably
nutrients,
high contents of photosynthesis-related
would be palatable to decomposers. This would
explainin functionaltermswhygreenautumncolorationis an indicatorof fastleafweightloss in deciduous woody species,independentof taxonomy.Further study is needed to test whether the fast
decompositionof shed greenleaves is a consequence
in compoundsthatpromote
of thelack ofinvestment
or of highernutritional
value. It also
stress-tolerance
remains to be determinedwhethercoloration of
senescedleavescan be used to predictdecomposition
ratein herbaceousplants.Leaf colorationcan change
rapidlyduringsenescence(Sanger 1971)and thismay
581
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 correlatedwithtaxonomy.Planttaxa are probablyto an
important
extenttheoutcomeofmultipleadaptations
to the environment.
Some primitivetraitsthat have
remainedin higherplanttaxa are probablystilladaptive in the species' presentenvironmentsand may
thereforepartiallycoincide with the traitsof functional 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 comparativelysuccessfulin disturbedand productive
habitats(Hodgson 1986;Grime1986).It can therefore
be expectedthat'primitive'familieswitha low index
are moresuccessfulin environmentally
stressedhabitats. Since the leaves of plants in such habitatscan
be expectedto be well protectedand theirlitterto
decompose relativelyslowly (Grime & Anderson
1986),thepositivecorrelationbetweenSporne'sPercentageAdvancementIndex and relativeweightloss
in dicotfamiliesmaybe an evolutionary
as wellas an
ecologicalrelationship.
leaf habit,autumnfoliagecoloration,
Life-form,
taxonomyand evolutionaryadvancementindexconstitutespeciesfeatureswhichareeithereasyto observe
in thefieldor whichcan be obtainedfromtheliterature.In viewof theircorrelationswithleafdecompositionrate,thesefeaturesmay proveusefultools for
predictingdecompositionratesin vegetationsdifferingin 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 decompositionprocesses.He arguedthatshiftsin speciescompositionin 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
different
fromthepotentialleafdecompovegetations
sitionratesoftheircomponentspecies.It mayalso be
possible to inferpotentialdecompositionrates for
speciesnot includedin this studyfromsome of the
functionaland taxonomicfeaturesshownhereto be
correlatedwithdecompositionrate.
c 1996British
Ecological Society,
JournalofEcology,
84, 573-582
Acknowledgements
I thankGeorgeHendryforhisacademicand physical
contributionsduringvarious phases of the experi-
ment;JohnHodgson,Ken Thompsonand Sue Hillier
forprovidingherbaceousleafmaterial;StuartBand,
Suzanne Hubbard,Debbie Cornelissenforassistance
in thelab; JonathanKieltyforidentifying
soil invertebrates. 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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Taylor,B.R., Parkinson,D. (1988) Aspenand pineleaflitter
decompositionin laboratorymicrocosms.II. Interactions of temperatureand moisturelevel. Canadian
Taylor,B.R., Parkinson,D. & Parsons,W.F.J.(1989) Nitrogenand lignincontentas predictorsof litterdecayrates:
a microcosmtest.Ecology,70, 97-104.
Tietema,A. (1993) Mass loss and nitrogendynamicsin decomposingacid forestlitterin the Netherlandsat in20,45-62.
creasednitrogen
deposition.Biogeochemistry,
Van Vuuren,M.M.I., Berendse,F. & De Visser,W. (1993)
in thedecompositionof litSpecies and sitedifferences
tersand rootsfromwetheathlands.CanadianJournalof
Botany,71, 167-173.
Van Vuuren,M.M.I. (1992) Effectsofplantspecieson nutriPhD thesis,UtrechtUniversity,
entcyclinginheathlands.
The Netherlands.
Westoby,M., Leishman,M.R. & Lord, J.M. (1995) On
correction'.Journalof
the'phylogenetic
misinterpreting
Ecology,83, 531-534.
Wieder,R.K. & Lang, G.E. (1982) A critiqueof the analyticalmethodsused in examiningdecompositiondata
obtainedfromlitterbags. Ecology,63, 1636-1642.
Received24 August1995
revisedversionaccepted15 April1996
J. H. C. Cornelissen(1996). Leaf decompositionratesappendix I
The status(N = native,Nd = naturalised,
P = planted.)
is
ofthestudyspeciesandtheirleaflitter.
Characteristics
Gardenorprivategardenin
to Stace(1991). * Leaves collectedinUniversity's
according
Experimental
Stace 1991):T = tree,S = shrub(max.heightbetweenIm and 10 m),
habit(following
Life-form/leaf
Sheffield.
