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Soil organic matter in European forest floors in relationto stand characteristics and environmental factorsAnnemieke I. Gärdenäs aa Department of Soil Sciences , Swedish University of Agricultural Sciences , P.O. Box 7014,Uppsala, S‐750 07, SwedenPublished online: 15 Dec 2008.

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Scand. J. For. Res. 13: 274-283, 1998

Soil Organic Matter in European Forest Floors in Relation toStand Characteristics and Environmental Factors

ANNEMIEKE I. GÄRDENÄSDepartment of Soil Sciences, P.O. Box 7014, Swedish University of Agricultural Sciences, S-750 07 Uppsala, Sweden

Scandinavian Journalof Forest Research

Gärdenäs, A. I. (Swedish University of Agricultural Sciences, Department of Soil Sciences,P.O. Box 7014, S-750 07 Uppsala, Sweden). Soil organic matter in European forest floors inrelation to stand characteristics and environmental factors. Received April 11, 1997. AcceptedFeb. 2, 1998. Scand. J. For. Res. 13: 274-283, 1998.

Published data on the quantities of organic matter in the forest floor (SOMfl) of Europeanforest stands were collated. Studies of SOM f l were included if the total sampled surface was atleast 0.15 m2, the surface of a single sample was at least 100 cm2, live material was sorted out,and ash or carbon content was determined. Data from 59 forest stands were compared withregard to stand characteristics and environment. Using a single variable, tree genus was mostimportant for the amount of SOMfl (R2

adj = 0.34). The amounts in spruce stands (41 × 103 kgha - 1 ) were significantly higher than those in larch, Douglas fir, oak and birch stands(4-11 × 103 kg h a - 1 ) . The best significant multiple model was genus combined with stand ageand basal area (R2

adj. = 0.72). The importance of litter quantity and quality for differentiatingthe tree genera is discussed. Key words: boreal, humus layer, litter, stand age, temperate, treegenus.

INTRODUCTION

Soil organic matter (SOM) in boreal and temperateforest systems plays a key role in the storage ofcarbon (Lugo & Wisniewski 1992, Schimel 1995).However, the balance between gains and losses of Cin forest ecosystems is vulnerable, and can be dis-turbed by, for example, intensive forestry practice,acid rain, high deposition of N and heavy metals aswell as climate change (Johnson 1992, Raich &Schlesinger 1992). Still, the effect of climate changeon SOM in temperate and boreal forest ecosystems isvery uncertain (Anderson 1991) and an increasedinterest in SOM is therefore justifiable.

Litterfall from trees is an important source forSOM in forest ecosystems. Quantity and quality oflitterfall vary with tree species, geographical positionand stand age (Albrektson 1988, Reurslag & Berg1993, Berg et al. 1995a). Ovington (1959) showedthat the amounts of SOM in forest floor (SOM,,) arerelated to stand age and Berg et al. (19956) tried toestimate the amount of SOM for Scots pine standsusing litterfall and litter quality.

The aim of this study was to synthesize data onSOMn amounts in European forest stands and toanalyse the importance of stand characteristics andenvironmental factors. The forest floor is defined asthe litter layer on top of mineral soil, i.e. the com-bined L-, F- and H-layers, also called the O-horizon

or the An-horizon. The forest floor is mostly a minorlayer of a soil profile, but with the highest concentra-tion of organic matter. The forest floor is believed tobest reflect the present climate and vegetation, and istherefore most suitable for comparing and analysingeffects of stand characteristics and environmental fac-tors.

METHODS

Published data on amounts of organic matter in theforest floor in European forest ecosystems were in-cluded when they met the following criteria: (/) thesurface area of each sample was at least 100 cm2, (//')the total sampled area was at least 0.15 m2, (///') livingmaterial such as ground vegetation had been re-moved, and (iv) the ash and/or C-content was deter-mined (Table 1).

Ashfree amounts of SOMH data were used forcomparison. For a few sites (Grunewald, SantaColoma, Soenderskov, Store Dyrehav, Sävar, TistedNoerskov, Vallgoguina), the amount of ashfreeSOMH was estimated by multiplying the measuredC-content by a factor of 2, i.e. assuming a C-contentof the organic material of 50% (Schnitzer & Khan1972).

