in Southe ,istern Eco . , Lems,F. G. Howell,J. t3. Gentry and M. H., Smith (eds.). ERDA Symposium Series.(UHF-740513) . pp. 609-624. 1975

Pflt!.70..JCTION, DECOMPOSITION, AND

C",(CLING IN A MIXED HARDWOOD

A WHITE PINE WATERSHED

NE Rmtrci:OMACIt,. J[, • and CARL. D. MONKDepartmeli«rf botany, Unr, • ersity of Georgia, Athens, Georgia

ABSTRACT

Lim!) production and decomposition data were obtained for a mixed-hardwood watershedaorl for a v.hite pine watershcil.Litterfall data were obtained for Laves, stems, flowers,acorns„ind ' miscellaneous debris in the hardwood watershed and for needltk, stems, andcones in the white pine watershed. Litterfall data obtained included biomass of litter;niirogen, phosphoms, potassium, calcium, and magnesium contents in hitter; and structuralorganic cou.iituents rlignin, cellulose, and total fiber) of leaf litter. Total annual litterproduction in the hardwood watershed (1970-1971) was 4369 kg ha' year -I of which2773 kg ha' year -1 (64") was leaf litter. Total annual white pine litter production(1970-1971) was 3253 kg ha year', 98% of which was needle litter in the young standrpianted I956). Litter decomposition data were obtained for weight loss rate and for lossrates of tula-ients. lirst-year litter breakdown rate of confined mixed-hardwood leaflitter was k -0.70 year -1 ; the breakdown rate of confined white pine needle litter was

= -0.46 year . Litter decomposition rates of chestnut oak, white oak, white pine, redmaple, and dogwood \* irc significantly correlated with senescent leaf car6on-to-nitrogenratio and sjerophyll nnlex, the sclerophyll index giving a better statistical estimate ofdecomposition rate.

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Nutrient cycling ant-energy flOw are two ecological processes that delineate thestructure and dynamics of ecolOgical syst' r rns. In terrestrial ecosystems -inimportant set of energy flows and nutrient transfers result from litterfall andfrom the subsequent decomposition of litter. Litterfall and litter decompositioncan be thought of as separate events in terrestrial ecosystems, but such ecological

'Present address: Department of Forest Management, School of . Forestry, Oregon StateUniversity, Corvallis, Ore.

609

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610 CROMACK AND MONK

processes as energy flow and nutrient cycling permit them to be linked togetherfunc;ionally in an ecosystems context.

Seventy pei . cent of the energy fixed in net primary production in a grazedmeadow ecosystem goes directly to decomposer organisms (Macfadycn, 1963).As much as 90% of terrestrial net primary production ultimately is utilized bydecomposers (Whittaker, 1970; Odum, 1971). It has been suggested thatdecomposer organisms may be just as important in aquatic ecosystems as theyarc in terrestrial ones (Pomeroy, 1970). Litterfall and litter decomposition alsoaccount for a substantial portion of internal nutrient cycling in terrestrialecosystems. In a study of a mcsic oak-dominated forest floor, littcrfallaccounted for approximately 60% (average for the elements phosphorus,potassium, calcium, and magnesium) of internal nutrient input to the forestfloor (Carlisle, Brown, and White, 1966a; 1966b).

Biodegradation of litter is a complex process in which microorganisms andsoil animals interact synergistically (Crossley, 1970; Witkamp, 1971). Both themicroorganisms and soil animals arc strongly influenced by such abiotic factorsas soil—litter 'temperature and moisture. Microorganisms predominate in litteroecompositi,on, primarily because they have enzymes capable of biodegradingstructural carbohydrates including such complex end products 2f carbohydratemetabolism as lignin (Witkamp, 1971). As much as 90% of litter materials canbe biodegraded by microorganisms (Macfadycn, 1963). MicroOrganisms are alsoimportant in the release and transfer of nutrients bound in the litter (Witkamp,1971; Stark, 1972; Todd, Cromack, and Stormcr, 1973). Soil animals, byinteracting synergistically With the microorganisms, function as one set ofbiological control factors which regulate rates of litter decomposition (EdWards,Reichle, and Crossley, 1970; Ausmus and Witkamp, 1974). In turn these animalsarc dependent, for their nutrition primarily upon the microorganisms they ingestsince soil animals lack enzymes necessary to biodegrade. all but the simplest'forms of structural carbohydrates (Nielson, 1962).

Considerable data exist for litter production and litter nutrients in forestecosystems, with major review papers summarizing forest-litter data fro a manydifferent sources (Ovington, 1962, 1965; Olson, 1963; Bray and Gorham, 1964;Rodin and Bazilc- vich, 1967). For the purpose of facilitating comparison of datafrom many different ecosystems, tota : , annual litter budgets are preYerred;subannual budgets for major litter components usually arc available in mostrecent studies (Carlisle, Brown, and White, 1966b; Sykes and Dunce, 1970; Gosz,Likens, and Bormann, 1972).

