The effects of slash burning on ecosystem nutrients during the land preparation phase of shifting...

14
Plant and Soil 220: 247–260, 2000. © 2000 Kluwer Academic Publishers. Printed in the Netherlands. 247 The effects of slash burning on ecosystem nutrients during the land preparation phase of shifting cultivation Christian P. Giardina 1,3 , Robert L. Sanford Jr. 1 , Ingrid C. Døckersmith 1 & Victor J. Jaramillo 2 1 Department of Biological Sciences, University of Denver, Denver, Colorado 80208, USA; 2 Instituto de Ecologia, Universidad Nacional Autonoma de Mexico, Campus Morelia A.P. 27-3 Xangari, Morelia, Michoacan C.P. 58089 Mexico; 3 Present address: Department of Agronomy and Soil Science, University of Hawaii-Manoa Beaumont Research Center, 461 Lanikaula Street, Hilo, Hawaii 96720 * Key words: cations, fire, nitrogen, nutrients, phosphorus, slash-and-burn, soil, tropical forests Abstract The most commonly observed change in soil following slash-and-burn clearing of tropical forest is a short-term increase in nutrient availability. Studies of shifting cultivation commonly cite the incorporation of nutrient-rich ash from consumed aboveground biomass into soil as the reason for this change. The effects of soil heating on nutrient availability have been examined only rarely in field studies of slash-and-burn, and soil heating as a mechanism of nutrient release is most often assumed to be of minor importance in the field. Few budgets for above and belowground nutrient flux have been developed in the tropics, and a survey of results from field and laboratory studies indicates that soils are sufficiently heated during most slash-and-burn events, particularly in dry and monsoonal climates, to cause significant, even substantial release of nutrients from non-plant-available into plant-available forms in soil. Conversely, large aboveground losses of nutrients during and after burning often result in low quantities of nutrients that are released to soil. Assessing the biophysical sustainability of an agricultural practice requires detailed information about nutrient flux and loss incurred during management. To this end, current conceptual models of shifting cultivation should be revised to more accurately describe these fluxes and losses. Introduction Shifting cultivation is practiced by 300 million people annually, and affects 400 million ha of the planet’s 1500 million ha of arable land (Brady, 1996; Kleinman et al., 1996; Seubert et al., 1977). Effective management of fire-converted land for sustained pro- duction relies on detailed knowledge of the fluxes and losses of nutrients incurred during and after the burn- ing of slash biomass (Raison, 1979). This information is particularly important for making land management decisions when fertilizers are used minimally. Studies of shifting cultivation have attempted to assess the bio- physical sustainability of shifting cultivation primarily by identifying the magnitude and persistence of in- creased soil fertility following slash-and-burn clearing of forest land (De Rouw, 1994; Juo and Manu, 1996; Nye and Greenland, 1960; Seubert et al., 1977) and * FAX No: 808 9744110. E-mail: [email protected] secondarily by quantifying fluxes and losses of nu- trients from these converted ecosystems (Giardina et al., 2000; Kauffman et al., 1993, 1995). These data have not yet been used to rigorously evaluate currently accepted conceptual models of nutrient flux during slash-and-burn that were developed nearly 40 years ago. Shifting cultivation is a multi-step process that in- cludes site selection, clearing forest by slash-and-burn, cropping, abandonment and fallow. The slash-and- burn phase of shifting cultivation, the focus of this paper, most often includes cutting down forest ve- getation at the beginning of a dry spell or the dry season, drying slashed vegetation in place for weeks to months, and broadcast burning the dried slash be- fore the onset of the rainy season (Ewel et al., 1981; Maass, 1995; Nye and Greenland, 1960; Seubert et al., 1977). The most detrimental changes in site nu- trient status occur during this phase of the shifting cultivation cycle.

Transcript of The effects of slash burning on ecosystem nutrients during the land preparation phase of shifting...

Plant and Soil220: 247–260, 2000.© 2000Kluwer Academic Publishers. Printed in the Netherlands.

247

The effects of slash burning on ecosystem nutrients during the landpreparation phase of shifting cultivation

Christian P. Giardina1,3, Robert L. Sanford Jr.1, Ingrid C. Døckersmith1 & Victor J. Jaramillo21Department of Biological Sciences, University of Denver, Denver, Colorado 80208, USA;2Instituto de Ecologia,Universidad Nacional Autonoma de Mexico, Campus Morelia A.P. 27-3 Xangari, Morelia, Michoacan C.P. 58089Mexico; 3Present address: Department of Agronomy and Soil Science, University of Hawaii-Manoa BeaumontResearch Center, 461 Lanikaula Street, Hilo, Hawaii 96720∗

Key words:cations, fire, nitrogen, nutrients, phosphorus, slash-and-burn, soil, tropical forests

Abstract

The most commonly observed change in soil following slash-and-burn clearing of tropical forest is a short-termincrease in nutrient availability. Studies of shifting cultivation commonly cite the incorporation of nutrient-richash from consumed aboveground biomass into soil as the reason for this change. The effects of soil heatingon nutrient availability have been examined only rarely in field studies of slash-and-burn, and soil heating as amechanism of nutrient release is most often assumed to be of minor importance in the field. Few budgets forabove and belowground nutrient flux have been developed in the tropics, and a survey of results from field andlaboratory studies indicates that soils are sufficiently heated during most slash-and-burn events, particularly in dryand monsoonal climates, to cause significant, even substantial release of nutrients from non-plant-available intoplant-available forms in soil. Conversely, large aboveground losses of nutrients during and after burning often resultin low quantities of nutrients that are released to soil. Assessing the biophysical sustainability of an agriculturalpractice requires detailed information about nutrient flux and loss incurred during management. To this end, currentconceptual models of shifting cultivation should be revised to more accurately describe these fluxes and losses.

Introduction

Shifting cultivation is practiced by∼300 millionpeople annually, and affects∼400 million ha of theplanet’s 1500 million ha of arable land (Brady, 1996;Kleinman et al., 1996; Seubert et al., 1977). Effectivemanagement of fire-converted land for sustained pro-duction relies on detailed knowledge of the fluxes andlosses of nutrients incurred during and after the burn-ing of slash biomass (Raison, 1979). This informationis particularly important for making land managementdecisions when fertilizers are used minimally. Studiesof shifting cultivation have attempted to assess the bio-physical sustainability of shifting cultivation primarilyby identifying the magnitude and persistence of in-creased soil fertility following slash-and-burn clearingof forest land (De Rouw, 1994; Juo and Manu, 1996;Nye and Greenland, 1960; Seubert et al., 1977) and

∗ FAX No: 808 9744110. E-mail: [email protected]

secondarily by quantifying fluxes and losses of nu-trients from these converted ecosystems (Giardina etal., 2000; Kauffman et al., 1993, 1995). These datahave not yet been used to rigorously evaluate currentlyaccepted conceptual models of nutrient flux duringslash-and-burn that were developed nearly 40 yearsago.

