1

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USDA

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.

Forest EcobgyandManagement 70 (1994) 311-318\

j

Effect of burning and brush treatments on nutrient and soil .physical properties in yoting longleaf pine stands

William D. BoyeP, James H. Miller! Fores service. Sourhem Forest ExperiiFjterrt Station. George ?iJ Andrew ForestqSciences~Uboratoty. Auburn -C%vrsity.

Aubvm. AL 36849. USA

Accepted 22 March 1994

FOUNDING EMTORFOUNDING EMTORLaurenoe L R&e, Murrbe, IrelandLaurenoe I.. Roche, Mu&, lreland

EDlTORS-lN-CHIEFEDlTORS-lN-CHIEFFor the Amer icas, Austra l ia ;For the Amer icas, Austra l ia ; ‘ . ‘ f‘ . ‘ f I)I)Newzealand and the Pacifii:Newzealand and the Pacifii:

&&&&at~&*&y _ _ *.&&&&at~&*&y _ _ *.

..E.P. Farrell‘E.P. Farrell‘

RF. FisherRF. Fisher Resouw ManagementResouw Managemento fo f Forest Sciehce+ Texas A&MForest Sciehce+ Texas A&M

, TX 77843-2135,, TX 77843-2135, USAUSAUnivemityUnivemity ~$fublki,.~e~ _ _~$fublki,.~e~ _ _

ROOK REviEW EDlTORROOK REviEW EDlTOR0. Binkley, Forest amI Wood Sdenca, Cdorado State UnM&ty, Fort Collins, Co 80523, USA0. Binkley, Forest amI Wood Sdenca, Cdorado State UnM&ty, Fort Collins, Co 80523, USA

EDlTORIAL ADvIsqrRY BOARDG. Atbtha$sen, Agricultural University of Norway, As,

M.;s&y Genetics & Fore&

F. Andersson, &v&ii Un-&etsity of A&u?@ral Scien%%s,

Rd!&%%%%%te of Terre&&l Eooloov. Penicuik.klkllothii. OK

1s.

J.Dk~$sti$e of Terrestrial Ecology, Peniiik,

. A-E.%R i o

;, l~n$on~I&Mute of Tro#cal Forestry,

ill;rx;cnCRAF Agr&forestry Pr@ct,

Maine. Or&i, ME, USAtUtiiv~t l&kGemlany

D.cI~~ofEdii~jEdinbu ,UK9sh Forest Research Institute. antaa.

Finiand -..JJl ‘f&Lean. Uniirs*~ of Bitish Columbii Vancouver ,

J.B. Hall, Unii College of North Wales;,“SG”JJ.aklavei Land Consultants, Wqneroo,

, UK.A,

P. l-bgberg, Ukkxsity of l@e&,,Um+,Wenyue Hsiung, Nanlrng Fore&Y UmveE Klmo, ~nivetsity of Agriculture, B

B.h?s Kera4 Agricultural University Karata IndiaJ.P. Lanly, i.A.0.. Forest Resources Divi&m. F&&e, ltalyJ.N. Long, Utah State University, Logan, UT, USA

GENERAI. INFORMATION~~~~~~~~~~~~ub~~s 8(

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ELSEVIER Forest Ecoi~gy kid Manageme&’ ( 1994) 31123&8

Effect of burning atid br&h%~eatments otitibtrient and soil:’ ~ >I..physical pmpetiies in Young kmg&pine stands

Wilbm Dc Boy&$ ~Jams H. M&x ’ n (USDA Foresi Service, Southern For&t Experiment St&o&,’ George W. Andrews &&try Sciences Laboratory, A&burn University,

Auburn. AL 36849, USA ‘,

Accepted i2 March 1994 :

Abstract

Over a period of 16 years, unburned longleaf pi& @inus palustris Mill. ) pole’hands ,grew an average of 27%more volume than similar stands regularly burned. Treatments includedbieu&al burr& in w&&r, SpriugzI,andsummer plus au unburned check, each of which was combmed with thre&upp&&mal treatments; namely, initial ’herbicide injection of all hardwoods, repeated handchxring of #l woo&y stetis,, and’ no treatment. All unbum&and winter-burned plots were paired to study this growth reduction relative tb t&&m&is. The Stat& of tiitrogen,’phosphorus, available moisture holding capacity, bulk density, and macrbpore spitce was determined in both SUF-

face and subsurface soils. Foliage from pines on Sampled plots was analyzedfof N, P, K, a, Mg, Mn, Cu, Fe, andZn. Burning did not significantly affect either soil N and Por foliar nun&n& Hewever, bumiugteduced availablemoisture holding capacity and macropore s,paee.,a& inseased the.bulk density of surface soils, and also reducedthe moisture-holding capacity of subsurface soils..The.results from this and otherstudies suggestthat growth gassesare due, at least in part, to increased moisture stress associated.lvith changes in soil physical properties. _.

