Successional Patterns Associated with Slash-and-Burn Agriculture in the Upper Rio Negro

Region of the Amazon Basin

Author(s): Christopher Uhl, Howard Clark, Kathleen Clark and Pedro Maquirino
Source: Biotropica, Vol. 14, No. 4 (Dec., 1982), pp. 249-254
Published by: The Association for Tropical Biology and Conservation
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Successional PatternsAssociated withSlash-and-BurnAgriculturein the

Upper Rio Negro Region of the Amazon Basin
Christopher Uhl,' Howard Clark, and Kathleen Clark
Instituteof Ecology, Universityof Georgia, Athens, Georgia 30602, U.S.A.
and
Pedro Maquirino
San Carlos de Rio Negro, TerritorioFederal de Amazonas, Venezuela

ABSTRACT
In a studyof plant successionduring and followingslash-and-burn
agriculturenear the town of San Carlos de Rio Negro
in southernVenezuela we foundthat: 1. Weeds establishedjust as readily in plots with Manihot esculenta (the principal
crop plant of the region) presentas in plots with M. esculen:a removed. 2. Repeated farm-plotweedings caused woody
plants to decline in numbersand biomass and herbaceousplants to increase. 3. Forbs and grasses dominated immediately
followingfarmsite abandonment,but by one year these had begun to senesce and fast-growing
successionalwoody species
(particularlyVismia spp.) were common. Standing crop biomass at one year was 773 grams dry weight m-2. 4. Several
microhabitattypeswere presenton abandoned farmsites. Grasses and forbs showed no microhabitatpreference,whereas
successionalwoody individualshad their best establishmentnear slash and under fruittrees.

RESUMEN
En un estudio de sucesion,realizado durantey luego del abandono de una parcela de agriculturade cortey quema, en los
alrededoresde la problacion de San Carlos de Rio Negro (Venezuela), encontramosque: 1. Las malezas se establecieron
con la misma rapidez en las parcelas de Manihot esculenta (que es el principal cultivo en la region), que en las parcelas
donde se habian eliminado las plantas de M. esculenta.2. El desmalezado repetidode las parcelas cultivadasprodujo una
disminucionen el nuimeroy biomasa de plantas leiiosas, y un aumentoen las plantasherbaceas.3. Inmediatamentedespues
de ser abandonada la parcela cultivada fue dominada por plantas herbaceas,pero al anio siguienteestas ya habian comenzado a senescer,y ya eran comunes las especies leniosasde crecimientorapido (particulamenteVismia spp.). La biomasa
de esta parcela era de 773 gramos de peso seco in-2 al primer anio. 4. En las parcelas abandonadas despues de ser cultivadas se encontrabanmuchos tipos diferentesde microhabitats.Las plantas herbaceas no mostrabanninguna preferencia
por determinadomicrohabitat,mientrasque los individuosde especies leniosasse establecian mejor cerca de los troncos
cortados y debajo de arboles frutales.

fluenceof the Guainia and Casiquiarerivers (10

is thepredominant
farmingmethodon roughly30 percentof the ex- 56' N, 670 03' W, 119 m elevation).A species-rich
ploitablesoils of the worldand supportsover 250 evergreenforestwith a canopyheightof 25-35 m
millionpeople (Hauck 1974). In theAmazonBasin was initiallypresenton thisplot.This forestwas cut
an estimated36,000km2of non-flooded
forestwere and burnedin late 1976. The plot was thenfarmed
practices(Uhl and
clearedby slash-and-burn
agriculturists
from1966 to forthreeyearsusingtraditional
1975, makingthisthe mostcommonformof land Murphy1981) and thenabandonedto naturalsucuse in thisregion (Meyers1979). This paper de- cession.A 30 x 50 m plot in thecenterof the farm
scribessuccession
duringtheperiodof farm-plot
use plot was designatedas the studyplot.
and for the firstyear followingabandonmentin
aseasonal
The climateof the regionis relatively
the upperRio Negro region.
of 26?C and mean
witha meanannualtemperature
annualrainfallof 3500 mm.The monthsfromOctorainber throughMarchare less rainy,but monthly
THE STUDY SITE
exceeds100 mm.
The studywas conductedon a 0.25 ha tierrafirme fallstillgenerally
(i.e., non-flooded)plotlocated4 km eastof thevilThe terrainconsistsof rollinghills,with only
lage of San Carlosde Rio Negro,close to the con- thehighertierrafirmeareasbeingused forfarming.
IContributionof the InternationalAmazon Project coordin- Soils on thehigherareascontainan A horizoncomated by the Ecology Center of the InstitutoVenezolano de posed mostlyof fine sand that overlieskaolinite
InvestigacionesCientificas(IVIC), Caracas,Venezuela. This clayand abundant
concretions
(Herrera1979).
ferric
work was funded by the U.S. National Science Foundation
becauseof inThese soils are verylow in nutrients
in cooperationwith IVIC.
SLASH-AND-BURN AGRICULTURE