HM = herbaceous(graminoid)
SS = subshrub
(max.heightlessthan1 m),C = woodyclimberor scrambler,
e = evergreen.
HD = herbaceousdicot.d = deciduous,se = semi-evergreen,
Mean relativepetioledry
monocot,
initialleaf
weightis expressedas % ofwhole-leaf
dryweight.n.m.= notmeasured.Classesofpredominant
orB & Y
coloration
(deciduouswoodyspeciesonly):B = brown,BB = blackishbrown,BY = brown-yellow
YG = yellow-green
orY & G mixture,
G = green,MC =
Y = yellow,YR = orangeorY & R mixture,
mixture,
ofat leastyellow,redandgreen).
multi-colour
(mixture
LifeLitter
Litter
form %petiole
seiila
nta
and
dry
Native/naturalised/planted
leafinta
speii
leaf
weight (mm/mg) colour
Family
Woodyspecies
habit
[N
[Nd
](Td
][Td
[9
[Aceraceae
[Nd
d
16
11
I
Acercampestrej[Aceraceae
Acer
platanoides[Aceraceae
Acer
Aseudoplatanus [
Aesculus
P/Nd
Hippocastanaceae
hippocastanum
|N
Alnusglutinosa I[Betulaceae
Amelanchier [Rosaceae
lam arckii__
Andromeda
polifolia
_
_
_
_
_
_
[Ericaceae
_
_
_
_
_
unedo
Arbutus
ricaceae
Aucubajaponia
!Comnaceae
_
II
_
_
Betulaceae
Betulapendula |Betulaceae
|uddlejadavidi |Buddlejaceae
3uxus
i|Buxaceae
sempervirens
.1
Calluna1vulg
icaceae
betulusBetulaceae
Carpinus
Castaneasativa ||agaceae
Td
11
J[6
[*Nd
I.]
Td
8
|*N
SSe
2
Se ~
[]
*N
lP
*N
||
||d
]S*N
11
IN
|P(Nd)
d__
|d
|Sd
S3
SSe
135.3
BY
21.3
JIG
_
_
_
__
[YR
_
_
_
_
_
_
_
_
_
ZI7.3
1118.6
6
2.3
19
||16.1
4
|1
H
I [ ]
15
_
_
_
_
7.9
114
IIId
||d
YG
21.1
_
6Sd
SSd
25.9
][MC
11_1
jISe jI3
ISe
IIYR
1
17
__1
]|Td
Berberis
vulgarisBerberidaceae IN?
etula
nana
15
it
]22.8
]j18.8
16.7
|11.4
15.7
_
_
______
1!
MC
IBY
1Y
IMC
||.
10.8
21
|28.3
_
G
IB
1
Corylusavellana ||Betulaceae
Cotoneaster
Rosaceae
horizontalis
Crataegus
monogyna
JjN
*Nd
[[Td
FSSd
N
[Sd
1130.8
14
IIY_
IMC
1111111111
1Rosaceae
11
Cytisusscoparius ]|Fabaceae
J|N
DryasoctopetalaJ[Rosaceae
J[*N
Empetrum
||Empetraceae
|N
Ericacinerea
Buonymus
europaeus
Fagussylvatica
Ficuscarica
j[Ericaceae
S[N
n_i_gT_____um
114
F6
[Celastraceae
I.
[9
11
20.4
[1
[Se
|SSe
J[0
16
||n.m.
13.4
SS
6
18.4
J16
112.8
I[Sse
N
Sd
6
20.4
YR
=
I___
||
1_
_____
MC
I_
11
1
|[Fagaceae
J[N
I[Td
J[Moraceae
I*Nd
J[5
j[Td
I1
11
[9.1
][24.6
[Oleaceae
[N
[Td
|14
]|16.6
__G_
[Onagraceae
[*P/Nd
1111
[Sd
7
][27.1
1
]{MC
[Se
0
111.3
I|N
|Ce
(14
]|14.4
[N
]|SSe
7
1111
120.7
||
][Sd
i
3
1
]
11
J[SSe
[o
112.7
11._
Ilex aquifolium ][Aquifoliaceae j[N
]Te
_
__5
Juniperus
rxcelsior
FFuchsia
chsia[
magellanica
Hebex
franciscana
Hederahelix
Hippophae
Hypericum
calycinum
Laburnum
anagyroides
Scrophulariaceae*Nd
1111
[Araliaceae
'[Ele
Elaeagnaceae
ir
11
[*N
Hypericaceae 1*P/Nd
11111111
11-
11
][BY
_IG
_
11
17.9
]15.7
IIG
_____
11
j[Cupressaceae
[*N
ISe
[o
113
1___
IFabaceae
I*P/Nd
][Td
[15
127.1
IIYG
IIP
|]Td
I[0
1115.5
]IYR
j[*p
][Td
]120.4
]IYR
]4.7
||.