Keyworok Prescribed fire; Pinus pulustris Mill.; Stand gmwth, Hardwood control; Hetbicide injection; U&ermry clearing

1. Introduction , this growth- pxiwziiin as&iated;with burning are

Over a period of 10 years, biennial wiiter,no& ~ppan2nt.; R&e i&x&ties were low, partly asa result of W3fqzx#&noy of burning that kept fuel

spring, and summer prescribed fires significantly aequmulatio~ low,, Crow,a sc6rch did not seemreduced the height and diameter growth of sa- to be a faotor, av&raging Loss :than 5Oh for fourpling-pole size longleaf pines (Boyer, 1987 )- successive, series of winter burns. Crown scorchStand volume growth from Age 14 to 24 aver- alone is not’expqsedto affect pine growth before ’ .aged 27% higher in unburned than in burned it exceeds omz&ird of the live crown (Waldropstands. Growth wasunaffected by season of bum. and Van Lear, 1984; Lilieholm and I-& 1987).The volume growth differential between burned Results from p&St studies have been inconciu-and unburned stands has continued at the same sive. Surf?= fires of low to moderate intensityrate to stand Age 30 (Boyer, 1994). Reasons for have reduced growth of small tongleaf pines

(Garren, 1’943; W&hle&xxg, 1946; Bruce; 1947 ),* Corresponding author. presumably’ ovving -to defoliation, but were not

,

3 1 2 @!D. Boy&, J.H. Mi&er / Eore~t E~obgpanci Mamgernenr 70 (1994) 316318 . I

expected seriously to affect development of this 1993). Replacement of this N loss can occur ”fire-tolerant spe&es ~onc.e beyond the sapling through fir&stimulated symbiotic and non-Sym- ‘-stage. In mature or maturing longleaf star@& no l&&k .N-fixqitkm ir(~;Wells3-~ 197 1; Jorgensen and ”growth reductions were observed with periodic Wells, 1971). Thus, non-volatile nutrients areburning (Garren; 1943; Sackett, 1975 ), whereas mineralized throu&h combustion and trans-annual burning for long periods (30 years) re- ported into the soil, and volatile N is lost, al-duced both height and diameter growth ,(Bruce, though N repiaoement can be initiated with a fire1947). Stone (1942) repap&hat ,$i$~ mu&xl ey@ ~~~~~~t.~~~ 1987).in diameter growth reductions over a vGd& rani& *!“r’kies$$~$ &#&&&e tis sti#e&mfio$ed on’

of tree sixes, indicating that frequen@ burned a l@@$erm b&n&@ &u&7 (.Royer, B&7) to de-longleaf pine stands will be unsuitable for prep- terminewhether 10 years-of repeated prescribedaration of yield tables. Diameter .growth reduc- fim, & ~bi&#&n with additional woody planttions have also been noted in. 2% to 3Qear+% ,, “: eenu@l .treatme&s, had* affe@ed sojl N and P., . ,longleaf stands durini the year fohowing a’win- .av&abie @$stuge ho$ding capacity, bulk den-ter burn (Zahner, 1989). This growth loss was sity, and: ‘ms?ropore .space or the nutrient statusgreatest in dry years, and less or non-existent’in of pine .foti&@. ff causes for the redu&d pine vol-wet years, suggesting a,cotmection with moisture ume growth assisciated with burning oan be iden-availability. tified, ‘it mtiy become possible to manage’ pre-