Shifting cultivation is a multi-step process that in-cludes site selection, clearing forest by slash-and-burn,cropping, abandonment and fallow. The slash-and-burn phase of shifting cultivation, the focus of thispaper, most often includes cutting down forest ve-getation at the beginning of a dry spell or the dryseason, drying slashed vegetation in place for weeksto months, and broadcast burning the dried slash be-fore the onset of the rainy season (Ewel et al., 1981;Maass, 1995; Nye and Greenland, 1960; Seubert etal., 1977). The most detrimental changes in site nu-trient status occur during this phase of the shiftingcultivation cycle.

248

Figure 1. The conceptual model proposed by Nye and Greenland(1960) and Sanchez et al. (1991) for nutrient flux during slash burn-ing. Ash from consumed biomass is incorporated into soil resultingin an increase in soil fertility. Carbon and N are largely volatilized; Pand cations are efficiently transferred from biomass to ash and thento soil. During rain some cations may be leached. Soils are relativelyunaffected by burning, but are modified by ash after rainfall.

In 1960, Nye and Greenland reviewed availableinformation on changes in soil fertility followingslash-and-burn clearing of forest, and proposed the‘nutrient-rich ash’ hypothesis to explain post-burn in-creases in soil nutrient availability. Their hypothesiswas straight forward: burning converts slashed veget-ation into nutrient-rich ash that is deposited on the soilsurface and incorporated into soil by rainfall and cul-tivation (Figure 1). In addition to providing an input ofnutrients, the incorporation of ash into soil increasessoil pH, which in acidic soils can increase nutrientavailability. The direct effects of heat on soil nutri-ent availability were hypothesized to be small whencompared with those of ash.

Temperate field studies of slash burning (reviewedby Raison, 1979 and Walker et al., 1986) and labor-atory heating studies of tropical soil (Kang and Sajja-pongse, 1980; Sertsu and Sanchez, 1978) have shownthat the view of slash-and-burn summarized in Fig-ure 1 is over-simplified. Nonetheless, field studies andreviews of tropical slash-and-burn agriculture have re-lied on this conceptual model to explain increasesin post-burn soil fertility (Beck and Sanchez, 1994;Brady, 1996; Buschbacher et al., 1988; Capistranoand Marten, 1986; Christanty, 1986; De Rouw, 1994;Kleinman et al., 1996; Lal and Cummings, 1979;

Lessa et al., 1996; Maass, 1995; Sanchez, 1976;Sanchez et al., 1991; Seubert et al., 1977).

To adequately assess the biophysical sustainabilityof slash-and-burn agriculture, the quantity of nutri-ents released from above and belowground biomass,and soil organic matter during and after slash burn-ing should be examined (Raison, 1979; Walker etal. 1986), however few studies of shifting cultivationhave examined nutrients at this level of detail (Juoand Manu, 1996). In the following sections, we re-view extant studies of slash-and-burn agriculture toconstruct budgets for total and available nitrogen (N),phosphorus (P) and cation pools in the soil and above-ground biomass of tropical forest. We track changesin these budgets through slash burning, and review theconsequences of burning on biological properties ofsoil. Where tropical field data are not available, weinfer from laboratory or temperate forest studies.

Slash-and-burn effects on aboveground biomass

Nutrients are more concentrated in fine plant material,such as leaves, twigs and small branches, than in thelarger components of a tree. These fine materials makeup a small portion of total aboveground biomass, butoften contain a large portion of the total abovegroundnutrient stock (Buschbacher et al., 1988; Kauffman etal., 1993, 1995). These fine materials are the quick-est to dry following slashing and are the most readilyburned components of a forest (Kauffman et al., 1995;Raison, 1979). Total aboveground nutrient loss dur-ing slash-and-burn clearing of forest land is amongthe highest of any disturbance known. AbovegroundN can undergo large, nearly parallel losses with car-bon (C) during biomass burning (Raison et al., 1985),with release rates ranging from 30% to 90% of totalaboveground stock (Buschbacher et al., 1988; Kauff-man et al., 1993, 1995). In the years following a burn,atmospherically derived N in rainfall and biologicalN2-fixation can replace losses of N (Raison et al.,1985, 1993). Phosphorus in aboveground biomass canalso undergo large volatilization and convective lossesduring slash burning (Giardina et al., 2000; Kauffmanet al., 1993; Raison et al., 1985). The new sourcesof P to replace losses include non-plant available P insoil and in parent material (except in highly-weatheredsoils), the generally low levels of atmospherically de-posited P and fertilizer (Raison et al., 1985). Nutrientcations, such as calcium (Ca), magnesium (Mg) andpotassium (K), have higher volatilization temperaturesthan N or P, but are equally subject to convective losses

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during a burn and to erosional and leaching losses fol-lowing a burn (Ewel et al., 1981; Raison et al., 1985).As with P, the primary sources of these cations are par-ent material, precipitation and fertilizer. In addition tosubstantial volatilization and convective losses of nu-trients during biomass burning (Kauffman et al., 1993,1995; Mackensen et al., 1996; Raison et al., 1985),nutrients may be lost following a burn by wind andwater erosion of ash (Khanna et al., 1994; Raison etal., 1985).

Across the sites reviewed here, element content ofash was significantly correlated with nutrient contentof the standing crop (Table 1, Figure 2). On average,3%, 49%, 50% and 57% of total aboveground N, P,Ca and K, respectively, were returned to soil as ash.The slopes of the regression lines indicate that elementtransfer, as a proportion of standing stock, increased asfollows:

N� P= Ca< K.

Most N in consumed biomass will be lost from a site,and ash generally contains little N. However, the Ncontent of ash was sometimes substantial when stand-ing stocks of N were large. The large variance for Nin Figure 2 is consistent with the fact that N is muchmore susceptible to volatilization losses than P, Ca orK.