In this paper we compare nutrient cycling from litter production and litterdecomposition in a mixed-hardwood ecosystem with the same nutrient-cyclingproccsxs in an adjacent white pine Wilms strobus L.) watershed in the southernAppalachians. We also compare indexes of foliage-litter substrate qualityobtained from the two different vegetation types and discuss nutrient cyclingimplications of foliage-litter nitrogen and phosphorus levels in the context ofcam bon all ocation to such subStrates as lignin and cellulose.

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LITTER PKODUCTION, DECOMPOSITION, NUTRIENT CYCLING 611

The two forest watersheds used in thiS study arc located within the U. S.Forest Service Coweeta I lydrologic Laboratory, Franklin, N. C. The two fieldsites were chosen such that litter production and litter decomposition in amixed-hardwood ecosystem could be compared and contrasted with correspond-Mg litter d y namics in a white pine ecosystem. The mature mixed-hardwoodstand, designated by the U. S. Forest Service as watershed 18, is 12.5 ha in area.The white pine stand, planted in 1956, is 13.5 ha in area and was designated aswatershed 17. Broad environmental differences between the two watersheds areminimized in that they, have similar slopes, aspects, altitudinal ranges, soils, andprecipitation and are contiguous for approximately one-half their verticaldistances (Kovner, 1955; Johnson and Swank, 1973).

NATERIALS AND METHODS

Sc,uare traps 0.5 m on a side were used for hardwood leaf -ter, flower litter,and debrL To make all traps equal regardless of ground slope, we placed eachtrap on wooden legs and leveled each with a spirit level during installation. Thetraps were checked periodically thereafter. Thirty of the litter traps were placedat random in the hardwood watershed in the fall of 1969. Acorn litter wascollected from 20 60-cm b y 60-cm cloth-bottomed deadfall traps.

White pine needles were collected in 200-cm by 15-cm aluminum troughsUsed in a throughlall study of watershed 17 (Best, 1971). Nineteen of thesetroughs were used; 10 were located in one random transect and 9 were located inanother random transect through the watershed.

During the fall periods, leaf and needle litter were collected at approximately2-week intervals. Since some of the white pine needles became lodged inbrandies, additional litter collections were made at intervals throughou' toey ear. No hardwood collections were made in winter because little leaf litterremained on the trees after December. Beginning in late spring and continuingthrough the summer at monthly intervals, flol,..'er litter and debris were collectedin the hardwood watershed.

Stem and branch litter were collected from 18 10-m by 10-in ground plots inthe hardwood stand. During the summer of 1970, the total accumulatedstanding crop of woody litter was estimated. Within each plot, two 2-m by 2-msubplots, located on opposite diagonal corners of the large plots, were used tocollect stern litter less than 2.5 cm in diameter. In the whole plot, woody litter2.5 ens or greater in diameter was collected. American chestnut logs andbranches were excluded from the standing crop totals since they representedlitter input from a catastrophic event in the watershed. The followin i, yearcol feet ion was made (Jr :111 woody litter that had fallen into the plots.

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612CHOMACK AND MONK

In the summer of 1970, 103.3 3-rn by 3.33-rn woody-litter lots ere setin e . white pine st::rl; collections

were made in the summer op

f 19 71. Most ofthe (Lad branches remain on the trees. Consequently woody-litter input to theforest floor is as yet relatively unimp

the white pine stand.ortant forWoody-litterfall data from the ground plots of both watersheds

were correctedto a horizontal area basis for average slopes of 53% intersbothed watershe

correds. Thispermitted co

mparison of woody litter from the ground plots with li tter-trap dataon a common basis.

Data for first-year leaf-litter breakdown of single species were obtainedduring two co nsecutive 1-year periods, b eginning in the fall of 1969. Single treespecies for which leaf-litter decomposition data were obtained included chestnutoak (Quercus • prinus L.), white oak (Querruc alba L.), . red maple ,(Acer

rub •,lin C.), and flowering dogwood (Cornus floritia L.) in the ,hardwoodwatershed and eastern white pine in the pine stand

. The four species used fordeciduous-litter breakdown were chosen to give a range of deco mposition ratesand to represent the more important species groups

pro ducing leaf litter withinthe system . in the hardwood stand additional leaf-decomposition data wereobtained for mi xe d-species leaves in litter bags.

Data for litter breakdown of single species were obtained using decimetersquare fiber glass litter bags (Crossley and I loglund, 1962), eac cotainingappro ximatelypp ximately 2.5 g of leaves. In the de ciduous stand the integratedh deconmposi-tion rate was

obtained for mixed-hardwood leaves confined in the large,0.125 m 2 , litter bags each containing appro ximately 40 g of decu us,leaves. Inaddition io the larger size, a different bag design

wasused foridtheo mixed-litterbags to permit easier entry of soil a nimals. The tipper half of each large bag was

made of 2.54-cm-mesh nylon netting, while the lower half was fine 2-mm-meshnylon to inhibit loss of small leaf fragments from the bag. The decimeter-sizedfiber glass lit; r bags used for single-species leaves from cht , ,red maple, and d ogwood and needles of white pine were al/

es1-tnu Oakmm meshwhite oak

.Littcf bags were removed from the field at approxiMately mon• lny intervals.The bags were weighed to determine percentage weight loss from originals

amples before the samples were ground in a Wiley mill for nutrient analysis.Litter weight loss was quantified for single-species and for mixed-deciduouslitter bags using the exponential decay model of Olson ( 1 963). Nutrient lossrates from mixed de

ciduous litter were also quantified using the same approach.Leaves, other small litter, and woody litter were air-dried for storage where

practical or dried at 50°C when wet litter or green leaves were collected. Leavesthat were to be used for decomposition studies were air-dried only to prevent

possible Chemical changes that might affect decomposition. For nutrianalyses, including nitrogen, litter was dried at 70°C. All final dry-weigent

htbiomass calculations were made on litter subsamples dried at 70°C. Sampleswere oven-dried at 50°C for analysis of organic constituents of litter. Nutrientanalyses included determinations of sodium and the following major elements:nitrogen, phosphorus, calcium, potassium , and magnesium.