Early studies of surface fire effects on nutri- scribed fires so as to obtain the needed benefits “Itional and physical properties of soils i.n the with minimum impact on growth.southeastern coastal plain suggested;fliat, on&al- : , :

ante, fm had slightly de$imenta! eff&& on !I.. * I. ., , . )’physical properties a~dslightly,$zm$c&&xt~ &~&tici;&?j&-l @+&a.~u~ef :,on nu t r i t ion . Sou the rn so& prote&&f&& ‘fift i j ., c - :- ‘ i , ’ ‘. ; : . _were more penetrable and porous than &e- _quently burned’ soils (Wahlenberg, 193%;, Hey- I* i 1. $tu& @fix i ,:_ ‘.. ”ward, 1936), but recovery appeared rapid after .** . .-,Iburning was stopped (Heyward, 1937 ). Early re- t The g@&y w!&p&&&&. @73 to &-g&f, .ports also indicated that surface soi$‘,in:&rned $eh@. ~&&&~~ &&&$ cof”~ve&;hardwood -stands tended to have higher levels of N;Ca, @d i e&rtr&L @&r&erzt$ on ‘rs&mp&&n and-stmctnreother minerals, more organic matter, and h&her of the untiersto@+d also growth of overstory,1 ..:pH ( Wahlenberg, 1946 ). pin&The s$dy was esf&i&d on the Escambia ’

More recent studies tend to support these ear- Experimental Forest (mG&&ed~by ,the Seuth-her findings and expand the understandingof.fire ern Forest Experiment Station, USDA Foresteffects on N pools and losses. Periodic buds -in” ‘~Service, in coop&ration with the T.R. Miller MU .tcoastal plain pine stands increased ma&&- ~ CC& ) ,.in southwest Alabama, on a typical coastaltxients (Metz et al., 1961; Hough, 1981;~~cKee; ‘, plain lon&afp&e site. Soils were primarily fine1982; Linnartz, 1984) while not adve&ly af: sands -of ‘the ~Troup: seties (lo-y,- &he&&s, G -’fecting N and organic matter in surface soils :: thermic CrossarenicP~euduhs)!~ with some Do- ,-(Metz et al., 1961; McKee, 1982). fncreased than (fine-&amy, siliceous, th&mic.Ptinthic Pa-availability of N in surface soils following-burn-. ^ Ieudults), Wagram: (loamy; $ie$aus;, .thermic + -.ing has been reported (Schoch and &&ley, Arenic PaJeudults), and Fuquaj~ (do&y, sili- ’ ”1986). However, losses in forest floor N and or-. ceous, thermic Arenic Plinthic Kandiudults) se-- ’ganic matter were shown by McKee ( 19.82 ) with r&s represented, ail with slopes of less thtin 5%. :increased frequency of burning. Furthermore, N Study areas contained even-aged longleaf pineis lost from burned forest sites through volatili- stands originating primarily from the I958 seed ‘.zation, from 30% to ,600/o of fuel cu~tent!‘(Del&ll cro-Q and released from residual seed ‘trees inand Ralston, 1970; Wells, 197 I; Vase and Swank,;’ winter I96 I. All study areas had been, periedi- -0

,

W.D. Bayer, J.H. Miller /FOYest Ecology and Management IO (1994) 31 I-318 313

tally burned in the past, although the last fire be- cm depths and corn&sited by plot and depth. Atfore the study began was in Jam&y 1962. every other sampling point (six per plot) two

undisturbed soil core samples (45 cm3) were2.2. Study design taken from the 025 cm and 15-20 cm depths.

Samples were collected in July 1984, after a bumThe parent study consists of three,-separate in the preceding January, and held in cold stor-

blocks, each with 12 square 0.16 ha plots. All age (4°C) until analyzed.plots were thimed to a density of 1236 trees ha-’at study establishment. After thinning, residual 2.3.2. F&age

~pines averaged 6.7 m in height, 8.1 cm diameter;

at breast height (dbh) and 6.9 m* basal areaFoliage was collected from five randomly se-

ha-‘. Indicated Age 50 site index (Farrar, 198 1)lected sample pines on each net plot. Six com-

for the three study blocks, derived from domi-plete fa$cicles from the first growth flush of 1984

nant/codominant tree heights on unburned5 plotswere collected from each sample tree in late June

at Age 30, averaged 23-25 m. Twelve treatment1984, S&&les were placedsin plastic bags as col-

combinations were randomly assigned amonglected ax&t&en frozen until prepared for labora-

plots in each block using a factorial design, Eachtory analysis. Foliage from each sample tree was

of four burning treatments (biennial prescribed.collected again in spring 1986. Twenty complete

fires in winter, spring, summer, and an un-fascicles from,the first flush of l985 growth were

burned check) was combined with three suppIe-:taken from shoot terminals or laterals in the up-

mental treatments: ( 1) initial hardwoo&,contr$per third of the crown. All 100 ne@Ie fascicles

by stem injection with 2,4-D, (2) repeated hand:per plot were combined, frozen-on return from

,clearing of all woody vegetation 1.4 m or more

the field, and kept frozen pending laboratory

in height, as needed, (3) an; untreated, ,check.analysis.