The 50% transfer of aboveground Ca to ash islarger than the 31–34% loss reported by Raison etal. (1985); however, large quantities of Ca are se-questered in tree boles and the low intensity firesexamined by Raison et al. (1985) would have a smallerimpact on these Ca stocks than the intense slash firesreviewed here. The reason why the Mg content of ashcorrelated poorly with Mg content of standing crop(R2 <0.01, P=0.92; Table 1) is not clear. Raison etal. (1985) suggested that organically bound Mg maybe more sensitive to volatilization losses than Ca or K,despite a high volatilization temperature for inorganicMg (1170◦C). Alternatively, the high recovery for Ca,K and P at the two sites with very little recovery of Mg(Mackenson et al., 1996; Zinke et al., 1978) may in-dicate that the Mg numbers from these two sites are inerror. Overall, when the wide range in burn conditionsand the diversity of methodologies are considered, thestrength of the relationships presented in Figure 2 aresurprising.

Fine ash is very susceptible to wind loss and post-burn losses of ash from a site may represent a siz-able export of nutrients, possibly equal to or greaterthan volatilization losses during a burn (Raison et al.,

1985). However, few estimates of ash loss from trop-ical slash-and-burn studies exist. Sampling for ash isdifficult because the distribution of ash is spatiallyvery heterogeneous, and ash is difficult to fully re-cover in the field. Ash is also very heterogeneous withregards to nutrient content. Fine gray or white ashrepresents a small portion of total ash, but is highlyenriched with nutrients and may represent a large por-tion of total ash nutrient content (Raison et al., 1985).Two of the studies included in Table 1 assessed wind-related losses of ash following burning (Giardina etal., 2000; Kauffman et al., 1993). In both, losses ofN and P in ash were substantial. At a wet forest site inCosta Rica, Ewel et al. (1981) also noted substantialash loss due to wind and water in the month followingburning. However, wind related losses were not quan-tified and leaching loss into deeper soils may not haverepresented transport off the site.

Studies of slash-and-burn have viewed the nutri-ents contained in ash as being wholly plant-available(Nye and Greenland, 1960; Seubert et al., 1977; VanReuler and Janssen, 1994). Few tropical studies, how-ever, have assessed the plant-availability of nutrientscontained in ash. Stromgaard (1984) measured plant-available P in ash from a Zambian miombo woodlandsite using a mild acid extract (Bray and Kurtz, 1945)and found that ash contained about 0.8 kg plant-available P per ha. In a temperate study, Ohno andErich (1990) found that 1.9–8.7% of the P, 30–72%of the K, 31–54% of the Mg and 54–87% of the Cain wood ash were extractable with heated ammoniumcitrate. The wide ranges were attributed to species dif-ferences in the source of ash. Khanna et al. (1994)found that 11% of the P, 60% of the sulfur (S), 54–69% of the K and<20% of the Mg and Ca containedin ash from burnt litter, bark and twigs ofEucalyptuspauciflorawere soluble in water. Nutrient availabilityof ash derived from tropical forest would be expectedto follow similar patterns.

With few exceptions (Alegre et al., 1988), mostfield studies of slash-and-burn document increasedsoil pH after burning (Table 2). Burn-related increasesin soil pH are due to the acid neutralizing capacityof ash (Fritze et al., 1994; Khanna et al., 1994; Nyeand Greenland, 1960; Ohno and Erich, 1990; Raison,1979; Romanya et al., 1994; Sanchez, 1976; Singh,1994) and to consumption of hydrogen ions duringthe combustion of organic acids in soil and the forestfloor (D. Binkley, pers. comm.). In acidic soils, in-creases in soil pH may increase the availability ofnutrients such as P (Romanya et al., 1994; Sibanda and

250

Table

1.A

shm

ass,

nutr

ient

cont

enta

ndlo

sses

afte

rtr

opic

alsl

ash

burn

ing

Loc

atio

nM

ean

Sta

ndS

tand

Nut

rient

cont

ento

fN

utrie

ntco

nten

tof

Nut

rient

cont

ento

fash

Re

fere

nce

Ann

ual

Age

Bio

mas

saS

tan

dB

iom

ass

ash

afte

rbu

rnin

gre

ma

inin

ga

fter

loss

es

Ra

infa

ll(y

r)(M

g/ha

)N

PC

aM

gK

NP

Ca

Mg

KN

PC

aM

gK

(mm

)(k

g/ha

)(k

g/ha

)(k

g/ha

)

Ch

am

ela

,Mex

ico

75

0>

100

110

944

2715

35N

D34

628

1169

6N

D90

75

ND

ND

ND

Gia

rdin

ae

tal.,

2000

Dry

deci

duou

sS

teel

ee

tal.,

1998

Per

nam

buco

,Bra

zil

790

nativ

eN

DN

DN

DN

DN

DN

D35

1234

847

61N

DN

DN

DN

DN

DL

ess

ae

tal.,

1996

b

Sem

iarid

deci

duou

s

Per

nam

buco

,Bra

zil

803

1674

551

37N

DN

DN

D16

20N

DN

DN

D7

9N

DN

DN

DK

auffm

ane

tal.,

1993

Sem

iarid

deci

duou

s

Lua

For

est,

Tha

iland

1400

ND

ND

143

1622

811

211

010

ND

567

24N

DN

DN

DN

DN

DZ

inke

etal

.,19

78D

ryse

mi-d

ecid

uous

TaiF

ores

t,Iv

ory

Coa

st18

854

350–

560

351

1725

474

298

2611

125

4212

4N

DN

DN

DN

DN

DV

an

Reu

ler

&Ja

nsse

n,19

93L

owla

ndev

erg

reen

2035

0–56

054

213

401

7820

127

811

847

70N

DN

DN

DN

DN

D

Per

u,Y

urim

agua

s21

0017

ND

ND

ND

ND

ND

ND

676

7516

38N

DN

DN

DN

DN

DS

eube

rtet

al.,

1977

Mo

iste

verg

ree

n

Pa

ra,B

razi

l20

8840

3434

27

311

3575

62

929

20N

DN

DN

DN

DN

DM

acke

nse

ne

tal.,

1996

Hum

idtr

opic

al40

9583

424

783

9527

04

517

716

54N

DN

DN

DN

DN

D(t

ree

s>7

cmdb

hha

d7

3120

79

291

4373

53

112

1522

ND

ND

ND

ND

ND

be

en

ha

rve

ste

d)