BIOTROPICA 14(4): 249-254

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1982

249

tensiveleachingunderhumidtropicalconditions
for two to threeyears.Farmersmay continueto visit
millionsof yearsand the absenceof unweathered theirplots for severaladditionalyearsto harvest
parentmaterialwhichcould serveas a freshsource fruitsor an occasionalpatchof matureyuca.Howof nutrients
(Fittkauet al. 1975).
ever,becauselittleor no weedingoccurs,such sites
wereregardedas abandonedforthepurposesof this
study.

METHODS

We recordedthe heightand identityof all in-

PRE-ABANDONMENT
SUCCESSION.-Farm plotsinthe dividuals?: 5 cm tall in the permanent
1 x 1.5 m

San Carlosregionare plantedprimarily

to Manihot plots at 4, 10, and 16 monthsfollowingthe final
esulentaCrantz.(knownlocallyas yuca), withpine- farm-site
weeding.
apple plantsand fruittreespresentat low densities.
We studiedtheeffectof yucaon weedestablishment Biomassof all plants< 2 m tall was estimated
by harvesting
all plants
thenumberof weedsunderyucaplants one yearafterabandonment
bycomparing
present
(including
pineapple
plants)
in
25
plotsof
with the numberpresentin plots whereyuca had
1
M2. Harvestedplantswereseparatedby life form
been artificially
removed.All weeds were removed
and weighed.The biofrombothtreatments
at thebeginning
of theexperi- and plant part,oven-dried,
mass
all
(in
grams)
of
plants
> 2 m tall in the
ment.Treatments
werepaired,2 x 2 m in size,and
study
plot
was
estimated
by
the
regressionof direplicated
eighttimes.A 2 m buffer
stripwas cleared
ameter2
X
height
(both
in
cm)
on stemweight
aroundthenon-yuca
plots.
[y = (0.49x) -29.5; n= 111 and leafweight[ySan Carlos farmplots are usuallyweeded once (0.17x)
+76.4; n 111. Diametermeasurements
in thefirstyearand twotimesa yearthereafter
until weremadeat 10 cmheight.Valuesof r2were> 0.90
plot abandonment.
All weeds are pulledout of the in bothcases.Rootbiomasswas estimated
byseparatsoil exceptforsproutsor an occasionallargeindivid- ing all rootsfromsoil to a depthof 50 cm in five
ual whichmustbe cut.The effectof repeatedfarm- 0.25 m2randomly
locatedplots.
plot weedingson density,biomass,and life-form
At the timeof farm-site
abandonment,
thereare
composition
of recolonizing
vegetationwas studied
several
distinct
Much
of the
microhabitats
present.
by pullingall non-cropplants from27 randomly
area
contains
decombare
soil,
but
large,
partially
located1 x 1.5 m permanent
plotsat 10, 16, 21, 26,
and 31 months.While thismimickedlocal practices, posedlogsstillcoverthesoil in someplaces.Clumps
reproducing
pineappleplantsand isoit resultedin an underestimate
of rootstandingcrop of vegetatively
lated
fruit
m
are also present.
Finaltrees
(4-10
tall)
becausea portionof the rootswas leftin the soil.
The harvested
plantsin each plot weregroupedas ly,soil is moundedand plantedto yuca in patches
forbs,grasses,successional
woodystems,foresttrees, whereit appearsthata finalcrop is possible.The
typeson the establishand vines.Plantsin each groupwere dividedinto effectof thesemicrohabitat
ment
of
successional
vegetation
followingfarm-site
root,stem,andleaffractions,
oven-dried
andweighed.
in a 25 x 25 plot locatedin
was
tested
abandonment
The effectof farming
on thesoil seed bankwas
a recently
abandonedfarmsite locatedabout 150 m
studiedby comparing
the numberand speciescomfrom
main
the
studysite.The plotwas clearedof all
positionof seedsgerminating
in soil samplestaken
and
slashexceptforfivecashewtrees.A
vegetation
fromthe farmsiteat thetimeof abandonment
and
x
m
was
formedaroundeach cashewtree.
3
3
plot
froman adjacentundisturbed
forestplot. Each soil
The
treatments
wererandomly
four
remaining
placed
samplewas 20 x 20 x 5 cm deep. All sampleswere
m
2
in
x
with
m
3
border
between
3
plots
strips
plots.
spreadin traysand set 1.7 m abovethegroundin a
The bare soil plotswere simplymarkedoff.Pinesunnylocationon large tables. Contamination
by
were transplanted
into theirassigned
wind- or animal-borne
seeds was checkedthrough apple plants
Soil
in
the
was
mounded
plots.
yuca
plotsand yuca
of fiveadditionaltrays,
surveys
containing
soil which
stem
stocks
inserted
into
the
mounds.
Large
logswere
had beenoven-dried
at 250? C forfivehours,placed
in the slash plots.All herbaceousand woody
amongthesampletraysunderstudy.Traysweresur- piled
in each of these
plantsthathad becomeestablished
veyedfor new germinations
two timesa week for
counted
and
identified
were
after
one year.Sixplots
two months,afterwhichtimethe soil in each tray
teen
soil
cores
cm
diameter
x
cm
(6
5
deep) were
was turnedoverand mixed.Surveyscontinueduntil
from
each
the
and
taken
of
bare
fruittree
soil,
slash,
no further
germinations
werefound.
of theexperiment.
The cores
plotsat the conclusion
POST-ABANDONMENT SUCCESSION.-Slash-and-burn fromeach plotwerepooledand set in a sunnyspot
farmsitesin theSan Carlosareaare actively
of soilseedsas describedabove.
used for to allowgermination
250