Supplemental treatments were initiated in spring1973. All burning treatments beganwith acon?., 2.4. Analysis proc+?dures Iditioning winter bum in, January 1974, with as:signed seasonal bums beginning during the suc- 2Al. soilsceeding year. Burning techniques were adapted ‘Composite soil samples were air dried andto site and weather conditions. Sixty per cent of crushed to pass through a sieve of 2 mm mesh,plot burning was done with strip headfires, 24% Soil P was extracted from quadruplicate 5 g soilwith backfires, and the rest with flank fires. samples using 20, ml of a weak double acid’ solu-

This study began in 1984. The three unburned tion (vehlich, 1953) and determined using theplots were paired with the three winter-burned method desc&bed’~y lvlurphj and Riley (1962)plots in each block. As pine growth responses to yield an. estimate of available P. Total soil Namong seasons of bum were similar, causes are was. deter&red from quadr@$ate samples us-,expected to be the same. The winter ,burning ing ~ISjeldal$ digestion. and .anammunia-specifictreatment was selected because most prescribed- ion electrode (Bremner and Tabatabai, 1972;burning in longleaf pine forests has been done Eastin, 1?76). Available moisture holding ca-during the winter. pacity, as per cent by volume, was calculated

from undisturbed soil cores, using pressure ex-2.3. Field sampling traction. The d,i,fference in moisture content at

0.03 and 1.5 MPa tension provided an estimate2.3.1. Soils of the capacity. of sampled soils to hold moisture

In each of the six plots per block, soil samples within the range available for plant uptake. Soilwere collected from two depths alOng a diagonal. ,bulk density was ,determi~ned from cores aftertransect across 0:04”ha. net plot& !I’wel\Fe~ qamples oven-drying. tot’s coqstant weight at 105’OC. Pereach were collected .from,the O.-J 5;cm and 1.5~30 cent macropore space was obtaiped. from core

.

3 1 4 W.D. Boyer,J.H. Mili~~Forest~~~~~dMan~ 7~~~994)311-3ld .’

Table 1 : I.,.,_’Available moisture holding cap&y (mf m”’ ) ~~$hsoi&a.~af&+ted by hardwood control &eatments

,. ‘ . .

Burn Supplemental tregment ’ ,; Average‘1 ‘

inject ‘Cl&r” None; . I, ,,

O-5 cm depth ‘ , ‘I. ” . , .^.’None 15.3 12.2 /_(, lS.O,/ ” “PO 14.2aWinter 9.9 11.1, IO.0 s\ i0.3b ‘. IAverage _,12.6a . ‘11.6a 1X5& ’ 12.2 ‘~

_. ;15-20 cm dqpfh .,. ,,

None 13.1 1 2 . 9 ‘ I c u.9 : 13.3a,Winter 11.1 rw ‘ , . .1$x8 ‘ , ,y mgb ,, ,,Average

/12.la * 1 pi 12.3a 12.1 ,.- . __,,‘ i:. , “ ..,c.,_

Cohmm or row means foliowed by thg same letter da n@t&f&r~~~ the 0.05 @el r&&&k&e. ) . ’ ”,.’

Table 2 2.5, Stat&t&l andyesBulk density and macropore space of s&he soil saqpks * -.: _ , : i.~,

Bulk densi ty hh73pore Analysis ofv&&n& proi$edures were used to(Is=“) space(%) i

, -..,: : .-No burn 1.22 47.1 ‘.Winter bum .1.26 44.5

samples as the water volume difference between edand-m* ”saturation and 0.03 MPa tension. bmed pi&s &e ‘to a iignificaia-t.:.&&p& ,_ /J

i+a&on;“~ “1 :,- ,

2.4.2. Foliage 1 .: IIFoliage samples were oven-dried’ (7@@),

‘.‘$Resdts. ” ._ ‘, “: :