Pa

ra,B

razi

l20

88pr

imar

y29

213

9062

90N

D54

547

1847

7N

D21

1N

DN

DN

DN

DN

DK

auffm

anet

al.,

1995

Mo

iste

verg

ree

n

Pa

ra-B

razi

l20

88pr

imar

y43

523

0087

1285

ND

945

5247

640

ND

506

ND

ND

ND

ND

ND

Kau

ffman

etal

.,19

95M

oist

eve

rgre

en+

logg

ing

Ron

doni

a,B

razi

l23

54pr

imar

y29

020

7057

360

ND

420

157

2219

2N

D24

8N

DN

DN

DN

DN

DK

auffm

anet

al.,

1995

Moi

stev

erg

reen

+lo

ggin

g

Ron

doni

a,B

razi

l23

54pr

imar

y36

124

2062

955

ND

500

4035

486

ND

316

ND

ND

ND

ND

ND

Kau

ffman

etal

.,19

95M

ois

teve

rgre

en

Tur

rialb

a,C

osta

Ric

a27

008

5245

914

439

8123

996

1655

510

719

0N

DN

DN

DN

DN

DE

wel

etal

.,19

81P

rem

on

tan

ew

etf

ore

st

aA

bove

grou

ndon

ly.b A

shco

ntam

inat

edw

ithso

ilan

dun

burn

tsla

sh.

251

Table

2.P

re-a

ndpo

st-b

urn

soil

nutr

ient

cont

enta

ndav

aila

bilit

yfo

rst

udie

sof

slas

h-an

d-bu

rn

Loc

atio

nA

nnua

lS

tand

Sta

ndS

ampl

eS

ampl

eTo

tal

CTo

talP

Ava

ilabl

eP

Tota

lNM

iner

alN

Soi

lpH

Re

fere

nce

Ra

infa

llA

geB

iom

ass

dept

hnu

mbe

rP

reP

ost

Pre

Pos

tP

reP

ost

Pre

Pos

tP

reP

ost

Pre

Pos

t(m

m)

(yr)

(Mg/

ha)

(cm

)(M

g/ha

)(k

g/ha

)

Ch

am

ela

,Mex

ico

75

0>

100

110

0–2

12a4.

43.

298

105

731

.975

769

029

117

6.5

8G

iard

ina

etal

.,20

00d

Dry

deci

duou

s2–

512a

ND

ND

137

143

3.7

15.1

720

768c

cN

DN

DS

teel

ee

tal.,

1998

Per

nam

buco

,Bra

zil

803

1674

0–2

125.

15.

550

57N

DN

D44

343

3N

DN

DN

DN

DK

auffm

anet

al.,

1993

d

Sem

iarid

deci

duou

s2–

512

6.0

6.3

7277

ND

ND

522

558

ND

ND

ND

ND

5–10

126.

96.

910

111

7N

DN

D62

566

9N

DN

DN

DN

D

Ibed

an,N

iger

ia12

5015

ND

0–10

ND

2326

.7N

DN

DN

DN

D39

5446

14N

DN

D6.

69.

0L

ala

ndC

umm

ings

,197

9L

owla

nd

rain

fore

st

Lua

For

est,

Tha

iland

1400

9N

D0–

51

17.3

17.7

ND

ND

210

.195

097

0N

DN

D6.

06.

7Z

iuke

eta

l.,19

78D

ryse

mi-d

ecid

uous

Nam

Phr

om,T

haila

nd15

00ol

d33

00–

53

20.5

32.5

ND

ND

3.3

44.4

1500

2100

11.5

33.6

6.3

7.2

Kyu

ma

eta

l.,19

85M

oist

sem

i-dec

iduo

usfo

rest

10–1

53

6.8

12.7

8.8

27.9

555

1045

6.3

19.6

5.6

6.0

Meg

hala

ya,N

.Ind

ia18

5015

ND

0–7

4013

.311

.2N

DN

D2.

52.

518

2017

50N

DN

D5.

17.

8M

ishr

aan

dM

onso

on10

ND

0–7

4012

.611

.9N

DN

D2.

42.

518

2017

50N

DN

D5.

37.

6R

am

akris

hnan

,198

3b

sem

i-dec

iduo

us5

ND

0–7

4011

.211

.2N

DN

D2.

32.

514

7014

00N

DN

D5.

57.

5

Per

u,Y

urim

agua

s21

0017

ND

0–10

4a

6.3

6.9

ND

ND

5.2

16.2

605

690

4164

4.0

4.5

Seu

bert

etal

.,19

77M

ois

teve

rgre

en

Pa

ra,B

razi

l20

88pr

imar

y29

20–

2.5

5N

DN

DN

DN

DN

DN

D36

829

0N

DN

DN

DN

DK

auffm

anet

al.,

1995

Moi

stev

erg

reen

2.5–

105

ND

ND

ND

ND

ND

ND

799

733

ND

ND

ND

ND

Pa

ra,B

razi

l20

88pr

imar

y43

50–

2.5

57.

06.

2N

DN

DN

DN

D62

866

5N

DN

DN

DN

DK

auffm

anet

al.,

1995

Moi

stev

erg

reen

+lo

ggin

g2.

5–10

520

.614

.2N

DN

DN

DN

D16

0316

03N

DN

DN

DN

D

Ron

doni

a,B

razi

l23

54pr

imar

y29

00–

2.5

510

.810

.331

53N

DN

D77

680

3N

DN

DN

DN

DK

auffm

anet

al.,

1995

Moi

stev

erg

reen

2.5–

105

18.8

13.5

8810

0N

DN

D14

7911

29N

DN

DN

DN

D

Ron

doni

a,B

razi

l23

54pr

imar

y36

10–

2.5

512

.3N

D43

ND

ND

ND

938

ND

ND

ND

ND

ND

Kau

ffman

etal

.,19

95M

oist

eve

rgre

en+

logg

ing

2.5–

105

17.1

ND

65N

DN

DN

D14

82N

DN

DN

DN

DN

D

Tur

rialb

a,C

osta

Ric

a27

008

520–

34

20.4

13.4

ND

ND

0.78

0.64

1511

1078

ND

ND

5.6

6.2

Ew

elet

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252

Figure 2. The relation between nutrient content of aboveground biomass and nutrient content of ash deposited on soil surface immediatelyfollowing slash burning at various tropical sites. Points are all taken from Table 1. The study of Ewel et al. (1981) was not included because Pand cation content of ash was higher than that of pre-burn slash biomass.

Young, 1989), reduce the availability of phytotoxicelements such as aluminum (Sanchez, 1976; Walker etal., 1986), and increase microbial activity and nutrientmineralization rates (Fritze et al., 1994; Khanna et al.,1994; Ohno and Erich, 1990).