Uhl, Clark, Clark, and Maquirino

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RESULTS AND DISCUSSION

actualpresence of yuca plantshad no apparenteffecton the

weedafterfarm-plot
abilityof weedsto reestablish
ing,althoughthe yuca canopydid reducelightand
soil-surface
temperature(table 1). More than 80
in bothtreatment
percentof all establishments
types
were the forb species,Borreria latifolia (Aubl.)
Schum.and Eupatoriumcerasifolium(Sch. Bip.)
Baker.Grassescomprised
an additional10 percentor
in both treatments
with
moreof all establishments
PaspalumdecumbensSw. the mostcommonspecies.
The processof succession
beganand was curtailed
throughweedingfivetimesbeforethefarmsitewas
finallyabandoned.These repeatedweedingshad no
cleareffecton the numberof reestablishing
plants
(table 2). Plantdensitywas low at the first,
second,
and fourthweedingsand relativelyhigh afterthe
thirdand fifthweedings.Weedingsthreeand five
occurredtowardthe end of the wet season,while
weedingstwo and fouroccurredat the end of the
PRE-ABANDONMENT SUCCESSION.-The

TABLE 1. A summaryof differencesbetween plots with

yuca presentand adjacent plots where yuca was
removed.
Yuca present Yuca removed
7
Number of species present
9
Number of woody
establishmentsa
0.7 + 0.8b
0.0
Number of herbaceous
establishments
15.6? 8.53
12.3?+9.6
Total numberof
establishments
16.3 ? 8.42
12.3? 9.6
Midday light at soil surface
on overcastday (lux)
7,500?+9,500 22,750?+9,700
Midday temperatureat soil
surfaceon sunnyday
46.8?4.4
(0C)
40.5?0.9
Soil percentagemoisture
after three days
withoutrain
6.4?0.93
6.9?+1.16
aOnly plants ?! 5 cm tall were consideredas established.
bNumberof plants presentper 2 x 2 m plot ? 95 % confidenceinterval.