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LITTER PRODUCTION,

DECOMPSITION, NUTHIENT CYCLING,‘ rni O

crojeldahl technique was used

for nitrogen deternlinations. e n

kj../.Thi procedures were those

of the Soil Testing and Plant Analysis

. 673IMoratory of the Agronomy

Department, University of Georgia, a s ivgen in

their 1971 laboratory manual

(1,aboatoi:y Procedures for

.1HalysiS Of Soils,

l'f.e,/, 11://cr, mul I'laut Tissue). All

other elements the

were determined by

direct-reading spark emissions

spectroscopy (]ones

and Warner, 1969). Ana/ysesfor the 'percentage of acid-detergent

fiber andand

lignin) and non-cell-wall

plant material followed the methods of Van

Soest (1963, 1966, and] 96 7). The

sclerophvIl index 71%is calculate

following the method of Loveless (196). Since

the scicrophyll index as originally proposed uses the

ratio of crude fiber to crude

protein x 100, crude fiber

was estimated (Ellis, Atatrone

and Maynard, 1946) as

ioi:ii acid-detergent fiber X 0.64. Crude protein teas calculated (Loveless, 196 I)

s percent nitrogen X 6.25. The percentage

of carbon in leaf litter ud for

leaves using a regression given

by, Van Soes

decomposition studies was estimates! from the percenta sege lignin content in

To amtlyze for t (1 963).•changes in nitrogen and total fi ber ,

we harvested green le;Ives

in August &bin se veral of the more important tree species.Jn

the hardwood stand

and from the white pine s t and.

species.

To analyze nutrient differences between

senescent leaves and needles, we collected

additional leaf samples in mid-

coniferous :Ind deciduo

October. I;or Comparative purposes we have ill ClUded a few values r.r westernRESULTS

AND DISCUSSION

Results arc given as annual bu dgets of nitrogen,

phosphorus, calcium potas-

slum, a:Id magnesium in litterfall

and biomass of different

hardwood and white

pine litterfall. components in 'Fables 1 and 2. The

leaf-litterfall c;ttegory in the

hitrdwood

watershed does not include litter from shrubs, such as li,iloO,147zifolia

L. il'bodo,/drou. uiaximum L., or

herbaceous litter. Such litter Was estimated

(flay, 19 71) to total 370 kg ha-1 year-1. Herbaceous

itterfall +as not

the closed canopy.estimated in . the

white pine stand; ground cover ryas sparseleaf l, however, o wing toTotal litter production in the hardwood

watershed was 4 369 kg ha-1 year-1,

which is within the lower 95%

confidencethe annual litter

production, totaling 4800 ± 548 kg ha-1 year-1 interval of

for II other North American

warm temperate deciduous forests

(Bray and Gorham 1964). hardwood_ 't

production tV:i 5

only 14';,; greater than the mean annual

litter production of

3730 ±. 260 kg ha-1 year-1 in 67 cool temperate

ann ual forests (Bray and

Gorham, 1964). It, an oak-dominated British forest with a

similar rainfall regime

as Coweeta, cm soils of low nutrient sta tus., a total annual litter b udge t of 3858

kg ha-1 year-1 was observed (Carlisle Brown, and White,

19 66a) . On calcareous1970).

kg ILt year-1 was reported (Sykes

and Bunce,

.soils in Aka thop Wood, Grea t Bri tain, an average

total litter p action of 5130rod

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G1 4 CF-10MACK AND MONK

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LITTER PliODUCTION, DECOMPOSITION, NUTRIENT CYCLING 615

TAB LE 2

WI HIT PINE WATERSHED 17: ANNUAL LIVIIR FALLBUDGET FOR BIOMASS AND MACRONUTRIENTS

FOR 1970-1971

Gaclury Needles Stems. Pine cones To t

Biomass,kg ha 1 year ' 3184 ± 293 38.5 i 14.0 30.6 ± 8.1 325 3

of biomassi\lacronutrient.,

kg ha l year'

97.9 1.2 0.9 100.0

Nitrogen 26.27 0.16 0.11 26.54Phiisphorus 3.66 0.03 0.03 3.72Potassium 5.32 0.10 0.12 5.54Calcium 19.08 0.11 0.003 19.19Magnesium 2.71 0.002 0.000 2.71

Most dead stems remained attached co the trees.