ground to pass a screenof0.0425 mm mtih, and “- - _’ - 1: . r;.,retained in cold storage for nutrierit~ana$yses. The 3.Js’sqils s _ ’ __ _y ’ L, _ ,.1984 foliage collection was a&y&d for“N “and _(.. _F ‘/,i’ ‘, : _P. The 1986 foliage collection w&san&zed for The av@able &roisture h6ldmg capacity ofN, K, Mg, Mn, Ca, Cu, Fe, and Zn. FoliarN was .w& was ~~~~~~~~~‘i~~~~, by 27$$&&~ ad ‘”determined from duplicate 0.1 g san&es ‘of 18% subsu&&e$with the v&ter+b!qn than withground foliage following the same procedure as the ~no&z& tre@nent (Tab@ 1 j+ Stpplemental sdescribed above for soil samples. Foliar mineral tr&f&&&&& nb &&$j~& &j&t. :, ,:.elements were determined from 0.5 g duph&te Winter burning significantly iqreased bulk ‘*. Isamples after dry-ashing at 450°C for at &a& 6’ .’ den&y and red&ed:per cent macropore space ofh. When cooled, 20 ml of 0.4 N HCl with 012% ‘s&face but not subsurface soils. Values fo.r sur- .‘lanthanum was added. After mixing and filter- face soil samples are given in Table 2. Suppie-ing, P was determined by the vanado-molybdate mental treatments had no effect on these twomethod (Jackson, 1958). The concentration of variables, “.., ;.

K., Ca, Mg, Mn, Cu, Fe, and Zn in clear, undi- “Burning did not sigr@zan$ly affect total N orluted extract was determined by atomic ,abSorp- (. ,available :P itiS3rh~r sutifaceor &b&&face soiis,‘tion spectrophotometry., ,’ z but, suppE@~otd’treaP~~~~s -didi,,Periodi.c feh-,I

.

E

V-D. Bayer, J. fk Miller / Foresl Ec&gy and Maiiagetneni 70 (1994) 31 I-318

Table 3Soil nitrogen-per cent as affected by hardwood control treatments

315

Bum Supplemental treatment

Inject Clear None

0- 15 cm depthNone 0.044a’ 0.045a 0.04laWinter 0.037a 0.057a 0.04OaAverage 0.04Ob’ 0.05 la 0.041b <\,

IS-30 cm depth : .None 0.032a 0.025a ’ 0.02OaWinter 0,018b.‘ 0.034a 0.026aAverage 0.025ab 0.029a 0.023b

’ Column or ’ row means foilowed by the same letter do liot differ at the 0.05 level of significance.,

Average

i0.044a0.044a0.044

0.026a0.0X+0.026

Extractable soil phosphorus (mg kg”) as~afkcted by hardwood control treatments

Bum Supplemental treatkknt . Average

Inject clear None

O-15 cm depthNone 0.38a’ 0.52a 0.7OaWinter ’ 0.75a 0.8Oa 0.52aAverage 0.57a2 0.66a 0.61a

i 5-30 cm depthNone 0.33a 0.33a 1 0.47aWinter 0.2Oa 0.53a 0.33aAverage 0.27b 0.43a 0.4Oab

’ Column or 2 row means followed by the same letter do not differ at the 0.05 level of significance.

0.54a0.69a0.61

0.38a0.36a0.37

Table 5Nitrogen and phosphorus contents in pine needles

.Nitrogen Phosphorus

No burn 0.88 0.03 Table 6Winter bum 0.84 0.03 Nutrieni values for burned and unburned plots