relatively
dryseason.Uhl et al. (1981) have shown
thatreducedrainfallat San Carlos lowersthe rate
in
and seedlingestablishment
of seed germination
disturbedsites.
The repeatedweedingsdid appearto affectthe
vegetalife-form
compositionof the reestablishing
densityat the firstweedtion (table 2). Forest-tree
plot-' (all of sproutorigin),
ingwas 0.65 individuals
had declinedto 0.11.
butbythefinalweedingdensity
Hence, the repeatedweedingshad apparentlyexhaustedthe sproutingreservesof manycut forest
trees.Successionalwoodyspeciescomprised64 perplot-') of thetotalstems
cent (i.e., 12.1 individuals
presentat the firstweeding,but by the fifthweedaccounting
for
ing theyalso had declinedin numbers
only 2 percentof the total individualspresent.In
contrast,
thedensityof forbsand grasseswas low at
thefirsttwoweedings,
(i.e.,more
buthighthereafter
than25 individuals
plot-' forweedings3, 4, and 5).
Weed biomass measurementsparalleled density
values. Havel (1960) in New Guinea, Snedaker
(1970) in Guatemala,and Kellman (1970) in the
Philippinesreportedsimilartrends.
on the abanHigh forband grassestablishment
froman abundance
donedfarmplotmayhaveresulted
surfacesthere.In the San
of favorablegermination
Carlos region,root mat (a surfacemat of forest
treeroots5-30 cm thick),charredwood,and bare
soil surfacesare the main seed-bedtypespresent
followingforestcuttingand burning.The bare soilsurfacetype increaseswith time because the root
mat decomposesand the charredwood is washed
awayin heavyrains.For example,21 percentof the
farm-site
surfacearea was bare soil one year after
69
forestclearing,but by the timeof abandonment
percento'fthissitehad exposedsoil. In separatesurfacecolonization
experiments
(Uhl et al. 1981), forb
on these
and grassspeciesshowedgood establishment
bare-soilsurfacesas comparedto successional
woody
species.

The resultsof the seed-banksurvey(table 3)

TABLE 2. The mean numberof plants and grams dry-weightbiomass in 27 1 x 1.5 m permanentplots at each of five
weedings.a
Forbs and grasses
Weeding

Density

Biomass

1
2
3
4
5

6.1? 2.7b
9.7? 5.6
72.6?30.8
28.2+16.2
53.1?22.6

7.1? 5.4
55.2?24.9
41.7?16.3
24.2?12.5
27.5?16.7

Total individuals

Woody plants
Density
12.7?6.0
2.2?0.9
2.2?1.2
1.8?0.8

Biomass

Density

Biomass

16.6+6.5
8.5?4.7
3.4.2.3
5.0+4.1
3.2?2.5

18.8? 7.4
11.9? 5.7
76.0?30.8
30.4?16.2
54.9?22.7

23.7? 9.1
63.7?24.9
45.1?16.7
29.2?13.0
30.7?16.6

aIndividuals < 3 cm tall not consideredin densityestimates.

"95 % confidenceinterval.
Slash-and-Burn Agriculture in Amazon Basin

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251

also help to explainthe observedshiftfromwoody ed out beforetheycan produceseed locally,and

to herbaceous
life-form
dominanceduringtheperiod afterseveralweedingsthe seed bank thatgives rise
of farm-plot
use. The estimatednumberof germin- to thisgroupis greatly
reduced.
able seedsm-2 was 581 in the farmsiteat the time
POST-ABANDONMENTSUCCESSION.-Four months
of abandonment
and 177 in theadjacent,undisturbed
afterthe last farm-plot
weeding,grassesand forbs
forestsite. Anotherforestsite, severalkilometers
?? 5 cm
accountedfor93 percentof theindividuals
away,was alsosampledat thistimeandhad a germinin
tall
the
27
1
m
permanent
X
1.5
study
plots
(table
able seed bankof 752 seedsm-2 (Uhl et al. 1981)
The
4).
most
abundant
at
this
time
were
forb
species
suggesting
thatseed-banksize varieswith location.
Eupatorium
cerasifolium
and
Phyllanthus
sp.
with
3.2
Successionalwoody species dominatedthe germinand
4.1
individuals
plot-',
most
respectively.
The
able seed bank in both the forestsites; however,
mostof the identified
germinations
in the farmsite commongrassspecies in the plots were Paspalum
were forbsor grasses.Forb and grassseeds increase decumbensand PanicumpilosumSw. Total plant
densitydeclinedat the 10- and 16-monthsurveys
in abundancein farm-site
seed banksbecausetheir
because
manygrassesand forbshad died.Mostdeath
lifecyclesareshorter
thantheinterval
betweetweedresulted
eitherfromshadingby tallerplantsor from
ings.In contrast,
successional
woodyspeciesareweedsenescencefollowingfruiting.
All otherlife forms
either
increased
or
remained
the
same betweenthe
TABLE 3. The numberof germinableseeds in soil samples
surveys.Foresttreeshad very
taken fromthe farm plot at the time of aban- four-and 16-month
donment and from an adjacent mature forest low densitiesthroughout
the studyperiodand were
plot.
onlypresentas sprouts.
At 16 months,successionalwoodyplantsdomiFarmed site
Forestsite
nated
the site withthe tallestindividuals3-4 m in
Number of samples
15
13
height.Of 199 individuals- 2 m tallpresentin the
Mean numberof
germinationsper sample
23.3 12.88a
7.08?2.29
30 X 50 m studyplotat 16 months,
61 percentwere
Number of germinationsm-2
581
177
Vismia lauriformis
(Lam.) Choisy, 19 percent
Percentageof total germinations
growingto an identifiablestage 74
Foresttrees,percentageof total
identifiedgerminations
0
Successionaltrees,percentage
of total
identifiedgerminations
8
Forbs and grasses,percentage
of total
identifiedgerminations
92
'95% confidenceinterval.