Total annual litter production in the white pine watershed was 3253 kg ha-1year -1 , considerably less than the mean annual litter production of 5425 ± 950kg, ha -1 vear 'l in 8 other North American warm temperate coniferous forests(Bray and Gorham, 1961). llowever, the white pine needle-litter production of3184 ± 293 kg ha 1 year does not differ significantly from annual needle-litterproduction, averaging 3640 ± 510 kg ha t year I in 10 North American warmtemperate coniferous forests (Bray and Gorham, 1964). North American warmtemperate forests average 37';•■ nonneedlc litter, while cool temperate forestsaverage 23% (Bra y aod Gorham, 1964). On this basis, the white pine watershedat Coweeta could average 4500 to 5000 kg ha -1 year I total annual litterproduction when mature. Needle-litter production did not differ significantlyfrom 'white pine needle production, averaging 3386 ± 375 kg ha - ' year a. 'torthree stands located iu the northeastern United States (Chandler, 1944).

Comparing the nutrient budgets of these three northeastern white pit*:aands with data in Table 2 shows that the former were cycling 26% morenitrogen, 10% more potassium, 54% more magnesium, nearly the same quantitiesof calcium, and 16% less phosphorus. Comparison of total annual litterfallnutrient budgets in Tables 1. and 2 shows the hardwood stand was cycling morepotassium, calcium, and magnesium in hail-fall than the white pine stand. Theappreciably greater amounts of potassium, calcium, and magnesium being cycledby the hardwood ecosystem than by the white pine watershed arc consistent withother reports (Chandler, 1944; Ovingron, 1962; 1965); temperate hardwoodecosystems generally .cycle more cations than coniferous ecosystems with similarproducti y ities. In the present case, total littcrfall and nutrient return in lit ter areull.tantially less in the white pine watershed.

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616

CROMACK AND MONK

Litter front tlae reproductive materials and fine 'debris in the hardwoodwa terhed had equ.d or hi 'her

g percentage concentrations of all macronutricnts,except calcium , as did deciduous leaf litter. Branch litter contained smallertilltricnt concentrations than de ciduous leaf litter. lowcr l hich wasmostly from oaks, and miscellaneous debris contained relatively mo

itter, wre macro-

nutrients, .except potassium, than did acorns. Partitioning

bss s into

component parts permitted the nutrient unit of the to acorn

be aessed.

Acorn pro duction in 1970 to 1971q

was heavy and totaled 9.4% of total lit:terfallbiomass. Threeyears of acorn production in an oak-dominated forest in GreatBritain averaged 2.5% of total litter (Sykes and Bunce, 1970.

a e mcorns were collecte

noticeably less.) Al though nod at

(Sykesduring the subsequent year, production

:.ced

"Total litter nutrient input b udgets in thro ughfall and litterfall 'arcgiven inTable 3. Data are for ipotassum, calcium, magnesium , and sodium in both thehardwood ;the

nutrind t white pine stands.a mounts o Both wa tersheds are cycling nearly similarf ents thro ughfall , but they differ substantially in quantities

TABLE 3

CO,V.PARISONLITTEIZFALL ANDTIIRO UGIII"ALL

NUTRIENT BU01' BETWEEN HARDWOOD ANDW1111• PINE STANDS FOR 1970-1971Nutrient budgets,

kg ha' year-IMg Na•

Ilardwood

Annual litterfall,total

18.07 44.49 6.55 0.009Annual throughfall,torah

Total annual30.50 8.10 3.10 8.61.0budget

4-8.57 52.59 9.65 8.609Annual lit terfaq,of total

White

3/.2 84.6 67.9. 0.1pineAnnual litterfall,

total

5.54 19.19 2.71 0.002Annual throughfall,tontlt

Total annual30.50 6.30 2.00 7.20

budgetAnnual litterfall.,

36•54 25.49 4.71 7.202of total • 15.2 75.3 57.5 0.03

'Sodium co nceiii ration is 2rpm in hardwood litter and0.5 rpm w hitt . pine lit ter .1 From Itcu ( 97 1 ).

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LIT-10{ PRODUCTION, DECOMPOSITION

NUTRIENT CYCLINGproportions of nutrients

617.a bsolute a„md in total

contributed by li tterfa II. In terms ofm o unts and percentage contribution by ht terfall to- the total nutrient1u-ought-all than is the

bt.dgets, the deciduous system has more

elements in litterfall re lative tocase for the white pine stand.

When the total nutrient b udgets in the hardwood watershed

forest in Britainarc compared

with those of a mesic oak-d(Carlisle, Brown, and

• White, 1966b) for

it isthe elements potassium, calcium, and magnesium, i

seen

that the percentages contributed by litterfall in the Coweeta hardwood stand arc

all ;reater by an a verage of 22%. Total a mounts of potassium and calcium cycled

at Cowecta in both throughfall and litterfall also are greater

by 20 while 27%more total magnesium was

cycled in their system. Sodium

bu dgets arc very,

different in the two deciduous forests with.

nearly seven times more total

sodiurn being cycl,:d in the B ritish oak woodlandugh detailed budgets'are

not given in l iable 1, actual sodium Rittercontent of litt in the British system

Warlisle Brown , and White, 196610 was three orders of magnitude

greater thanthat of the haidwood litter at

Coweeta.In Table 4 the data for organic constituents in laves sllow several trends.