Additions of ash often increase soil microbialactivity, because labile C in ash is added to soil or min-eralization of native soil C is stimulated by changes inpH and additions of nutrients. Khanna et al. (1994)found that ash additions of 25 Mg per ha increased Crespiration rates by 50–100% in acid soils with widelyranging organic matter contents. Increases were smal-ler in soils with low organic matter content, and atlower ash addition rates. In a Finnish forest soil,in situadditions of up to 5 Mg ash per ha did not significantlyincreasein vitro respiration rates of field moist soils 10d after ash application (Fritze et al., 1994). When soils

were brought to 60% of water holding capacity, respir-ation rates did increase by∼30%, but not at lower ashaddition rates. Raison and McGarity (1980) also foundthat ash additions increased soil respiration rates, butnoted respiration responses were much larger when incombination with soil heating.

Soil heating

Soil heating is controlled by a variety of factors thatinclude fuel quantity, quality, moisture and spatial dis-tribution on the soil surface (Martin, 1990; Raison,1979; Tomkins et al., 1991; Walker et al., 1986).Soil heating also will be influenced by soil texture,soil moisture and initial soil temperature (Dunn et al.,1979; Raison, 1979; Sanchez, 1976; Serrasolsas andKhanna, 1995a; Walker et al., 1986). Topography and

253

Figure 3. Soil temperatures attained at various depths during trop-ical slash burning in Dry (a), Monsoonal (b) and Humid (c) forests.Data are from the following studies: Mexico (Giardina et al., 2000);Brazil (Kauffman et al., 1993); Nam Phrom, Thailand (Kyuma etal., 1985); Ban Pa Pae, Thailand (Zinke et al., 1978); Ivory Coast(Van Reuler and Janssen, 1993); Costa Rica (Ewel et al., 1981); andNigeria (Lal and Cummings, 1979).

meteorological conditions preceding a burn, during aburn and following a burn influence soil heating bymodifying burn times and intensity, as well as the dir-ection and speed of fire spread (Martin, 1990; Raison,1979).

Soil heating during slash-and-burn appears to in-crease with increasing length of dry season (Figure3), a trend that can be at least partly attributed toincreasing soil moisture from dry to humid climates(Ewel et al., 1981; Giardina et al., 2000). Comparis-ons like these are complicated by the diverse methodsused to estimate soil temperatures (temperature sensit-ive paints or crayons, thermocouples). Some methodsmay over-estimate heat flux into soil (e.g. backing ma-terials for paint that conduct heat), and most requiresome disturbance of the soil prior to burning. Whilethese data should be interpreted cautiously, they doprovide a starting point for linking laboratory studies

to the field. This link is important because the surface2 cm of soil can contain from 300 to 1500 kg of Nand 30 to 200 kg of P (Table 2), a large portion ofwhich may be biologically active (organic matter, mi-crobes, roots) and sensitive to heating. Results frommost laboratory heating studies do not accurately pre-dict the effects of soil heating on plant growth becausepotted plants are often grown in uniformly heated soil(Giovannini et al., 1990; Kang and Sajjapongse, 1980)and plant roots are able to grow in the field belowheated soil. Furthermore, few studies have combinedsoil heating with additions of ash. However, laborat-ory studies provide insight into soil changes that arespecific to a given temperature.

The quantity of nutrients in surface soils can equalor exceed the quantity of nutrients in aboveground bio-mass, yet soils have received limited attention in fieldstudies of slash-and-burn. In the following sections,we examine results from studies that have examinedsoil heating in the field and in the laboratory.

Slash-and-burn effects on belowground biomass

Belowground biomass is comprised of soil organisms(bacteria, fungi, macro-fauna), mycorrhizae and plantroots. Microbial biomass can represent a large pool ofnutrients, especially on less fertile sites (Chapin et al.,1986), in disturbed forest stands (Vitousek and Mat-son, 1984, 1985) and in tropical dry forests (Campoet al., 1998; Singh et al., 1989). Soil microbial com-munities also control organic matter decompositionand nutrient mineralization rates and influence nutrientuptake by plants (Paul and Clark, 1996). Burning canhave profound effects on microbial biomass and thecapacity of soil microbes to carry out these functions(Garcia-Oliva et al., 1999; Raison, 1979; Serrasolsasand Khanna, 1995a; Walker et al., 1986).

Heating soil for 10 min at 70◦C kills some fungi,protozoa and bacteria, while temperatures above 127◦C almost completely sterilizes soil (Raison, 1979).Heating of a temperate forest soil (15% water content)to 300◦C to a depth of 1 cm and to 125◦C to a depthof 2 cm killed 98% of soil fungi and 75% of soil het-erotrophic bacteria in the top 2 cm of soil (Dunn etal., 1979). Soil heating to 120◦C in a range of Aus-tralian soils resulted in 34–80% declines in microbialbiomass, while heating to 250◦C resulted in bio-mass reductions of 85–99% (Serrasolsas and Khanna,1995a). For all sites reviewed in Figure 3, slash burn-ing resulted in soil temperatures to 1 cm that were highenough to cause very high rates of microbial mortality.

254

Soil temperatures were sufficiently high at all sites ex-cept the Costa Rican and Nigerian sites, to kill mostmicro-organisms down to 2 cm.

In lieu of data from slash-and-burn plots, estimatesof the quantities of nutrients that would be releasedfrom heat-killed micro-organisms during a burn canbe made using microbial biomass and nutrient contentdata from intact tropical forest. For a range of dryforest sites, dry season soil microbial N averaged from45 to 153µg N per g soil, while microbial P averagedfrom 11 to 34µg P per g soil in the top 10 cm of soil(Campo et al., 1998; Jaramillo et al., in prep.; Singhand Singh, 1995; Srivastava, 1992; Srivastava andSingh, 1988). Jenkinson and Ladd (1981) estimatedmicrobial biomass of an intact Nigerian moist forestsite to contain 90µg N per g soil and 25µg P perg soil. With data from Srivastava (1992) (dry seasonmicrobial nutrient content of 88µg N per g soil and 34µg P per g soil), and the moist forest data of Jenkinsonand Ladd (1981), we illustrate the potential effects ofsoil heating on microbial biomass at these sites. Afterconverting these concentrations to an area basis (withan assumed bulk density of 1 g per cm3), then mul-tiplying by 0.9 to approximate 90% burn mortality,soil heating in the top 2 cm would cause the release of16 kg N per ha and 6 kg P per ha for the dry forest, and16 kg N per ha and 4 kg P per ha for the moist forest.These quantities of N and P are significant relative tothose contained in ash (Table 1).