111111111
Larixdecidua j[Pinaceae
Larixkaempferij[Pinaceae
Ligustrum
OlOeaceae
ovalifolium
11111
Lgustrum
|Oleaceae
'LOniCera
11
1111111111
[elianthemum Cistaceae
nummularium 111IL
Ihppophae
rhamnoydes
_
I
IIP
||N
Caprifoliacac NIM
1[o
13
__Se
.1111
Sd
ld
1331.3
j4S
1R
G__
MalussylvestrisRosaceae
Piceasitchensis Pinaceae
Pinus
sylvestris|Pinaceae
Populusigra Salicaceae
tremula|Salicaceae
Populus
Prunus
avium |Rosaceae
jN
p
|N|Te
|N/P
|N|Td
|
'runus
jRosaceae
laurocerasus 11
Prunus
lusitanica|Rosaceae
runuspadus |Rosaceae
ilex
Quercus
*Nd
|*P(Nd)
|N|Td
Nd
1*P/Nd
!Fagaceae
u1erCUs
Petraea FagaCeaeN
robur |Fagaceae
|Quercus
tharticus
cahamnius
;hododendron
ponticum
j[Ericaceae
1
||
|N|Td
|N
Sambucusnigra
CaprifoliaceaelN
dulcamara
[Solanaceae
j[N
*N
orbusaucupa |saceae
Carifoliaceae
ISymphoricarpos
18
120.1
Sd
Td
IIe
Td
|6
|12
| Z6
I0
17
6
||.8
.
|43.2IYI
| U7.7
10.O8
149
*Nd
115.7
J59
1111111
Sd 181121[2
LIIMc
iSd
19
1Z3
|Sd
{Sd
|Sd
|13
118
|12
|28
120.2
lTd
]ISd
SCd
I|Td
|29.3IY
__
IIY I
IYI
14.5
[8
|d
iYR
1|.
][Se
Sd
|YG I
18.7
1
38.5
[1]____
|N|Sd
|NISCse
IN
Rosaceae
|16
Sd
Ribesfnigrum Grossulariaceae*Nd?
|R.besrubrum |Grossulariaceae||d
Ribes
uva-crispaIGrossulariaceaeI*N?
Rosaarvensis !|osaceae
aria
Sorbus
|13
Y
13.4
N
IN]s
[*Nd
Salix
fragilis Salicaceae
15.61
Se
|Te
Z.71
11
12
MC
|23.3
14
Ribesalpinum
|Grossulariaceae|N
||osa canina |Rosaceae
|Rubusfruticosus
||osaceae
Rubusidaeus Rosaceae
|Salixalba
|Salicaceae
Saixcaprea
|Salicaceae
[
|d
11
11
9
||
]______
Rhamnaceae
d
Id
IK!Z
'runusspinos Rosaceae
yruspyraste Rosaceae
Td
Te
-
B
|IBI
1IG
___
||
I
_
I
l
!I
18
1|5.9
123.1
IMC I
14
114.9
17
||15.9
|GI
117
125.8
I
[43.6
IxY
X17.4
[
||YR
113
11
30.4
E15ZIiIY
111
ji 1
Z7
Td
II22
Sd
3
15.4
1
y
|M
By
I
I
Tamarix
gallica Tamaricaceae *P(Nd)
n.m.IYG
axusbaccata
Taxaceae
Te
0
)olytrichus
Lamiaceae!N
SSe
12
!3.
Tiliacordata
iliaceaeN
Td
10
24.3
Ulmusglabra
accinium
Ulmaceae
Ericaceae
Se
|| Z XM.I
1EinImI1ZIZ
|Td
||2
|N
N
___
]
vitis- Ericaceae
Vaccinium
idaea
i
11
Viburnum_lantana
ICaprifoliaceae ]
Se
SSd
4_1
SSe
3
]ISd
1]7
Viburnum
opulusIlCaprifoliaceae ]uN
Herbaceous
species
capi11arisPoaceae
Agrostis
]uN
Anthoxanthum Poaceae
iN
odoratum
1
_______
|Alvsthriscs
Apiaceae
Aylhenaterum 11
Arrhenaterum Poaceae
elatius
HM
0
31.7
HD
][n.m.