Most senescent leaves have higher total fiber and lignin but lower nitrogen and

M ost than do 'leaves in the mid-growing season.

For purposes ofcomparison with deciduous .s

pecies, we had to take composite sa mples of all age

classes of needles or leaves to obtain a representative

sample for Collarpothesized that foliar sclero

nutrientand Organic analyses.

It has been hyphyll indexes a bove 100 to 150

in dicate ,a vegetation that is adapted to gr

owing on phosphorus-deficient sites(Loveless, 1961; 1962). 'Species growing on such sites generally have lowerprotein content than vegetation with adequate phosphorus available (Loveless,1961; 1962).

Although there is some question as to when in the season to.

sample green l e..vc.; for sclerophyl/ determinations,

very early season values

probably would be misleading, o wing to higher nitrogen and phosphorns.-ontentand lower fiber content. When mid-August values for green hardwood leaves atCo weeta are com pared, all sclerophyll in dexes arc high, ind icating a scler0,phyl-Ions vegetation. All conifer species compared including

those from thew i:stern

UnitedStates, have high sc/erophyll indexes. Red alder and snolvbrush arcin cluded as exa

mples of nitrogen:fixing species. Nitrogen-fixing, species mighthave low .selerophyll indexes simply

because they have a readily available sourceof nitroen even in soils of low phosphorus availability. American chestnutsu rvives at Coweeta only •a

Evidence has been presented that astump sprouts. phosphorus content below 0.3% is1

associated with a lower protein content and a high scleroPhyll index (Loveless,

9 1961;

62). Most phosphorus values for green .leaves in the species listed inTable 4 were less than 0.2%, except yellow poplar and

Douglas fir, and all wereetation Occurs in

ist both wet and dry habitats (Loveless, 1962).

Phosphate deficiency is characterie

of acid soils in area of high rainfall

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SpeciesSpecies Location__________ % N % P

% total aciddetergent

fiber%

lignin

Scicro-phyllindex---

Chestnut oak (C)' Coweeta,______

Chestnut oak (S)2.0 0.18 32.8 12.6 168N. C. 1.2 0.12 48.3 25.5 430

Scarlet oak (C)Scarlet oak (S)

Coweeta, 1.8 0.16 31.0 15.5 177N. C. 0.9 0.12 34.8 16.7 397White oak (C)Wlnte oak (5)

Coweet a, 2.0 0.17 26.0 9.1 132N. C. 1.0 0.12 38.2 17.2 390fixed hickory (G)

Mixed hickory (S)Coweeta, 2.1 0.18 33.7 9.2 165N. C. 1.2 0.16 42.9 16.9 351

Red maple (C)Rcd maple (S)

Cowecta, 1.5 0.16 29.3 9.4 200N. C. 1.2 0.14 a0.0 12.7 260Yellow poplar (C)Yellow (5)poplar

Coweeta, 2.0 0.20 29.3 7.2 146N. C. 1.0 0.12 41.1 14.6 443Dogwood (C)Dogwood (S)

Coweeta, 2.0 0.18. 22.4 3.5 115N. C. 1.4 0.14 • 21.1 3.9 160American chestnut (C)American chestnut

Coweeta 1.9 0.18 30,3 7.6 163(S) N. C. 1.0 0.10 32.8 8.8 354White pine (G)White (5)pine

Cowceta, 1.4 0.17 38.8 16.4 296- N. C. 0.9 0.11 54.3 31.0 617Douglas fir (G) .Doilas fir (S) -

Corvallis, 1.0 0.27 28.3 16.5. 309Ore. 0.5 0.26 38.6 24.1 815Ponderosa pin.2. (G) Flagstaff,

Ariz.0.8 0.14 32.4 16.1 414

Engelman"' spew-c (G) Logan, Utah 0.6 0.07 27.4 13.3 510Red alder (G)Red alder (S)

Blue River, 2.5. 0.23 11.9 6.0 -' 49Ore. 2.1 0.16 19.4 9.5 94Snov.arosh (C)Snowbrush (S) '

131ue River, 2.0 0.13 15.7 6.1 80Ore. 0.81 0.06 21.0 10.0 265

IF1,0,17711.71,1, ,T,..,1411/7.-77.1M•111OrlInr

618CROMACK AND MONK

TABLE 4

REPR ESENTATIVE SPECIES DATA FOR FOLIAGEORGANIC-MATTER QUALITY

••

N

All samples represent mean values of composite samples taken from at least fivedifferent individual trees, except Ponderosa pine and Engclmann spruce. G = mid-growingseason, S = senescent.