Microbial biomass of a slashed but not yet burnedforest may be 2–3 times higher than for undisturbedforest because of higher soil moisture from reducedevapo-transpiration and the mulching effect of felledvegetation, reduced competition with plants and in-creased availability of dead roots and litter (Paul andClark, 1996). Additionally, when a site is slashed,microclimate of soil generally is much warmer thanin the previous forest. Therefore, the above estimatesof microbial release are likely conservative for dry,monsoonal and some moist forest sites (Figure 3). Inline with this suggestion, Jaramillo et al. (in prep) ob-served a 90% reduction in microbial biomass in 0–10cm depth soils following slash burning that released∼140 kg N per ha and∼30 kg P per ha.

Microbial recovery from burning is hypothesizedto be rapid and substantial (Nye and Greenland, 1960),because of elevated soil pH, moisture and temperature,increased nutrient supply and substrate availability,and reduced competition with plants. In a tropical hu-mid forest, the dynamics of microbial activity appearto fit this model (Matson et al., 1987; Nye and Green-

land, 1960). In contrast, slash burning in Mexicandry forest resulted in a dramatic reduction in micro-bial biomass that persisted in the top 10 cm of soilfor up to 2 years (Jaramillo et al., in prep.). Nitro-gen mineralization rates following burning at this siteshowed a similarly long recovery time (Døckersmithet al., 1999). In a temperate study of dry woodland,microbial biomass N was reduced for 90 days fol-lowing heating (the length of the incubation), withlarger reductions for initially drier soils (Klopatek etal., 1990). In another temperate study, reductions insoil microbial biomass were inversely related to soilheating, itself related to soil moisture at the time ofburning (Dunn et al., 1979), which suggests that mi-crobial recovery is increasingly delayed with increas-ing severity of soil heating. Serrasolsas and Khanna(1995a) found a mixture of responses and persistenceof these responses depending on degree of heating,soils type, clay content and the population of organ-isms in question. In the field, microbial recovery fromslash-and-burn will also be influenced by the dramaticchanges in soil micro-climate that accompany land-use. Without the moderating influence of the forestcanopy, soils will be warmer during the day and coolerat night. Additionally, soils will be wetter in clearedsites (Paul and Clark 1996), particularly at the start ofthe rainy season because canopy interception is elim-inated. Overall, assessing the long-term response ofsoil micro-organisms to slash-and-burn represents aresearch priority.

Coarse and fine roots in surface soil are killed andsometimes consumed during slash burning. For ex-ample, in a Mexican dry forest, slash burning reducedlive and dead fine roots (<1 mm in diameter) in theupper 2 cm of the soil by 55% and 32%, respectively(Castellanos, 1998). Notably, on an area basis fineroots contain low amounts of nutrients compared tothe other nutrient fluxes reviewed here (Kummerow etal., 1990; Lessa et al., 1996), especially at the endof a dry season when burning typically occurs andwhen fine root biomass is lowest (Castellanos, 1998;Kavanagh and Kellman, 1992; Roy and Singh, 1995).

Slash-and-burn effects on soil organic matter

Soil organic matter is comprised of the decay resist-ant remains and the precipitated by-products of plantresidue decomposition, and is affected by slash burn-ing on various levels. Thermally induced losses oftotal soil C, the focus of most slash-and-burn stud-ies, are typically small (Table 1). Larger losses may

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be incurred in surface soil when large quantities offuel are consumed. Soil heating can also alter thechemical composition of organic matter (Fernandezet al., 1997), either through soil desiccation, thermalmodification or release of microbial biomass. Finally,soil heating can thermally mineralize nutrients with orwithout loss of total soil C (Giovannini et al., 1990;Giardina et al., 2000). Because the effects of soil heat-ing are often element specific, we address individualnutrients separately below.

Phosphorus

The processes regulating the geochemical release ofinorganic P and the biochemical mineralization oforganic P are affected by soil pH, vegetation and mi-crobiology (Cross and Schlesinger, 1995; Giardina etal., 1995; Hedley et al., 1982; Stewart and Tiessen,1987), all of which are significantly, but not uniformly,affected by fire. Studies of slash-and-burn agriculturehave assessed changes in plant-available P by usingextracts that remove pools of soil P that correlate withplant uptake of P in controlled agronomic studies. Theutility of these methods in studies of slash-and-burnis unclear because the longer-term supply of P in soilis controlled by the replenishment of depleted solu-tion P pools and the turnover of organic P, not thesize of a specific extractable pool (Tiessen and Moir,1993). Because the capacity of an extract to removeP varies with the chemical properties of a soil, thiscapacity will also be influenced by fire. For example,the slightly basic, sodium bicarbonate solution oftenused to estimate plant-available P in soil (Olsen et al.,1954) functions by increasing soil pH and providingan overwhelming supply of anions that compete with Pfor adsorption sites (Tiessen and Moir, 1993). Burningalso causes an increase in soil pH and an increase inthe concentration of anions that compete with P foradsorption sites.

Plant-available inorganic P, as determined by pHdependent extracts, generally increases when soil isheated (Tables 3 and 4). Between 170 and 300◦C,these increases are matched by declines in organic Pas P is thermally mineralized from soil organic matter,often with little loss of total soil C (Giardina et al.,2000; Giovannini et al., 1990; Kang and Sajjapongse,1980). The heat-induced death of soil microbial pop-ulations is a likely source for part of the increase inextractable soil P (DeBano and Klopatek, 1988; Serra-solsas and Khanna, 1995b). Between 220◦C and 500◦C, P is liberated during the thermal oxidation of soil C

Figure 4. An alternative conceptual model for the fluxes of nutri-ents during slash-and-burn clearing of tropical forest (Modified fromWalker et al., 1986). Variable quantities of biomass are consumedand converted to ash, which may be lost from a site by wind orwater erosion, or incorporated into soil. Substantial quantities ofC, N and P are volatilized, and C, N, P and cations may be lostvia convective transport during the burn. Soils are influenced byburning; heating is variable and dependent upon the many factorsthat control heat penetration into soil. Soil heating causes the re-lease of nutrients from non-plant-available sources (roots, microbialbiomass, organic matter) to plant-available forms. Heating and ashalso cause an increase in soil pH, which can solubilize previouslynon-plant-available nutrients.

(Table 4; Andriesse and Koopmans, 1984; Giovanniniet al., 1990; Kang and Sajjapongse, 1980).