Single and multiple regression (SAS procedures"GLM" and "STEP", Anon. 1988, 1995) were usedto analyse the influence of annual precipitation andmean annual temperature, geographical position, alti-

© 1998 Scandinavian University Press. ISSN 0282-7581

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Scand. J. For. Res. 13 (1998) SO M in European forest floors 275

Table 1. Sampling

Total area sampled(m2)

10.4-3.750.6-0.80.180.4-110.41.7-3.752.51.080.750.61-21

and analytical methods

Sampling month

July-Aug.nmApril-Nov.AutumnJuly-Aug.MayMay-Sept.JulyJulyJanuaryApril, Nov.OctoberAugust

Drying temp.(°Q

8085-105

10570

105 '1058585

1054080

nm105

Loss on ignition(°C and h)

800, 2 h600, 2 h600, nm500, nmnm, nm*600, 2 h550, nm550, 2 h*nm, nm500-600*

Literature reference

(1)(2)(3)(4)(5)(6)(7)(8)(9)(10)(11)(12)(13)

nm = not mentioned.* No loss on ignition, but a C determination with CN analyser Carlo Erba Na 1500 or a Leco CHN 1000 Analyser.(t) Ovington (1954); (2) Berg et al. (1993a); (3) Persson & Wirén (1995); (4) Williams (1983); (5) Nykvist (1971); (6)Marschner & Wilczynski (1991); (7) Staaf & Berg (1977); (8) Nihlgârd (1972); (9) Kubin (1983); (10) Vesterdal et al. (1995);(11) Parkinson et al. (1980); (12) Matzner (1988); (13) Alriksson & Eriksson (1998).

tude, soil texture, and stand age on amount of SOM,,.Moreover, the effects of tree genus and of coniferousversus deciduous species were analysed. Forest standswith the dominant species belonging to the samegenus were grouped according to the taxonomy givenby Mitchell (1977). The statistical analysis of genus

Harads• Oulanka

• Sävar

Jadraás

Garpenberg

SkogabyT. Noerskov I Farabol

• •HasslovSt. Dyreha\WKonqalund

•SoertderskovWest ToftsF°99ebotzand

• t... » ' ^ • • *tf3runewaldAbbotswooob • Soiling

Bedgeburry

Fetteresso»

effect was limited to the genera with at least twostands, i.e. pine (Pinus), spruce (Picea), larch (Lari.x),Douglas fir (Pseudotsugd), beech (Fagus), oak (Quer-cits) and birch {Betula), in order to keep the numberof classes low (Table 2). Soil texture was grouped intosand, loam or clay. The tree genus and soil textureclasses were compared using the Duncan test andenvironmental factors with principal components(SAS option "Duncan" and procedure "Princomp",Anon. 1988, 1995). For the spruce class, the analysiswas repeated to see whether the factors could differ-entiate within the spruce stands. For comparison ofthe R2, ^justed w a s u s e d to correct for the differencein number of observations and explaining variables.# Ljmted was calculated as

(n-p)(1)

where n is the number of observations and p thenumber of degrees of freedom.

El Raso -Santa CoFomaïR a s o

#*Vallgorguîna

•Furadouro

»DonanaFig. 1. Approximate locations of the sites.

STAND DESCRIPTION AND COMPARISONOF METHODS

Of the 59 stands included, 18 were of Norway spruce(Picea abies (L.) Karst.) and only 11 of deciduousspecies (Table 2). Most of the sites are located near asea or the Atlantic Ocean. As a consequence, the

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276 A. I.