Winter bum No bum

ing of woody plants resulted in significantly more(mg kg-‘) 6% kg-’ 1

N in surface soils, and also raised N levels in sub- 8900 9000.surface soils compared with plots where woody

~~~~~m3490 3200

understory plants were not cut (Table 3). The C a l c i u m 2250 2080

highest N levels were found on plots where woodyM a g n e s i u m 950 990Manganese 170 201

plants were repeatedly cut and burned. Iron 28 30Nitrogen consistently ,decreaskd with depth, Zinc 26 24

and the lowest level of N occurred on plots that capper 2.8 3.1

316 W.D. BOJPC J.H. killer / Fo& Edogj a&i &k&qy& 70 (1994) 31 l-318

were both injected and burned. This was signifi- ityy, increased bu1.k density, and decreased ma-cantly lower than on injected-unburned plots. cropore space in surface soils; and also with a re-Contrary to some earlier reports (Metz et al., . duce&l moisture retention capacity in subsurface1961; McKee, 1982), frequent burning alone did. . soils. Wahlenberg et al. ( 1939) reported that an-.‘, not consistently result in a higher level of N in nuai burning on a coastal.plain soil increased soilthe soil. bulk density from 1.3 to 1.4 g crne3 and reduced f

Available P was not significantly affected by : porosity’ from 42 to 40% compared with similar.supplemental treatments in surfaoe soils. How- unburned soils, T&se changes are &nil@ toever, available P in sub&rfaee soils was signifi- those observed in this study. R&ton and H&t?&cantly higher on repeatedly cleared plots than on ell ( 197 1) noted that, with repeated moderatethose with the injection treatment (Table 4 ) . For burning over long, periods, decreases in macro-all supplemental treatments combined, available pore space, infiltr&on, and ~aeration can be- a-P, like N, was not consistently higher with pected. This results f&n exposure of mineral soilb u r n i n g . to r@ifi&. titlj ~cq3.nsegu3gt dispersal of, aggre-

gate5 ,that can clog soil pores (Bower, 1966;3.2. Foliage Moehring et al., 1 g-66). They also noted that re-

ductions in percolation rates are sometimeS ob-,..Neither burning nor supplemental treatments ~served .&er fiti& ,032 s&n& soils, as a, result of re-

significantly affected N and P content of pine fb- sistanoe to- wet&g that impedes downwardliage collected in 1984. Percentages of these ele- infi&ration of water. However, when prescribedments in pine needles are given in Table 5. fires do not completely remove surface organic

Nutrient analyses of foliage’collected in’ i986 ma&r, changes i!n ‘in&&ion and ‘pore &Race 1also did not reveal any significant differences may be toosmall to detect,owing to burning or supplemental treatments. The negative in&et of biennial burning ,onThe average values for burned and unburned‘ pine growth could&e associated with the regularplots are given in Table 6. removal of surface organic matter. The physical

removal oflitter from a longleaf pine pl&.&tionresulted ‘in a diameter growth reduction du.r&rg

4. Discussion and conclusions the year immediat+ly following removal fMc-Lead et al., l-979 ) ., Th$ rapid response to’ l&&r

Biennial winter burning did not significantly removal su@ested*a change in water entw andaffect either the N or P content of soils or of infiltration, as macronutrient concentrations inIongleaf pine foliage when compared-with simi- pine fohage were not, aqected. Purther?:.meirsiiie-lar stands unburned for 22-24 years, Burning also merits of xylem pre&re ‘potential indxc&ed in-did not affect foliar content of K, Ca, Mg, Mn, creased moiSure.stress in TV&% onplots with lit-Cu, Fe, and Zn. The longleaf pine growth reduc- ter removed compared with treea where littertions associated with biennial bums in this study remained intact @inter et @., 1979). This dif-do not seem to be due to changes in nutrient ference in ,moisture stress lfKtw.een treated tindavailability and utilization. This supports con- control plots continued throughout the first sea-elusions from a number of studies reporting no son regardle\ss of rainfall or drought. Heywarddeleterious effects of periodic fire on nutrient ( 1 $J+) ~‘jy&+ e& f&j& &%&tu& &etent ofavailability when low-intensity buming,.was used, surface so?l’hor$ons in unbur&d compared withexcept the volatile loss of N from the site (De- burned lotig@&f pine stands, suggested that theBell and Ralston, 1970; Wells, 197 I). Wells’s mulching effects of surface organic htter on un-( 197 1) findings in a pot study suggest less N burned soiismight be’partly responsible.availability on burned soils. It is dif%.iIt to believe that the relatively small