(Salisbury, 1 95 9); Coweeta watersheds arc situated in a region of high rainfall,and considerable weathering of the older Appalachian land surface has occurred(Kovner, 1955). Return of phosphorus in litterfall averages 0.11% concentra tionon a dry-weight basis, based on total litterfall biomass and total amount of

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LITTER l 'iODUCTION, DECOMPOSITION, NUTRIENT CYCLING 619

pnoTlionis given i.i Table 1. The white pine stand is cycling an equally low levelof phosphorus in litterfall (Table 2). The critical level is 0.2% for phosphorusimmobilizat.ion in decomposing litter (Alexander, 1961). Phosphorus values indecomposing litter at Coweeta arc all less than 0.2%, showing that this elementwould tend to be immobilized by microbial organisms. First-year phosphorusloss of 34% from mixed-hardwood leaf litter was considerably lower than thefirst-year weight loss of 50% (Cromack, 1973). Further evidence of phosphorusdeficiency in Cowecta soils comes from experimental data with these soils inwhich American sycamore (Platanus occidentalis L.) seedlings lacking m y corrhi-zae show phosphate-deficiency symptoms when grown in greenhouse containers(Best, Marx, and Monk, 1974); presence of mycarrhizal symbionts would permitsurvival and ., reproduction of tree, vegetation in the infertile soils characteristic ofCoweeta. Phosphorus-cycling levels remain low enough to indicate that thevegetation would have the high sclerophyll indexes observed.

High sclerophyll indexes are associated with lower foliage ash contents(Loveless, 1962). One general implication of this work is that ecosystems whicharc- characterized by highly sclerophyllous vegetation (possibly due to phos-phorus deficiency) are cycling smaller quantities of total nutrients, as reflectedby ash content, than is the case for vegetation with low sclerophyll indexes(Monk, 1 966). When comparative sclerophyll data become available from manyof the temperate ecosy stems for which litter nutrient data exist; then a betterjudgment can be made concerning the relationship between nutrient cycling inthe system and the sclerophyll indexes of their characteristic vegetations.Assessment of the role. of mycorrhizal symbionts in permitting survival andreproduction of tree vegetation in soils of low fertility is also needed.

Decomposition and nutrient-loss-rate data were obtained for both hardwoodand white pine litter.- On the basis of sets of litter bags put out in twoconsecutive years, ..Cirst-year, decomposition data for single species in bothwatersheds showed the following exponential weight loss rates: k = —0.46 year-1for white pine, k = —0.61 year -1 for chestnut oak, k = —0.72 year -1 for •hiteoak, k = —0.77 year -1 for red maple, and It = yearr 1 for dogwood. Amixture of ha rchs Ood leaveS contained in large litter bags - had an annualexponential loss rate of k = —0.76 year -1 , or in terms of percentage weight l6ss,50% weight loss occurred during the first year. By contrast, white pine averagedonly 37% Weight loss during the first year. The slowest decomposing hardwoodspecies was chestnut oak, which averaged 46% weight loss during the first year.Dogwood, the fastest decomposing hardwood species, averaged 68% weight lossduring first-year decomposition.

Release of nutrients during first-year decomposition showed that nitrogen,phosphorus, and calcium tended to be immobilized, while magnesium andpotassium were lost at greater rates than weight. The mixed-hardwood litter lostonly 12% nitrogen during the first year, and there was no significant loss ofnitrogen from white pine litter. Mixed-hardwood litter had-a carbon-to-nitrogenratio of 62. : 1, while white pine had a carbon-to-nitrogen ratio of 58 : 1;

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O20 CROMACK AND MONK

carbott-to-nitrogen ratios appreciably greater than 30: 1 result in increasingimmobilization -of nitrogen in the microbial populations' decomposing litter(AlexanL;r, 1961). •

Mixed hardwood litter lost 34% of phosphorus during the first year ofdecomposition, in contrast to a 50% weight loss; an indication that partialinunobilization of phosphorus occurred. In white pine there was no significantloss of total phosphorus in first-year decomposition; but the weight loss was37%. As indicated previously, the probable reason for the appreciably lower lossrate of phosphorus compared to weight loss from decomposing litter was the lowlevel of phosphorus (0.12%) in foliage litterfall of both watersheds. The criticallevel at which phosphorus is immobilized in organic material is 0.2%.

Calcium loss rate was also less than the weight loss of the deciduous litter,e.g., a 29% first- y ear loss of calcium compared to a 50% weight loss. There wasno significant loss of calcium from white pine litter during the same period.Evidence exists that litter- and soil-inhabiting fungi can concentrate substantialamounts of calcium (Stark, 1972; Todd, Cromack, and Stormer, 1973),c,luantities perhaps sufficient to result in partial immobilization of this nutrientin litter. M cronutrient quantities of calcium as well- as other cations, such as•potassium and sodium, can be used by fungi to neutralize oxalic•acid, which is alow-energy waste product of carbohydrate metabolism in fungi (Foster, 1949).Oxaloacetic acid, a key intermediate in respiratory metabolism, is hydrolyzed tooxalic acid and acetate by oxaloacetate hydrolase in fungi (Burnett, 1968). Thecommon metabolic production of this acid would result in substantial demandof cations, such as calcium, in the production of oxalate salts (Foster, 1949).