Table 2. Tree

Tree generaEnglish name

PinesPines

Pines

PinesPinesPinesPinesSprucesSprucesSprucesLarchesLarchesLarchesLarchesDouglas firsFirsHemlocksRed cedarsCypresses

OaksOaksOaksOaksBeechBirchesBirchesSweet chesnutAlders

Gärdenäs

species and their genera,

Tree species

English name

Scots pineCorsican pine

Austrian pine

Stone pineMaritime pineMonterey pineLodgepole pineNorway spruceSerbian spruceSitka spruceJapanese larchA hybrid larchEuropean larchSiberian larchDouglas firGrand firWestern hemlockWestern red cedarLawson's cypress •

Sessile oakNorthern Red oakEnglish oakOak, unspecifiedCommon beechWhite birchSilver birchSweet chestnutGrey alder

Scand. J. For. Res. 13 (1998)

average ashfree SOMß amounts and number of stands

Scientific name

Pinus sylvestrisPinus nigra varcorsicanaPinus nigra varaustríacaPinus pineaPinus pinasterPinus radiataPinus contortaPicea abiesPicea omorikaPicea sitchensisLarix leptolepsisLarix sp.Larix deciduaLarix sibiricaPseudotsuga taxifoliaAbies grandisTsuga heterophyllaThuja plicataChamaecyparislawsonianaQuercus petraeQuercus rubraQuercus roburQuercus sp.Fagus sylvaticaBetula verrucosaBetula péndulaCastanea sativaAlnus incana

- Average and (SD)(kg 103 ha"1)

26.418.2

26.0

13.322.922.914.642.220.931.910.77.7

31.413.710.18.49.99.6

10.1

4.73.13.14.1

19.23.59.53.43.6

(19.6)(3.9)

(7.6)

(23.8) •

(5.1)

(22.0)

No ofstands

53

1

2231

1811111131111

I11131111

Literaturereference

(1, 2, 6, 8, 13)(1)

(2)

(2)(2)(2)(H)(1, 2, 3, 5, 9-13)(1)(1)(1)(1)(1)(13)(1)(1)(1)(1)(1)

(1)(1)(1)(1)(1,9, 13)(U(13)(1)(1)

(I) Ovington (1954); (2) Berg et al. (1993a); (3) Persson & Wirén (1995); (4) Williams (1983); (5) Nykvist (1971); (6)Marschner & Wilczynski (1991); (7) Staaf & Berg (1977); (8) Nihlgârd (1972); (9) Kubin (1983); (10) Vesterdal et al. (1995);(II) Parkinson et al. (1980); (12) Matzner (1988); (13) Alriksson & Eriksson (1998).

variation in mean annual temperature (—0.9 to16.6°C) and annual precipitation (400 to 1150 mm)were not as large as one might expect within Eu-rope (Fig. 1, Table 3). The most common soil typeswere Podzols and Cambisols. Further stand descrip-tions and references to the original site descriptionsare given in Table 3.

Comparing the methods used to sample SOMfl, itwas evident that there was a large variety in samplesurface (100-5000 cm2), number of replicates (4-44) and thereby in the total sampled area (Table 1).

Staaf & Berg (1977) and Matzner (1988) sampledsystematically along a transect or grid system, whileothers used a random approach. Some of the sam-plings were made before the main litterfall, for ex-ample, Ovington (1954), Nykvist (1971) and Kubin(1983)—others just after the main litterfall—suchas those of Williams (1983) and Matzner (1988).Two different studies were performed at the standSoiling II, one by Parkinson et al. (1980) in thewinter 1976/77, the other by Matzner (1988) duringthe years 1966 to 1983. In the statistical

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Scand. J. For. Res. 13 (1998)

Table 3. Some stand characteristics

Stand

AbbotswoodBedgeburryDoñanaEl Raso IEl Raso IIFarabolFetteressoFuradouro IFuradouro IIGarpenberg IGarpenberg IIGrunewaldHaradsHasslövJädraas Ih VKongalund IKongalund IIOulankaRoggebotzandSanta ColomaSkogabySoenderskovSoiling ISoiling liaSoiling libSoiling IIISoiling IVStore DyrehavSävarT. NoerskovVallgorguinaWest Tofts