Biennial winter burning was associated w&h a changes .in soil &&ire holding capacity ob-significantly reduced moisture retention capac- sew& irI bs’hdy cotild be reSpons&ie for an

.

W. D. Bow, J. H. Miller /Fore;, Ec$ogy and Mawgenrenr 70 (I 994) 311-318 317

annual pine volume growth reduction in all win- affect soil moisture availability and uptake by theter-burned plots averaging 2.17 m3 ha-’ (20%) tree.for the 16 years from Age 14 to 30. The averagqdiameter growth reduction over this period was9%. Zahner (1989) reported that, withother Referencesfactors held constant, longkaf pine ring widthduring the year following a winter burn was re- Bower, D.R., 1966. Surface soil recovers quickly after burn.duced an average of 13% in stands from 20 to 28 US For., Serv. Res. Note, S&46,2 pp=years old. A 12% diameter growth difference inmature longleaf stands (50-60 years old) wasobserved between unburned stands and similarstands with biennial prescribed fires (Bayer,

Bayer, W.D., 1987. Vohunegrowth loss: a hidden cost of pe-riodic prescribed burning in longleaf pine?South. J. AppLFor., 11: 154-157.

Boyer, W-D., 1994, Eighteen years of seasonal burning :nlongleaf pine: effeits on ‘overstory growth. in: Proeeed-

1987). Highly variable reductions in longleaf ings 12th International Conference on Fire and Forestpine radial growth following fire were reported

)Meteorology, 26-28 October 1993, Jekyll Island, GA So-

by Stone ( 1942). Radial growth reductions av- eiety of American Foresters, Bethesda, MD, pp. 602-6 IO.

eraged 23% during the year after a bum for DinesBremner, 3-M. and,Tabatabai, M.A., 1972. Use of ammonia

under 15.2 cm dbh, and 19% for larger trees.,electrode for thi‘determinacion of ammonia in Kjeidahlanalysis of soils. Commun. Soil Sci. Plant AnaL, 3: 159-

Stone assumed that growth reductions were due 165.

to defoliation by fire. Bruce, q., 1947. Thirty-two years of annual burning in lon-

The major impact of fire (Stone, 1942; Zah-gleaf pin& J. For,, 45: 809-S 14. \

DeBell, QS. and Ralston, C-W., 1970. Release of nitrogen by =c.ner, 1.989) or surface litter removal (McLeod et burning light forest fuels. Soil Sei. Soe. Am. Proe., 34: 936- Ial., 1979) on longleaf pine growth occurred dw;- 938.

ing the first year after treatment, and was much Eastin, E-F;, 1976. Use of ammonia: electrode for total nitro-

diminished or absent in the second year.- The fact gen determination in plants. Commun. Soil Sei. Plant

that growth responses to both litter removal and ”Anal., 7: 477-481.’

Farrar, J?, R.M., L98f. A‘site-index function for naturalIyy-fire were so similar suggests that causal factors regenerated longleaf pine in the east Gulf area. South. J.are more probably related to changes in soil Appl. For.; 5: 156-l 53.

moisture conditions than to defoliation from Garren, K.H., 1943. Effects of fire on vegetation of the south-

crown scorch. That the growth reduction is con-eastern United States. Bot. Rev., 9: 617-654.

fined largely to the first year after fire or litterGinter, D.L, McLeod, K.W. and Sherrod, Jr.,C., 1979, Water

stress in longleaf pine induced by litter removal. For. Ecol.removal is not entirely consistent with a perma- Manage., 2: 13-20.

nent change in soil physical properties resulting Heyward, F., 1936. Soil changes associated with forest fires

from the absence of fire; unless, of course, con-in the longleaf pine region of the South. Am. Soil Survey

ditions improve so rapidly that by the second orAssoc. Bull., 17: 4 I-42.

Heyward, F., 1937. The effect of frequent fireson profile de-third year after a fire, soil moisture availability velopment of longleaf pine forest soils. J. For., 35: 23-27.and related tree growth conditions are essen- Heyward, F., 1939. Some moisture relationships of soils from

tially equivalent to those within a stand un- burned and unburned longleaf pine forest soils. Soil Sci.,

burned for 10 years or more.47: 3 13-327.