][N
]HM
0|o36.1
--------
]1
]
rizamedia
IPoaceae
][N
]uHM 10
jentaurea
nerastium
fontanum111
Chamerion
angustifolium
al1um
Conyza
Asteraceae
]N
[N
[Caryophyllaceae
Onagraceae
111
[N
Chenopodiaceae N
Asteraceae
Nd
]IHM
[0
[0
-
___
1[22
1|20.9
1120.3
n.m.
19.3
[HD
0
26.3
[1
23.3
[Ii
21.4
1
HD
___
33.4
HD
][HD
1nl
|HD
11
__G_
]_39
|Poaceae
][N
YR
_____
____
]
]uHM
_
|IY
7.9
]131.2
i
BY
19
]1o
1
BY
1I
]IHM
][N
][N
23.4
____
7.6
pinnatum
BromopsiserectaIPoaceae
Carexflacca
]Ccyperaceae
||29.8
1__
_9
ISd
___________]___
11-----
IEmI
lTd 112
Tiliaceae |(N)/P
Tiliaxvulgaris
Ulexeuropaeus abaceae
Fabaceae
Ulexgallii
6.5
29.3
11
11
1!
J_
!
f.
I
_
1
_
[
_
_
Dactylis
glomerata
Deschampsia
flexuosaI
Poaceae
N
HM
Poaceae
N
HM
26.3
0
][o_____
[21
I[29.5
J
Digitalis
purpUrea][Scrophulariaceae
]IHD
]
Epilobium
[
HD
FI
][
IN
][HM
11.9
]_01_____
N
][HM
Onagraceae
hirsutum
_
Eriophorum
vaginatum
I
_
Cyperaceae
Festucaovina
j|Poaceae
IPoaceae
estucarubra
Galiumaparine ][Rubiaceae
Helianthus
Atrca
annuus
Helictotrichon
Poaceae
Poaceae
Holusan]us
PaeeliEi
|Koeleria
macrnt]aPoaceae
Leontodon
Asteraceae
N
11
I
11
][HM
[N
]|HD
[N
IN
[HD
i
I|N
][HM
[N
[HD
[N
HD
IHD
Plantago
1111111
rHD
[NRosaceae
rtc
doca
Utiaea.
I1
JIo
1130.2
5I3
][35.3
1
___
|1I
1!
I
_
122.3
][23.4
][21.1
1I
_
11
__
11
_
11
HD
_
][n.m.
620
26
.r
11
I
20.1
1.9
_
]1_ I
15
1_
1_8.4
N
HD
___
25.1
J[4
I[HM 1[?
j[HD
]16.9
125.6
k
1
]39.9____
3
11l
J[N
RumexacetosellaPolygonaceae[N
1[28.4
]22.2
]HM
[N
Rubus
[O
[N
1
chamaemorus I
l
J[
[
IffiCinarUm
!
||Poaceae
[0
]HM
111
Poa annua
23
02.
[N]HM
iPlantaginaceae [N
23.9
111
pH
Loliumperenne ]|Poaceae
Lotus
[Fabaceae
|o___latus
comniculatus
Origanum
vulgare|[Lamiaceae
Pilosella
IAsteraceae
lanceolata
l
Zf
I
__
_
J. H. C. Cornelissen (1996). Leaf decompositionratesappendix2
as % ofinitialdryweightin fivetreatments
Dryweightloss ofleaflitter
(see text).In thespeciesmarkedwith
*, samplesweighing
0.5 g wereused,and% dryweightloss was multiplied
bya factorof 1.11,as derivedfrom
withBetulapendula(1.16 timesmoreweightloss in 0.5 vs. 1 g samplesin W8, 1.13times
a controlexperiment
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
W8F: (1) 0-12(2) 12-24(3) 24-36(4) 36-48(5) 48-60%
weightloss classes(see text)werein eachtreatment:
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) 6885 % 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.
iZ
I
S8F
||W20C|i
*LIiIZ n
St.error
MeanSt.errorMean St.errorMean St.errorMeanSt.errorl
class
classMean
--|WTreatment
l
ILI
*IiiW20FIL
* L
Woodyspecies
Liii
LI]Z
1][
251190.72
0.77 j|1.74
45.7 0.85
41.1
Acercampestre
15.2 0.96
18
1136.3 |1.7 II42
18
4i I
.81
A,cerplatanoides
2.5 3.13 553 111.54 J[81.3_4.13
Acerpseudoplatanus40.3_1.65
Aesculus
i139
3 H4O117
hippocastanum
I
15.7 1.9
14.7 1.6
A.lnus
139.5 |1.35
glutinosa
Amelanchierlamar 5.20.4
|6615.72
116.6 10.54
Andromeda1polifo5 056
rbutusunedoEfi
EZi
:
Aucubajaponica
|Berberis
vulgarils
Betula1nana
35.2|0.52
17.20 jE6
etulapendula 2 .60.42
471
23.9 1[3737.7 1.0
177.3 |1.85
1124.1
EZZZLjEZE
I
2.07
=lZ
122.711.6
56.5
||46
][.