Lat./longitude

51°49'N;02°30'W51°05'N;00°27'E38°07' N;06°12' W41°47'N;05°26'W41°47' N;05°26' W56°25'N;14°35'E56°58'N;02°15'W39°25'N;09°15'W39°25' N;09°15' W60°16'N;16°13'E6O°16'N;16°15'E52°28'N;13°14'E66°08' N;20°53' E56°24'N;13°00'E60°49'N;16°30'E55°59'N;13°10'E55°59'N;13°10'E66°22'N;29°15'E52°34' N;05°47' E41°52' N;02°38' E56°33'N;13°13'E54°44'N;11°46'E51°45'N;09°35'E51°45'N;09°35'E51°45'N;09°35'E51°45'N;09°35'E5t°45'N;09°35'E55°54'N;12°22'E63°64' N;02°30' E56°47'N;10°0rE41°40' N;02°30'-E52°3O' N;00°43' E

Alt.(m)

213762

7607601402008080

2101853058

190185120120357- 520010510

390505500440500651055

26031

Meantemp.

(°O

10.99.7

16.611.411.46.38.3

15.215.25.04.08.91.36.53.86-76-7

- 0 . 910.315.67.08.26.26.26.46.26.47.62.97.4

13.69.5

Ann.prec.(mm)

765839557400400680528607607595650580650

1100609800800541826750

1150690

10391039103210391032660662710736517

Tree species

see comments*see comments^Stone pineStone pineMaritime pineNorway spruceSitka spruceMonterey pineMaritime pineNorway spruceNorway spruceScots pineScots pineNorway spruceScots pineCommon beechNorway spruceNorway spruceAustrian pineMonterey pineNorway spruceNorway spruceNorway spruceNorway spruceNorway spruceNorway spruceCommon beechNorway sprucesee comments1

Norway spruceMonterey pinesee commentsf

SO M in European forest floors 277

Standage(y)

23-4617-224430-4030-407127232345-123

12340

12941

13045-13055

25033193049398797

1151324927493121

Basalarea(m2 ha"1)

n.g.n.g.n.g.n.g.n.g.n.g.n.g.n.g.n.g.

29.8n.g.n.g.

9.736.715.031.455.620.0

n.g.n.g.

32.655.135.544.8

n.g.37.7

n.g.45.625-3252.5

n.g.n.g.

Soil texture

sandy loamsilty claysandn.g.n.g.silty tilln.g.sandsandsandy tillsandy tillsandclayey tillfine sandsandy loamfine sandn.g.sandy tillsandn.g.n.g.n.g.n.g.n.g.n.g.n.g.n.g.sandloamloamn.g.sand

Literaturereference

(la,b,c)(la.b.c)(2)(2)(2)(3a,b)(4,1c)(2)(2)(5a,b)(2)(6a,b)(2)(3a,c,d)(7a,b)(8a,b)(8a,b)(9a,b)(2)(2)(3a,e)(10)(lla.b)(lla.b)(12,11b)(lla.b)(12,11b)(10)(13)(10)(2)(la,b,c)

* Abbotswood; Scots pine, Corsican pine, Norway spruce, European larch, Douglas fir, Grand fir. Common beech, Sweetchestnut, English oak and an unspecified oak sp.t Bedgeburry; Corsican pine, Norway spruce, Serbian spruce, a hybrid larch, Douglas fir. Grand fir, Lawson's cypress,Western hemlock. Western red cedar. Sessile oak and Northern red oak.í Sävar; Norway spruce, Scots pine, lodgepole pine, silver birch and Siberian larch.f West Tofts; Corsican pine, Japanese larch, Douglas fir. White birch and Grey alder.n.g. not given.(1) a: Ovington (1954), b: Ovington (1953), c: Anon. (1989a); (2) Berg et al. (1993a); (3) a: Persson & Wirén (1995), b:Hallbäcken & Popovic (1985), c: Popovic pers. comm., d: Andersson pers. comm., e: Bergholm et al. (1995); (4) Williams(1983); (5) a: Nykvist (1971), b: Anon. (19896); (6) a: Marschner & Wilczynski (1991), b: Marschner et al. (1992); (7) a:Staaf & Berg (1977), b: Axelsson & Brâkenhielm (1980); (8) a: Nihlgârd (1972), b: Nihlgârd (1971); (9) a: Kubin (1983), b:Havas & Kubin (1983); (10) Vesterdal et al. (1995); (11) a: Parkinson et al. (1980), b: Bredenmeier et al. (1995); (12) Matzner(1988); (13) Alriksson & Eriksson (1998).

analyses, only the sampling in 1979 by Matzner (1988)was used.