Alternatively, heating in heavily weathered acidsoils can reduce P availability by increasing the P ad-sorption capacity of soil (Sibanda and Young, 1989);heating soil to 200◦C destroyed carboxyl groups onorganic matter that compete with P for adsorption siteson Fe and Al minerals. Organic matter reductionsof <15% can correspond to large (>50%) losses ofcarboxyl functional groups, and increased P adsorp-tion capacity may largely offset any increased supplyof P from mineralized organic matter. In contrast,Romanya et al. (1994) found that despite higher P ad-sorption capacity in intensively heated soils, seedlinggrowth and P content in these soils was higher thanin either unburnt or lightly burnt soils. Seven monthsafter burning, Bray-1 extractable P was significantlyhigher in burnt soils, with the largest increases foundfor the most intensely burnt soils. These increasesin Bray-1 extractable P corresponded to significantreductions in organic P.

Burning can profoundly affect phosphatase activ-ity, the primary exo-enzymes responsible for miner-alizing P from soil organic matter. Serrasolsas andKhanna (1995b) reported a steady decline in phos-

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Table 3. Effects of laboratory heating on soil C and N. Values for sand, clay, C and N are % of soil mass

Soil Sand Clay Total C Total N NH4+NO3 Referencetype (%) (%) (%) (%) (mg/kg soil)

Australia a20 60 120 250 20 60 120 250 20 60 120 250 Serrasolsasand Khanna,1995a,b

Yellow podzolic 73 8 1.7 1.7 1.8 1.7 0.06 0.05 0.05 0.05 10 11 13 39Leached sand 98 0 6.9 6.6 6.8 ND 0.22 0.12 0.12 ND 19 19 39 35Yellow podzolic 72 11 3.2 3 2.9 3.4 0.08 0.08 0.07 0.07 8 9 23 38Yellow earth 65 12 6.2 6.3 6.4 6.3 0.12 0.12 0.14 0.17 20 21 59 51Red earth 34 26 10.6 10.4 10.5 10.9 0.36 0.37 0.37 0.39 39 40 122 60

Italy a25 170 220 460 700 25 170 220 460 700 25 170 220 460 700Giovannini etal., 1990

Arnino Silty Clay 2.4 2.3 2.1 0.3 0.0 0.13 0.122 0.11 0.008 0.005 13 22 54 8 7Torretta Sandy Loam 1.1 1.0 0.9 0.1 0.0 0.099 0.078 0.064 0.005 0.004 12 20 42 8 7

Nigeria a25 100 200 500 600 25 100 200 500 600 25 100 200 500 600Kang andSajjapongse, 1980

Paleustalf 63 21 1.75 1.7 0.8 0.1 0.05 0.24 0.24 0.21 0.03 0.01 ND ND ND ND ND

USA, India& Ethiopia a25 100 200 400 600 25 100 200 400 600 25 100 200 400 600Sertsu and

Sanchez, 1978Ethiopian 12 50 3.4 3.9 2.5 0 0 0.85 1.19 1.2 0.6 0.21 1 2 75 1 0Paleudult 15 32 2.9 2.4 1.8 0 0 1.1 1.1 1.1 0.65 0.6 1 1 12 1 0Vertisol 25 55 2 2 1.9 0 0 0.95 0.8 0.94 0.59 0.61 2 2 14 1 0

a Incubation temperature (◦C).

phatase activity with increasing temperature, and anear complete loss of phosphatase activity in soilsheated to 250◦C. Burning on dry pine soils re-duced phosphatase activity to 30% of unburnt controls(DeBano and Klopatek, 1988). This loss of activitypersisted for the 90 day duration of the experiment.Losses of phosphatase activity were also observed inburnt juniper soils (dry or wet), but declines in controlsoil phosphatase activity complicated interpretations.Phosphatase activity in burnt, wet juniper soil re-covered after 45 d and surpassed control soil activityby day 90.

Few field studies of tropical slash-and-burnprovide enough information to assess changes in soil Pavailability (Table 2). One month after slash burning ata site in the Peruvian Amazon and several weeks intothe growing season, Seubert et al. (1977) measuredincreases in bicarbonate extractable P (Olsen P) of 11kg per ha. These authors found that ash contained 6kg P per ha immediately after burning and suggestedthat ash contributed 6 of the 11 kg P per ha increase.The size of the increase in Olsen P was likely under-estimated for several reasons. Soil pH increased afterburning, potentially reducing the capacity of the ex-tract to remove P. Only a small portion of P in ash

appears to be plant-available (Khanna et al., 1994;Ohno and Erich, 1990). Furthermore, in the monthbefore sampling, post-burn soils were moist, and bio-logical and geochemical fixation (especially importantin the heavily weathered, acidic soils of this Peruvianstudy) would have reduced the elevated quantities oflabile soil P.

At a Mexican dry forest site, soil heating dur-ing slash burning transformed 35 kg of non-plant-available organic and occluded soil P per ha intoplant-available mineral forms (Giardina et al., 2000).About 7 kg per ha of volatilized P were transferredduring burning from biomass to soil (Table 2). Ashfrom consumed slash contained 11 kg P per ha im-mediately after burning (Table 1), 6 kg of which weretransported off the site by wind. Because soils at thisdry forest site were near neutral before burning, ash in-duced increases in soil pH probably played little part inincreasing soil P availability (Giardina et al., 2000). Incomparing inputs to soil, P from ash was of secondaryimportance compared with immediate changes withinthe soil. The longer-term supply of mineralized P wasnot examined.

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Table 4. Effects of laboratory heating on soil phosphorus and pH. Available P in the Australian and Nigerian soils were determined usingBray-1 extract (Bray and Kurtz, 1945), and by Olsen extract (Olsen et al., 1954) in the others

Soil Total P Available P Soil pH Referencetype (mg/kg soil) (mg/kg soil)

Australia a20 60 120 250 20 60 120 250 20 60 120 250 Serrasolsasand Khanna,1995a,b

Yellow podzolic 252 ND ND ND 23 21 22 42 ND ND ND NDLeached sand 45 ND ND ND 1 0.5 1 4 3.6 3.5 3.4 3.1Yellow podzolic 68 ND ND ND 0.5 0.5 1.5 4 4.2 4.7 4.6 4.3Yellow earth 94 ND ND ND 0.5 0.5 0.7 3 4.0 4.7 4.6 4.1Red earth 458 ND ND ND 3.5 3.5 7 8.5 ND ND ND ND