Potassium and magnesium loss rates were greater than weight loss rates forboth mixed-hardwood litter and white pine litter. Potassium loss during the firstyear for hardwood litter was 83%, while potassium loss was 82% from white pinelitter. Magnesium loss was 77% from mixed hardwood litter and 68% from whitepine. It has been shown that I 37 Cs (an analogue of potassium) loss rates weregreater than weight loss rates for three deciduous species (Witkamp and Frank,1969). In another study the loss rate of I 34 Cs was also considerably greater thanthe weight loss rate of white oak (Witkamp and Crossley, 1966). Potassium isknown to be significantly concentrated •)y both the vegetative and spoi)ocarpportions of fungi-inhabiting litter (Stark, 1972; Todd, Cromack, and Stunner,1973). In contrast to calcium, which tends to form insoluble calcium oxalatecrystals: in such vegetative structures of fungi as hyphae and rhizomorphs,potassium is readily translocated in fungi where it is accumulated in sporocarps(Stark, 1972). Although potassium oxalate can be formed, it is much moresoluble than calcium oxalate and would tend to be lost by lea ching whenexcreted by hyphae. Potassium is more easily leached from litter than any other -essential cation (Cromack, 1973). Magnesium was not significantly accumulatedb y fungi in litter substrates at Coweeta (Todd, Cromack, and Stunner, 1973),indicating a lesser use rate for the element than for potassium. Magnesium also'is

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LITTER PRODUCTION, DECOMPOSITION, NUTRIENT CYCLING 621n , )t

as easily leitelied from lit ter as potassium (Cromack, 1973), in part because itis less easily displaced from cation exchange sites in liner

s ubstrates than ispotassium.

Weight-loss regressions were calculated from exponential loss rates of white

pine, chestnut oak, white oak, red maple, and d ogwood (as previouslymentioned) and the chemical p roperties of the senescent leaves of those trees.Two simple linear regressions were calculated using te exponentil weight lossrates of the species as the dependent variables and theh carbon-to-nitroagen ratiosand scleroplivIl indexes of senescent leaves of these species (Table 4)

asindependent variables. The linear regressic n of exponential weight loss rate (y)and earbon-to-nitrogen ratio (x) is

y — 1.92 + 0.026x (n = 5, r = 0.86, p = 0.06) (1)The data for the carbon-to-nitrogen ratio in senescent leaves of the five speciesused tit •F,q. 1 were: white pine, 58.0: 1; chestnut oak, 42.2 :

i18.3: 1; red. 39.2: 1; and dogwood, 31.8 : 11; wh te oak,

. The li of theexponent ia! weight loss rate (y) and scl erophyll index (x)near. regression is

y + 0.001x (n = 5, r = 0.90, p < 0.05) (2)Co mparison of the two regressions shows that the s elerophyll index of senescentfoliage is a sta

tistically better variable with which to assess leaf deconapOsitionrate than is the se nescent leaf carbon -to-nitrogen ratio.The s clerophyll index incorporates information about organic-matter qualityfor deco mposition in terms of such structural compounds as cellulose and lignin,

as well as crudc protein nitrogen. The carbon-to-nitrogen ratio itself does notindicate the nat me of the carbon substrate. Alexander (1961) presents evidencefrom the work of others (Fuller and Norman, 1943; Peevy and Norr

,at„ 1948;Pine!: ,•\11ison , and Sherman, 1950) that litter decomposition rates may bebetter p redicted . . by knowing lignin confent than by knowing the carbpn-to-nitrogen ratio. The present -data on s eler.?phyll index, which inco rporates ligninand cellulose as total acid detergent fiber, plus arlinear regression relating specieslignin content to deco mposition rate (C r omack, 1973), • support his basichypothesis (Alexander, 1961).

possible co nsequence of selerophylly not considered by Loveless (1961;1962) was that leaf-litter decomposition rates would be less in those species withhigh, sclerophyll in dexes. As discussed p reviously, he established relationships

between high leaf sci c r ophyll indexes and low leaf levels of phosphorus,n itrogen, and ash. In forest ecosystems characterized by cycling low levels ofnitrogen and ph osphorus, the selerophyll index can be considered an index toslower organic matter decomposition rates and release rates of such nutrients as

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622CROMACK AND MONK

nClUICaEDGP.';ENTS

This work was supported in part by the Eastern De ciduous Forest Biome,Interritional Biological Program, funded by the National Science Foundation

under interagency' agreement AG-199, 40-193-69 with the U. S. gyCommission—Oak Ridge National

Laboratory; in part by National ScienAtomieEner ce

Foundation grant G B-20963 to the Co niferous Forest Biome, U. . AnalysisEcosystems, international Biological Program; and in part by theS

U. S. For ofestService Pacific Northwest Range and Experiment Station. We thank Samantha

Best and Ann Young for technical assistance, Douglas- M cGinty , Alan Tor-

renueva, and Robert Fogel for field

assistance, and J. Benton Jones, Jr., andLaraine L. Glenn for nutrient analyses.Con

tribution No. 209 from the Eastern Deciduous Forest Biome,International B iological Program and con tribution No. 175 from the ConiferousForest Biome.

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Magnesium Flux in a Mature HardwoodForest Watershed and an Eastern White Pine Forest

W atershed at Cowceta, M.S. Thesis,University of G eorgia, Athens, Ga.

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J. B. Cragg (Ed.), Vol. 2, PP. 101-157, Academic Press, Inc., NewYork.B urnett, J. 11., 1968, Punda mentals of Mycology, St. Martin's Press, New York.Carlisle, A of •, A. 11.