Laboratory standards differing in the choice oftemperature for analysing water and ash content (Table

1) could result in systematic errors (Donkin 1990).Naturally, these differences in sampling methods andlaboratory standards give systematic errors, but for theevaluation these discrepancies were disregarded.

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278 A. I. Gär denos Scand. J. For. Res. 13 (1998)

Table 4. Regression between

Regression

All standsSimple linear modelGenusConiferous vs. deciduousLongitudeLatitudeAltitudeStand ageBasal areaTexture

Exponential modelTemperature

Multiple linear modelGenus, ageGenus, age, basal areaGenus, longitudeGenus, temperatureLong, alt.Long, alt, genusLong, lat, genus

Spruce standsSimple linear modelLongitudeAltitudeStand ageBasal area

Exponential modelTemperature

Multiple linear modelAge, basal areaAge, temperatureAge, temp., prec.

the ashfree SOMß

R2 adjusted

0.340.120.300.00.70.270.00.2

0.7

0.500.720.440.00.400.490.47

0.290.260.570.42

0.49

0.760.630.66

amounts

n

5050505050501645

50

50165050505050

1717179

17

91717

and stand

P<

0.00030.00690.0001

n.s.0.03150.0001

n.s.n.s.

n.s.

0.00010.00420.0001

*0.00030.0001

*0.0001*0.0001

0.01520.02030.00030.0360

0.0010

0.0069*0.0004*0.00U

characteristics

Intercept

n.d.n.d.19.191.32

18.8810.0022.24n.d.

0.06*

n.d.n.d.n.d.n.d.13.25n.d.n.d.

25.624.411.798.98

Ó.193

68.448.531.8

and environmental factors

Slope

n.d.n.d.

1.420.440.030.300.24

n.d.

40.12s

n.d.n.d.n.d.n.d.

1.45, 0.03n.d.n.d.

1.70.070.47

-1 .33

147.1s

1.42, 0.36, -0 .030.31, -3.730.33, -3.37, 0.02

Conifer vs. deciduous = comparison of coniferous versus deciduous species.n.d. = not determinable because classed variables are included.n.s. = not significant.* = one or more explaining variables not significant.° and s = the first and second constant of the exponentional model.

RESULTS

Tree genus explained best the variation in SOMfl

amounts using a single variable (Rlá¡ = 0.34, Table4). The amounts found in spruce stands were signifi-cantly higher than those in larch, Douglas fir (Pseu-dotsuga taxifolia [Lamb.] Brit.), oak and birch stands.SOMfl in Norway spruce forests had a mean value of42.2 x 103 kg ha"1 and ranged from 10.9 x 103 to95.5 x 103 kg ha"1 (Table 2, Fig. 2). The lowest valuefor Norway spruce stands belonged to the reforesta-

tion stand at Sävar in northern Sweden, and thehighest to a stand in Denmark (Store Dyrehav). Theaverage amount in Scots pine {Pinus sylvestris L.)stands differed little from that of the other pinestands (Table 2). Of the Scots pine stands, the one atGrunewald had extremely high amounts of SOMfl;excluding this stand would reduce the SD (Table 2) toless than one-third. For other conifers such as differ-ent larch species, Douglas fir and Lawson's cypress(Chamaecyparis lawsonia (Murr.) Pariatore), the

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Scand. J. For. Res. 13 (1998) SO M in European forest floors 279

so

Fig. 2. Average ashfree SOMfl amounts related to treegenus.

amounts of SOMH varied between .7.7 x 103 and31.4 x 103 kg ha".1 (Table 2).

The SOMfl-amounts found in the combined class ofall coniferous genus were significantly higher thanthose in the combined class of all deciduous genus(Table 4), but the comparison was somewhat biasedbecause the deciduous stands were few in number.Common beech (Fagus sylvatica) deviated from theother deciduous species, birch, oak, Grey alder (Alnasinccina) and Sweet chestnut {Castanea sativa), with amean value of 19.2 x 103 kg ha"1 and a standarddeviation of 22 x 103 kg h a ' 1 (Table 2. Fig. 2).