Hough, W-A., 198 1. Impact of prescribed tire on understoryThe results to date strongly suggest that long- and forest floor nutrients. US For. Serv. Res. Note, SE-

leaf pine growth reductions associated with pe- 303,4 pp.

riodic prescribed fires may be due to changes in Jackson, M.L., 1958. Soil Chemical Analysis. Prentice-Hall,

soil-tree moisture relations. Further investiga-Englewood Cliffs, NJ, 498 pp.

tions on the impacts of prescribed burning onJorgensen, J.R. and Wells, C.G., 197 1. Apparent nitrogen tix-

ation in soil infiuenced by prescribed burning. Soil Sci.longleaf pine growth should concentrate onchanges in the physical properties of the soil, in-eluding soil organic layers, and possible changesin surface fine-roots and mycorrhizae, that may

Sot. Am. Proc., 35: 806-S 10.Lilieholm, R.J. and Hu, S.C., 1987. Effect of crown scorch on

mortality and diameter growth of 1 g-year-old loblolly pine.South. J. Appl. For., Ii: 209-211.

Linnartz, N.E., 1984. Effects of prescribed burning on some

318 W. D. Bqver, J.H. ~flik?r / l%wsl Ecologv and Management. 70 r&994) 31 l-318

chemical properties of forest. soifs. ,LSU ForestryNotes~ pine: US:For; Seti. South. For. Exp. Stn. Note, 43,3 pp.No. 141, Louisiana Agric. Exp. Sm., Baton Ruuge, LA, 2 Vase, J.M +tx$ Swank W,T., 1993, Site preparation burni.ngP P . to improve s&u$ber$ Appalachian pine-hardwood stands:

McKee, Jr., W.H., 1982. Changes in soil fertility following aboveground biomass, forest floor mass, and nitrogen andprescribed burning on coastal plain pine sites. US For. earban pbo’fs‘.l=sin. J. For. Res. 23:. 22552262.Serv. Res. Pap., SE-234,23 pp.

McLeod, K;W., Sherrod, Jr., C. and Porch, T.E., 19 79. Re-WahJenb&g, W&, f935: Ef%c;bf’fire and grazing on soil

properties.and.the znaturai reproductionof longleaf,pine.sponse of longleaf pine plantations to litter removal. For.Ecol. Manage., 2: 1-l 2.

J. For.,~33:.33&338.

Mehlich, A., 19 53. Determinations of P, Ca, I$& ‘K Na, and 1WahJenbexg, W.G, 194.6. LongJeaf pine: its use, ecology, re-

aNH, by North Carolina soil. t&ing laboratories. North j

ge*er&ii,~+&&i0*~ growth, and .management.‘Charles~

Carolina State University, Raleigh ‘(mime0 1.Latbrop Pa@ For&y Foun&tio& ,in ,&operation wi!hUSDA’For. S&%5 Washington; DC, 429 pp.

Metz, LJ., Lot& T. and Klawitter, R.A., 1961. Some effectsof prescribed burning on coastal plain forest soil. US For.

WahIenberg, W.G., Greene, SW, and Reed, HR., J 9 39 ;‘E&

Serv. Sta. Pap., 133, 10 pp.fects of&ead‘d cattle grazing on long&af pine lands as

Moehring, D.M., Grano, C.X. and Bassett, J.R; 1966. Prop-stq*ti..,at.:##eiIls Missipippi. US ,Dep. &ric.Tech.

et-ties of forested loesssoils after repeated prescribed burns. BdL,683,3@3p. ;

US For. Serv. ResNote, SO-40,4 pp.Waldrop, T.A. and VanLear, D-H., JY84. Effect of crown

”Murphy, J. and Riley, J.P., 1962. A modified s&r& spiutionseomh on survival and growth of young Ioblolly pine.

method for the determination of phospbate in natural ’ So&. JIi$pLF&, 8; 35-40. ,’

waters. Anal. Chim. Acta, 27: 3 l-36. Waldrop,%%, Va&&ar, D.H.&loyd, FT. and Harms, W.J$, 3

Ralston, C.W. and Hatchell, G.E., 19 7 1. Effects of prescribed. 198X I.&-terni stud&of prescribedburning inioblolly

burning on physical properties of soiJ. In: Proceed&s,. pin+ forests ofthe southeastern coastal p!ain. US For. Serv.Prescribed Burning Symposium0 14- 16 April ,197 1 J G&p. Tech. Rep., SE-45,23. PP.Charleston SC. US For. Servo, Southeastern For. Eq?, Stri., 0 WelIs, CG,, 1971. E&cts of p&cr&d -bu&ig on soilAsheviBe, NC, pp. 68-85. && &$&$$ ~~:n~~~ent,~~~bili2y.,. In: pros _

Sackett, S.S., 1975. Scheduling prescribed bums for hazard &&&$&&Qjgij I~u~~&2#xq@i41, _ J4- 16 Aprilireduction in the southeast. J. For., 73: 14%147+ r97@Cha$estcm, SC. US For.Serv.~Southeastem For.

Schoch, P. and Binkley, D., 1986. Prescribed ~burning in- Exp. Stn, Asheville,NC, ppl .86-YY. icreased nitrogen availability in a mature loblolly pine tin;er, &; :i $%Y,~Tme&+g ties relate&o stand and: em+;stand For. Ecol. Manage., 14: 13-22. ronmental factors in,&th Alabama h&leaf pine stands.

Stone, E.L., 1942. Effect of fire on radial growth of longleaf US Fat. Serv. Gen. +I%&. Rep,, Sb-74: 1 Y3*19 7.)_

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