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Nutrition Content of Freshly Fallen NeedleLitter of Some Northeastern Co nifers, Soil Sci. Amer., Proc., 8: 409-411.Cromack, K., Jr., 1973, Litter Production and Decomposition in a Mixed HardwoodWatershed and a White Pine W

atershed at Cowecta Hydrologic Station, North Carolina,Ph. I). Dissertation, U niversity of Georgia, Athens, Ga.Crossley ; D. A., Jr., 1970, Roles of Microflora and Fauna in Soil Systems, in

Pesticides inthe Soil: Ecology, Degradation and Movement,

International Symposium on Pesticidesin the Soil, pp. 30-35, Feb. 25-27, 1970, M ichigan State University.—, and M. P. H oglund, 1962, A Litter-Bag Method

for the Study of MieroarthropodsInh:t biting Leaf Litter, Ecology, 43, 5 71-573.Day, Jr., 1971, Vegetation S ( r ucture of a Hardwood W atershed at Coweeta, AES.Thesis, University of Georgia, Athens, Ga.

ipsnwunne 41...rnuovr,omnew,rowesnovwx.,,,,,,zirprI,

r,4 <.....,,,,,,,...

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LIT'FLII PRODUCTION; DECOMPOSITION, NUTRIENT CYCLING

623

Edwards, C. A., D. E. Reichle, and D. A. Crossley, Jr., 1970, The Role of Soil Invertebratesin Turnover 01 Organic Matter and Nutrients, in A nd iVALS; of Temperate Forestco •ystetio , D. E. Reichle (Ed.), pp. 147-172,

Sprin ger-Verlag Ncw York, Inc.Ellis, U. ' II., G. Matrone, and L. A. Maynard, 1946, A 72% II,504 Method for theDetermination of Lignin and Its Use in Animal Nutrition Studies, J. Anita. Sci., 5,2335-297.

Foster, J. W., 1949, Cbcolical Acti^ities of Fungi, Academic Press, Inc., New York.Fuller, W. II., and A. C. Norman, 1943, Cellulose

Deco mposition by Aerobic MesophdieBa cteria from Soil. Ill. The Effect of Lignin,f, Bacteriol., 46, 291-297.Cosz, J. R., G. E. Likens, and F. It B ormann, 1972, Nutrient Content of

Lit terfal(on theHubbard 13rook E xperimental Forest, Ncw I lampshire, Ecology, 53. 769-784.Johnson ,P. C., and W. T. Swank, 1973, Studies of Cation Budgets in the SouthernA ppalachians on Four Experimental Wa tersheds' with Contrasting

Vegetation, Neology,54: 70-80.-Jones, J. U,, Jr., and M. II, \Varner, 1969, Analysis of Plant-Ash

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jnt erpretation ofScl erophylly, Amt. Rot., (Londou), 26; 551-561.—, 1961, A Nutritional In terpretation of Scl erophylly Based on D ifferences in theCheinic:.1 Co mposition of Sci crophyllous and Mesophytic Leaves, Ann. Bo t. (London),25: 168-184.eldyen, A., 1963, A itnal Ecology, Sir Isaac Pitman and Sons, London.Shook , C. D., 1966, An Ecological S

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Ovington, J. D., 1965, OrgaMe Production, 'turnover, and Mineral Cycling in Woodlands,Rea., 40: 29::1 - 3 36.--, 1962, Quantitative Ecology and the Woodland

Ecosystem Concept, in Adv..- CC S in.Ecological kc.scarch , J. B. Craft; (Ed.), Vol. 1, pp. 10 3-192, Academic Press, NewYork.

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P roduction and Mineral Cycling in TerrestrialVegetation, Oliver & Boyd, Edinburgh:Salisbury, E. .1., 1959, Causal Plant Ecology, in Vistas in Botany, W.13. Turrill (Ed.),PP . 1244 44, Vol. 1. Pergamon Press, Inc., New York.Stark, N., 1972, Nutricdt Cycling Pathways and litter Fungi, Biosciener , 22: 355-36o.Sykes, .1. ,\1., and 13. C, II,

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•

. .1 1 . r A

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624 CROMACK AND MONK

Todd, it. I.., K. Clomack, Jr., and J. C, Storincr, Jr., 1973, Chemical Exploration of theAcrohabitat by Electron Probe Microanalysis of Decomposer Organisms, Nature, 243.544-5.16.

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—, 1963, Use of Detergents in the Analysis of Fibrous Feeds, II. A Rapid Method for theDetermination of Fiber and Lignin, J. Ass. Offic. AKric. Chem., 461 829-835.

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and D. A. Ctoydey, Jr., 1966, The Role of Arthropods and Microflora in Breakdown ofWhite WE Litter, Pe.lobiologia, 6: 293-303.

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Errata:

page 617, second paragraph, last sentence. Change.. . m three orders of magnitude greater" to "twoorders of magnitude greater".

page 617, next to last paragraph:. Delete Douglas-fir from the last sentence.

3. In table 4, page 618, reduce all phosphorus values

by 10 for western coniferous, western deciduousand western broadleaf evergreen species. Alleastern deciduous and coniferous species values

for P are correct.

!L page h20., second paragraph , last sentence:' TheAlexander, 1961 reference was left out in referringto critical level at which P is immobilized in

organic material.

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