Stand age explained 16% of the variation in SOMn

amounts using all data. The degree of determinationincreased to 27% when the strongly deviated sprucestands at Oulanka, Sävar and Store Dyrehav wereexcluded (Fig. 3a). Longitude was positively relatedto the amount of SOM,,, explaining 30% of thevariation (Table 4, Fig. 36), and was strongly corre-lated to temperature ( — 0.7) and stand age (0.7).Annual precipitation and mean annual temperaturedid not shown a linear response model. SOMH had a

tendency to decrease exponential with mean annualtemperature. Latitude and basal area were of noimportance as a single explaining variable.

The multiple model of genus combined with standage and basal area was the most robust and signifi-cant model (/?;dj. = 0.72, Table 4).

Within the spruce class, stand age was best atdifferentiating (Rl^, = 0.57, Fig. 4a)- Combined withbasal area, a significant model was produced (Table4). Basal area as a single variable had a negativeslope (Fig 46). This could be interpreted that at siteswith high basal area, turnover rates are also high orthat another factor, like stand age, is dominating.When using the multiple model age, basal area andproduct of these variables, basal area got a weakpositive coefficient. As found by Vesterdal et al.(1995) when comparing C amounts in SOMH at threeNorway spruce sites in Denmark (Soenderskov, StoreDyrehav and T. Noerskov) with different classes ofbasal area, the difference between the sites is mostimportant, but within a site, C-SOM,, is weakly posi-tively related to basal area.

Amounts of SOMn of spruce stands were fitted to anegative exponential temperature model (/?;dj. = 0.49,Fig. 4c). Stand age, together with temperature orwith temperature and precipitation, gave an alterna-tive model with a somewhat lower /?.̂ Uj than themodel of stand age and basal area had.

DISCUSSION

Johnson (1992) stated that the effect of tree specieson SOM is often significant, but inconsistent. Hereferred to studies in which two or four species werecompared on a few sites e.g. Alban ( 1982) and Turner& Kelly (1985). In the present study, seven generaclasses and 22 sites were used and tree genus was

100

^ GO

% 4 0

1 2 o

0

a)

0 30 GO 90 " 1 2 0 150 -10

Age (y)0 10 20

Longitude (°)

Fig. 3. Ashfree amounts ofSOMfl related to (a) stand ageand (b) longitude.

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280 A. I. Gärdenäs Scand. J. For. Res. 13 (1998)

50 100 150 200 250

Age (y)30 40 SO"

Basal area (m2 ha-')60

literature dataexponential model

-5 " 0 " 5 " 10 15 20Temperature (°C)

Fig. 4. Ashfree amounts of SOMfl

in spruce stands related to (a)stand age, (b) basal area and (c)mean annual temperature. Standsat Grunewald, Oulanka andSävar in parentheses.

found to be the most important single determiningfactor for SOMn.

The fact that tree genus is so important for amountof SOM,, may be due to litter quality, litter quantity,both or other factors. The effect of litter quality onthe extent of decomposition has been found to bemainly related to variation in N, lignin and Mnconcentrations of fresh litter (Johnson 1992, Berg etal. 1996). Tree species differed significantly in N andlignin content, while no difference in Mn content wasfound (Reurslag & Berg 1993). A large range in genusaverage SOMH (4.-40 x 103 kg ha~") was found for asmall range in genus average initial N content (5-10mg g~')> and spruce had an outlying value (Fig. 5fl).The average lignin content in fresh litter could becalculated for only a few genera (Fig. 56). Calculat-ing the N-lignin ratio for pine, spruce, beech andalder gives a ratio close to 0.02 for pine, beech andspruce, while for alder with its low amounts of SOM,,a ratio as high as 0.11.

The amount of litterfall limits the potential ofSOM,, accumulation. Staaf & Berg (1977) noted that

the heterogeneity of SOM,, of a Scots pine stand wasmainly determined by coarse litter and the distancefrom trees. The average amounts of yearly litterfalland SOM,, for spruce and pine were clearly distinct(Fig. 5c). For the other genera, the average amountsof litterfall were based on few studies; nonetheless,the combined data seem to indicate an increase inSOM,, with increase in litterfall (Fig. 5c).