Italy a25 170 220 460 700 25 170 220 460 700 25 170 220 460 700Giovannini etal., 1990

Arnino 650 650 650 650 650 5 23 42 92 35 7.6 7.3 7.0 7.6 10.7Torretta 700 700 700 700 700 8 37 55 100 40 7.8 7.3 7.2 7.9 11

Nigeria a25 100 200 500 600 25 100 200 500 600 25 100 200 500 600Kang andSajjapongse, 1980

Paleustalf 210 ND 206 ND 259 7.2 9.6 40 20 18 6.1 6.1 6.2 6.9 6.8

USA, India& Ethiopia a25 100 200 400 600 25 100 200 400 600 25 100 200 400 600Sertsu and

Sanchez, 1978Ethiopian ND ND ND ND ND 2 4 33 59 26 5.9 5.2 5 6.5 5.4Paleudult ND ND ND ND ND 10 13 49 48 56 5.1 4.9 5.1 6 5.3Vertisol ND ND ND ND ND 3 2 10 20 19 8.2 8 7 6.9 7.7

a Incubation temperature (◦C).

Nitrogen

Soil N is very sensitive to biological transformations,and to losses due to leaching, volatilization, oxida-tion and denitrification (Matson et al., 1987; Raison,1979). Volatilization of ammonia (NH3) and nitricacid (HNO3) increase with temperature. At temperat-ures above 300◦C, soil organic N is lost during thethermal oxidation of organic matter in the form of ox-idized N gases and N2 (Raison, 1979). When soil isheated above 100◦C, ammonium levels also generallyincrease (Table 3) as NH3 is released from microbialbiomass (DeBano and Klopatek, 1988; Serrasolsasand Khanna, 1995a), thermal decomposition organicmatter and protein hydrolysis (Russell et al., 1974)and the desiccation of soil minerals (Raison, 1979).Most slash-and-burn studies examining soil N have fo-cused on changes in total N (Table 2), which may notcorrespond to short-term changes in N availability. Intropical and temperate studies where N mineralizationhas been examined, the influence of slash burning onN mineralization rates appears variable, with studiesreporting increases (Matson et al., 1987), decreases(Døckersmith et al., 1999) or no significant change(Bauhus et al., 1993).

At a wet forest site in Costa Rica, slash-and-burnclearing of a 75-yr-old stand resulted in a 7–12 foldincrease in extractable NH+4 and NO−3 , and a 3 fold in-crease net N mineralization rates (Matson et al., 1987).Large differences between burnt and unburned controlplots persisted for up to 6 months. At a dry forestsite in Mexico, slash-and-burn clearing of a 100 yrold stand resulted in a decrease of 150 kg non-plantavailable N; 68 kg per ha was lost from soil and theremaining 82 kg per ha was transformed into mineralform (Døckersmith et al., 1999). Net N mineralizationrates in 0–10 cm depth soils decreased dramaticallyfollowing burning and continued to decline throughthe first growing season.

CationsCation availability and soil cation exchange capacitycan be affected by soil heating, but the influences arevariable. At low temperatures, cation availability hasbeen shown to increase, decrease or remain unchangedin response to heating. Sertsu and Sanchez (1978)found that both exchangeable Mg and Ca declinedsteadily while exchangeable K increased steadily withheating to 600◦C; effective cation exchange capa-

258

city declined steadily with heating. In contrast, Kangand Sajjapongse (1980) found a steady increase in ex-changeable cations and cation exchange capacity withincreased temperature to 200◦C, after which the avail-ability of these cations and cation exchange capacitydropped dramatically. In the field, slash burning usu-ally results in an increase in soil cation availability. Itis generally assumed that increases in an exchangeablecation represents the input of that cation from ash. In-terpreting these results is complicated, however, by theinfluence that ash, heating and changing pH can haveon cation availability in soil.

Soil pHThe influence of heating on soil pH are not consist-ent across studies, temperatures or soil type (Table 4).Several laboratory heating studies have documentedincreased soil pH after heating alone (Andriesse andKoopmans, 1984; Kang and Sajjapongse, 1980; Khareet al., 1982; Kutiel and Shaviv, 1989), with changesoccurring at temperatures as low as 200◦C. Con-versely, other studies have documented that soil heat-ing to 200◦C (Sertsu and Sanchez, 1978) or to 220◦C (Giovannini et al., 1990) lowers soil pH, and it isnot until soils are heated above 400◦C that soil pHincreases over that of unheated soils. Lowered soil pHhas been attributed to reduced buffering capacity ofsoils, newly exposed soil surfaces, and released Alduring heating. Elevated soil pH has been attributedto consumption of organic acids.

Conclusion

The biophysical sustainability of shifting cultivationhas been the focus of several recent reviews (Brady,1996; Harwood, 1996; Kleinman et al., 1996; Tinkeret al., 1996); however, assessing the biophysical sus-tainability of shifting cultivation is difficult because ofthe long time periods involved and the wide array ofsoils, climates and population pressures under whichit is practiced, and because adequate data are lacking(Juo and Manu, 1996). Biomass burning is known tohave significant long-term consequences for the nutri-ent balance of forest ecosystems (Raison et al., 1993),and Walker et al. (1986) have identified three mainsources of nutrients that supply post-burn increases insoil fertility. These include soil organic matter, soilmicrobial biomass and aboveground biomass (Figure4). The extent to which slash-and-burn alters these nu-trient sources, and the persistence of these alterations

is poorly characterized for most shifting cultivationpractices (Table 2). In perspective, the following com-ponents would be useful to address in a single study:(i) the quantity of nutrients contained in slash biomass;(ii) the nutrient content of ash and unburnt debris; (iii)the quantity of nutrients in ash that are lost from a site;(iv) the net quantity of non-plant-available soil nutri-ents that are thermally transformed into mineral formsavailable to plants (i.e. changes in both exchangeableand total soil nutrient pools); (v) the extent to whichpost-burn increases in soil pH modify nutrient avail-ability (e.g. P sorption capacity); and (vi) the impactof burning on microbial biomass and activity (e.g. Cand N mineralization rates). Ecosystem level studiesaddressing these questions are needed across a rangeof forest types and field conditions before the biophys-ical sustainability of slash-and-burn can be accuratelyassessed.

Acknowledgements

We thank D. Binkley, J. Fownes, R. Menezes and twoanonymous reviewers for ideas, comments and sug-gestions on this manuscript. We thank B. Kauffman,F. Garcia-Oliva and C. Rhoades for helpful discussionson fire and nutrient cycling.

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