69
1
95
1

Vismiajapurensis
Reich, and 9 percentwere Cecropia
ficifolia
Snethlage.An additionaleightsuccessional

woodyspecieshad fromone to sevenindividuals 2 m tall present.These successionalwoodyspecies

presumablyoriginatedfrom seeds dispersedonto
the site fromindividualsringingthe marginof the
clearing.The marginsof farm sites are seldom
farmedbecause these areas burn poorly and are
quicklyovergrown
withsuccessional
vegetation.
We

TABLE 4. Changes in densityand species numberin 27 permanent1 x 1.5 m plots during the first16 monthsof succession followingthe final farm-plotweeding.
Months following final weeding
4
Mean numberof individuals plot-1:
forbs'
grasses
successionalwoodye
primaryforestwoody
Total
Number of species, all plots:
forbsa
grasses
successionalwoodyc
primaryforestwoody
Total

7.8 + 3.9b
19.8?10.6
1.8+ 0.8
0.04?+ 0.08
29.4+13.4
5
8
6
1
20

10
7.8?+ 4.2
6.3 ?3.3
3.1? 1.0
0.04?+ 0.08
17.2?6.0
6
6
10
1
23

:Includes herbaceouscreepers.
% confidenceinterval.
'Includes woody vines.

h95

252

Uhl, Clark, Clark, and Maquirino

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16
3.1 ?+3.2
1.7+2.3
3.2?+1.0
0.07?+ 0.15
8.1?5.4
4
6
9
3
22