Deciduous species had lower SOM,, amounts thanconiferous species, but this does not imply that theSOM amounts in the whole soil profile are lower fordeciduous species. The SOM,, amounts in the beechstands in Soiling were almost seven times higher thanthose in Abbotswood and Kongalund, while soil typewas the same for Soiling and Kongalund. The highdeposition of N at Soiling might detain decomposi-tion, as Kowalenko et al. (1978) and Nohrstedt et al.(1989) found that N fertilization hampered respira-tion. In the pine stand at Grunewald, close to Soiling,and at other spruce stands with known high deposi-tion (Skogaby and Hasslöv) the amounts of SOMH

were also higher than could be expected from theirgenus and stand age.

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Scand. J. For. Res. 13 (1998) SOM in European forest floors 281

5 10 15 20 25 30 0

Initial N content (mg g1)

80 1G0 240 320 400Initial Lignin content (mg g"1)

JOCO

"o-S2 10

• c )

piney

i

Douglas fir

W. nemlock

, i

00 3000• i .

4000

spruce .

beechC)

Grand fir

i . i .

5000 G000 70

Fig. 5. Average SOMfl of treegenera as a function of (a) initialN content of fresh litter (Reurslag& Berg 1993), (b) initial lignincontent of fresh litter (Reurslag &Berg 1993), and (c) amounts ofyearly litterfall litter (Berg et al.19936, Reurslag & Berg 1993).Only species occurring in theSOM,, studies are included in theaverages, and averages based onless than three values are given inparentheses.

Amounts of litterfall (kg ha1 y')

Ovington demonstrated the importance of standage for SOM,, in a chronosequence of nine Scots pinestands (Ovington 1959). Ryan et al. (1997) found thattree production diminishes after maximum canopydevelopment is reached. This should result in a lowerlitterfall and eventually a lower SOMfl, which mightexplain the strong deviated position of the stand atOulanka (250 years old, Fig. 4a). However, far toofew stands older than 150 years are available toconfirm this theory.

For some environmental factors, like mean temper-ature and annual precipitation, linear response mod-els were not suitable. The exponential model formean annual temperature seemed more suitable. Theexponential model is often used for the temperatureresponse in decomposition models like Bunell et al.(1977) and Simunek & Suarez (1993). For furtheranalysis of the importance of temperature and precip-itation, I would prefer a process oriented carbonbalance model with a higher time resolution becausetemperature and precipitation could influence theamounts of SOMn in three ways: decomposition rate,

biomass growth and thereby litter production, andtransport of SOM from the forest floor to the mineralsoil by soil fauna or leaching. Comparing differentSOM models for forest ecosystems, McGill (1996)noticed that SOM models of forest ecosystems oftenoriginated from models for agricultural and grasslandecosystems. In agricultural systems, the permanentlitter layer is often missing, and this might be onereason why this layer is described with little detail inforest models. Chertov (1990) made a model speciallyfor the forest floor as a function of temperature,moisture, litter production and ash- and N-content oflitter for a Scots pine stand in the middle taigaregion.

Although sampling methods differed greatly andinformation about several important factors like his-tory of land use and minor abundance of otherspecies was lacking or limited, support for the impor-tance of tree genus was found. Too few data wereavailable to conclude whether litter quality and/orquantity differentiated the genera in amounts ofSOMfl, but it seemed to be worthwhile to continueresearch in this direction.

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282 A. I. Gärdenäs Scand. J. For. Res. 13 (1998)

ACKNOWLEDGEMENTS

Dr Gunnar Ekbohm and Dr Ewa Kasmierszak madesuggestions for the statistical analyses. Dr Lars Hy-lander and Hans Johansson assisted with the SASapplications. Dr Dan Berggren and Dr Henrik Ecker-sten contributed with valuable comments on themanuscript and Dr Mary McAfee corrected the lan-guage. Vattenfall Utveckling AB financed an earlierphase of this project, in which Dr Björn Berg wasalso involved. All are kindly thanked.

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