i1l
1
1
Callunavulgaris
IC
C'arpinusbetulus
Castanea
sativa
ClematisIvitalba
Cornus
sanguinea
avellana
ICorylus
.210.57
6_
I _1.35
371.35
15.3
129.911.0
8.1
13.71.35
125.5 095
20.2 :l
1.17
|Cotoneaster
horizontalis
11.4
I1E|.05
08
117.62
C
1
|
|IL
1134.2111.07 I49I7
1.3
73512.74
31.3
||35.6
33
I|6I73
|ZI
r
il m iL
11 11.4
134
111Z
1 1.4
13 .0-8
octopetala* j5S.3 [67.3
|Dryas
1[|
1|~
8||30.48
I.
1.
3.5
1
J[ II21 I
I3
1
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2
1
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IZIII
LI1||1.81
I
4.5
6
34.5S9
154 19.1
rCategusmongyna
1j26 fj31 117.31[4Z IIiL IZr 188.811j.6
11
] |1Z 5
97.31121
1p89|0.41
4
|IiiI
2II
I 62iI.29
11
I
I14
|86.1 |2.88
7E6.54
|71.6 |2.8
26.3 860 401 1.56
L
_
94 1I49 ]LZ i1I I] IIZi
11127.7
1122. 0.79
17.90.5
]
1.i.8
Buddlej
2 142.7282 183.28.1
adavidii 22 1.39 137.218
B uxussempervirens*
115.7 0.?95
] I2Ii
.
i7.49
10.64E IZ
.
.11._I
][2.5
I I
01.49
j73 j37.9
1.4
J
1.r
j1
[1
14-6
2.51g
m
2.5
13y
yy
7]
11
I
10.31 I.
nigrum* 117.2
Empetrum
Erica
cinerea
1131 0.53
Buonymus
europaeus [391 1.03
Fagussylvatica
Ficuscarica
E18.30.31
56.8 1.71
Fraxinusexcelsior 33
nummularium*
l
83.32.86
|98.6 |0.22
86.7 4.03
1 116
[
36
__2
10.34 17.3 099
Hypericum
calycinum iI3.1I0.41 114.7
|Hippophaerhamnoides
3 1.1
4][10.64
14150.65
| 1xaquifolium
communis5EZ 0.22
Juniperus
1
|Labumnumanagyroide5.8 0.91
71.4
Larix decidua
15j490.32
54
J 3
4014.87
Pi.ceasitchensis
1E20.6
|Pinussylvestris
Po:pulusnigra
1pulustremula
P|runusavium
27034
51E
|36.7 4.09
|E
33.81.34
31.5 1.31
271421.36
4?3.61.61
32fi6 0.72
28.9 1.43
pyraster
Pyrus
7.4 0.52
Quercusilex
13.20.35
Quercuspetraea
8.4 0.63
robur
Quercus
catharticus 4821.79
Rhamnnus
1hododendron
936.0778
I~ 0.78cu
~
~ ~~.
ir
l|E|0.32
1125
8.1
72|.4 3
60.49.52
361.7 5
.9 1.56
13.90.44
---9.60.36
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l
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le
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3.5IlI
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44.6 118.47 33 1.24
112.6 2.31
II
II
1142.7 1i0.94 |J75.8115.61
||7222.13
151.5 11.87
|52
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15.1 1.07
|26.3 10.74
116.3 11.13
12.11
6
6.9 1.07 11652 61
]j111 111071 16. 1.1
195.3 |1.46
1957 13.94
185./ 155
150.9
|25
|1.83
1116.5 11.36
11
1
1661
116.2
Z
114.
1
95.2 ||2.19
|
2I
33.8 0.8
G
]_
2.8
1[I0.
1
][
Z
[
21
]2.1
3
36.70.6
Z2IiZ
6p.06
1
.1
|98.1 10.47
|63.2
1.5
]
10.71
5.5 |1.46
3m0.70.03
LI
41.5114.06 1154.5I 1.48
15.210.43
30 2.05
11
||2||
56.812.23
25.2 4.22
runus
laurocerasus 17.4 0.69
|62.9 |4.41
10Z34
||53.8 13.57
80.6
1Z31
20j7i0.36
1runuslusitanica
'runus
padus
Prunus
spinosa
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An Experimental Comparison of Leaf Decomposition Rates

Transcription

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