have observedother newlyabandonedfarmplots

TABLE 6. The mean numberof grass and forband woody
in the environsof San Carlosand foundthemto be
colonizersin five treatments
afterone year.Only
individuals more than 10 cm tall were considsimilarto the studyplot in vegetationlife-form
ered as established.
composition
and structural
development.
Standingcrop biomassin the abandonedfarm
Mean numberof colonizers/plot
site one yearafterthe finalyuca harvest(i.e., 16 Treatment
Grasses and forbs
Woody plants
monthsafterthefinalweeding) was 773 gramsdry
0.0
184+ 65.7a
weightm-2, a valuesimilarto otherliterature
reports Bare Soil
Yuca
0.2?0.6
130+ 62.3
forabandonedneotropicalfarmsites (table 5). Of
Pineapple
137? 130.3
0.6?1.1
thistotal,430 gramswereaboveground
successional Cashew
10.2?8.0
146+108.0
speciesbiomassand 278 gramswere biomassfrom Slash
10.2?4.2
185+115.5
crop plantsstillpresenton the site.Leafbiomassof
the successional
vegetation
was 234 g m-2 and stem '95 % confidenceinterval.
biomasswas 196 g m-2 whilerootsaccounted
foronly
in woodyplant establishTreatment
differences
about 10 percentof thetotalnon-cropbiomass.The mentcouldoccur
becauseof differential
dispersalto
low root biomassvalue may be an artifactof the certain
Indeed, soil-germinable
seed
microhabitats.
smallsamplesize. In excavations
of entirerootsys- checksat the conclusionof the experiment
revealed
tems of severalcommonsuccessionaltree species, thatsoils underthe cashewtreeshad an estimated
root biomasshas generallybeen 15-20 percentof 932 germinablewoodyseeds m-2 (all successional
totalbiomass(Uhl, pers.obs.).
species) as comparedto 126 in the slashand 74 in
The woody species presenton the abandoned the bare soil plots.Fruittrees,suchas cashew,may
farmsiteappearedto havegreaterdensitiesnearde- supplyfoodas well as perchsitesforbirds.Differcayinglogs and underfruittrees.Grassesand forbs, entialgermination
of seeds once in a microhabitat
wereable to establishin open,baresoil and/ordifferential
by contrast,
mortality
of seedlingsfollowing
areasas wellas nearlogsand underfruittrees.These germination
may also be importantin explaining
establishment
preferences
weredemonstrated
by the higherestablishment
in fruittreeand slash microresultsof the microhabitat
experiment(table 6). habitats.Slashpresenceresultsin lowersoil evaporaThe cashewand slash treatments
had significantlytion and lowersoil-surface
temperatures
(Uhl et al.
morewoodyestablishments
(p < 0.01,DuncanMul- 1981), thereby
providing
favorable
soil-moisture
contiple Range Test) than did the othertreatments,ditionsforgermination
and seedlingprotection
from
while no significant
differences
in the numbersof middayheat.The presenceof fruittreesalso protects
forb and grass colonizerswere detectableamong
and steadylitterinputs
seedlingsfromdirectheating,
treatments.
Becausepineappleand yucagrewpoorly, fromthesetreesmaylikewisefavorestablishment.
thesetreatments
were probablysimilarto the bare
soil plotsin termsof lightand soil temperature.
CONCLUSIONS
The
earlysuccessionfollowingslash-and-burn
agriTABLE 5. The standingcrop biomassof a farmsite one
compoyearafterabandonment
at San Carlosde Rio cultureat San Carlosis similarin life-form
Negroand at siteselsewhere.
sitionand structure
to otherliterature
reportsfrom
New and Old Worldtropicalfarmsites.Symboth
Mass drywt, g m-2
thatabandonedfarmplotsin
ington(1933) reported
Abovegroundbiomass:
Malaysia were dominatedby grasses (particularly
Grasses
127
Imperatacylindrica),forbs,and successionaltrees
Forbs
9
Successionaltrees
272
and shrubs.Kellman (1970) foundthatearlysucPrimaryforesttrees
0
cessionalsites in the Philippineswere coveredby
Vines
22
standsof herbs,but afterone year,softwoodtrees
Cashew trees
39
beganto overtopthe herblayerand soon formeda
Pineapple plants
239
completecover. Clayton (1958) reportedthat a
Belowground biomass:
thicketof close-growing
treesand shrubsis formed
All groups combined
65
soon afterfarm-site
in Nigeria.In the
abandonment
San Carlos,Total
773
Guatemala, Total aboveground
Budowski(1961) and Ewel (1971) deneotropics,
(Snedaker 1970)
836
scribed early-fallow
vegetationin Costa Rica as
Costa Rica, Total aboveground
tanglesdominatedby a mixtureof grasses,herbs,
(Ewel 1971)
1067
vines,and woodypioneers;Snedaker(1970) reportSlash-and-BurnAgriculture
in AmazonBasin

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253

ed thatone-year-old
fallowsin Guatemalaweredom- burn.Forbs,grasses,and successional
treesdominate
inatedbywoodyshrubswhichformeda more-or-lesson newlyabandonedfarmsites because theyhave
uniform
canopyat 2 m; and Blum (1968) reported manyseedspresentin theseed bankand/orbecause
a similarvegetation
structure
afterone yearin Pan- they have easily dispersedseeds. Forest trees,by
ama.
contrast,
have few,if any,seeds in farm-site
seed
The timerequiredto regrowa foreston a de- banksand oftenhave large,porly dispersedseeds.
primary
forestmustbe close to the farm
siteat San Carlosdependson howtheforest Therefore,
forested
is removedand on the mannerof land use follow- clearingforforesttreesto colonize.In cases where
ing removal.For example,whentheforestis cut for clearingsare verylarge (i.e., seed sourcesmanykm
timber,recoveryis rapid becausemanyforesttree away) and land use prolonged(i.e., sproutingpospecieshave the abilityto sproutfromcut stumps tentiallost), it seemssafeto predictthatthe return
(Uhl et al. 1981). If sitesare cut and burnedand to primaryforestwill take manyhundredsor perof years.
then immediatelyabandoned,successionproceeds haps eventhousands
moreslowlybecauseburningreducessproutingpotentialand reducesthe woody species seed bank
(Uhl et al. 1981). If sitesare cut and burnedand ACKNOWLEDGEMENTS
and thenfarmedforseveralyearsbeforeabandon- Thanksare extendedto GetulioGomezand Paulo Livino
ment (this study),recovery
is even slower.The re- formanyhoursof helpin thefield;Drs. CarlJordan,
Bruce
MartinKellman,and an anonymous
reviewerfor
peatedweedingsduringfarming
usuallyexhaustthe Haines,
theircriticalreadingof themanuscript;
and SaundraGreen
sproutingreservesof those stemsthat survivethe forhelp in preparation
of the manuscript.

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