Regeneration of an Atlantic forest formation in the

Journalof Tropical
Ecology(1995) 11:147-152.With2 figures
SHORT
COMMUNICATION
Regeneration of an Atlantic forestformationin
the understoreyof a Eucalyptusgrandis
plantation in south-easternBrazil
MANOEL CLAUDIO da SILVA JUNIOR, FABIO RUBIO SCARANO
and FABIO de SOUZA CARDEL
deBrasilia,Departamento
deEngenharia
CP 04357,CEP 70919
Universidade
Florestal,
of Geography,
Drummond
Universiyof Edinburgh,
Street,Edinburgh
Department
EH8 9XP
KEY WORDS: Apuleialeiocarpa,Atlanticforest,Brazil, Eucalyptus
grandis,forestrecuperation,
regeneration.
This note aims to provide data indicatingthe great potentialof Eucalyptus
plantationsforthe recuperationof degradednative tropicalforestsin Brazil.
forcharcoal,paper or wood production
Cultivationof fast-growing
Eucalyptus
startedin Brazil at the beginningof thiscenturyand by 1966 400,000ha were
in the early 1970s
coveredwithspecies of thisgenus. Rapid industrialization
favouredthe expansionofthisarea to 1,052,000ha in 1973,theworld'slargest
area coveredwith Eucalyptus
plantationsat that time (Lima 1987). Current
estimatesare ofnearly3,900,000ha (Institutode Pesquisas Florestais- IPEF,
pers. comm.). In orderto establishtheseartificialplantations,however,many
nativeforests
werecut,mostlyin thecerradoand lower-montane
Atlanticforest.
The widespreadsubstitutionof natural heterogeneousforestsby introduced
forestmonoculturesproducedsome dramaticenvironmental
consequences,as
witnessedby the frequentproblemsthatthe Eucalyptus
plantershad withpests
and diseases (BertiFilho & Fraga 1987,Zanuincio& Lima 1975).
It has now become a priorityto restoreareas of native vegetationamidst
Eucalyptus
plantationswiththejoint aims ofprotecting
watersources,sheltering
local fauna,and providingalternativeresourcesforpestsand a breedingground
fortheirpredators.
To providebackgrounddata we carriedout a surveyin 1988 ofregeneration
ofnativetreesin the understorey
ofa Eucalyptus
grandisHill ex Maiden plantation.The sitewas at CAF (Santa Barbara Agro-forestry
Company),a charcoal
147
148
MANOEL
CLAUDIO
DA
SILVA
JUNIOR
ET
AL.
production unit in the municipality of Dionisio (190 500 S, 420 28' W) in
the State of Minas Gerais, south-easternBrazil. Prior to the introductionof the
Eucalyptusplantations, the area was covered by a lower-montaneseasonal forest,
belonging to the Atlantic forest complex (Rizzini 1977). Topography varies
from a slightlyrolling plain to mountainous terrain, with a mean altitude of
450 m. The climate is humid-subtropical with a mean annual temperature of
20-23?C. Mean annual rainfall is 1450 mm, seasonally distributed,with most
rain falling during the summer months (December to February). Winter
consists of four to five dry months with a moderate annual water deficit
of 30-90 mm (Golfari 1975). Predominant soil types are red-yellow podsols
(ultisol), 'Terra-roxa' (eutrustox) and eutrophic-red-yellow podsols (alfisols)
(Camargo & Palmieri 1979, Soil Survey Staff-USDA 1975).
In 1970, this area was cleared of natural vegetation and prepared for EucalyptusgrandisHill ex Maiden plantation at 2 m X 2 m spacing. The stand was
harvested in 1978, and the stumps were allowed to coppice. In 1984, a hilly
area with mean slope of 20%, located at the margins of Lagoa Nova, was
transformedinto a 48 ha reserve forstudies of the fauna. Since then no forestry
practice has been applied.
The point-centredquarter method (Cottam & Curtis 1956) was used to make
an extensive floristicsurvey of individuals >5 cm gbh regeneratingin the Eucalyptusgrandisunderstorey.Twenty-one sampling lines were established at 100 m
intervals between the rows of E. grandis,throughout30 ha of the reserve, from
the margins of Lagoa Nova to the summit of the hill. In total 222 points at
15 m intervalswere placed on these lines. Girth and heightof recorded indidviduals were distributed in optimum class intervals according to Spiegel (1976).
Specimens were identifiedusing the herbaria of the Rio Doce State Park and
the University of Brasilia (UB).
The sample covered an estimated area of 1.37 ha and native trees in the
E. grandisplantation understoreywere estimated to have a total of 634.5 individuals ha-' with a basal area of 17.9 m2 ha-'. The 888 individuals belonged
to 123 species and 67 families (Table 1). Since much of the material was sterile,
identificationwas often difficultand 57 species were undetermined. Species
richness could be attributed to the proximityof natural forestsof Rio Doce
State Park (c. 20 km away). This may have allowed long-distance seed dispersal
into the study area. Much of the regeneration may have also originated from
the soil seed bank or fromthe resproutingof remaining underground structures.
It is interestingto note that the number of species recorded at Lagoa Nova
(123) was almost identical to that found in a floristicsurveyof adult individuals
(dbh >3.5 cm) in the natural forestof the Rio Doce Park (124) (CETEC 1982).
Calegairio (1992) found 69 species of 41 families regeneratingunder Eucalyptus
grandisand E. paniculataSm. at Belo Oriente municipality. That area is similar
in topography and vegetation cover to Lagoa Nova but differsfromit in having
dystrophicsoils. This is evidence to indicate that E. grandisdoes not show any
in south-eastern
Regeneration
of an Atlantic
forestformation
Brazil
149
Table 1. List ofthe dominantnativetropicalforestspeciessampledin the understorey
ofEucalyptus
grandis
plantationat Dionisio,StateofMinas Gerais,south-eastern
Brazil. The parameterscalculatedwereabsolute
density(AD), relativedensity(RD), absolute basal area (ABA), relativebasal area (RBA). A list of all
determinedspeciesis providedas a footnote.
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
Species
Family
AD
ha-'
RD
%
ABA
cm2/
ha-'
RBA
%
Apuleialeiocarpa
(Vog.) Macbr.
Spondiassp.
Dalbergianigra(Vell.) Fr. All. ex Benth.
Brosimum
sp.
Cecropia
sp.
colubrina
Anadenanthera
(Vell.) Brenanvar
cebil(Griseb.)Altschul
brasiliense
Carpotroche
(Raddi.) A. GrayEngl.
Swartziasp.
Sennamultijuga
(Rich.) Irwin & Barneby
Sapindaceae
Tabebuiaheptaphylla
(Vell.) Tol.
Pipersp.
Allophyllus
sp.
Vermonia
sp.
Siparuna
guianensis
Aubl.
Tabernaemontana
sp.
Aegiphilla
sellowiana
Cham.
Machaerium
sp.
Ocoteaodorifera
(Vell.) Rohwer
Myrtaceae
Leg.-Caes.
Anacardiaceae
Leg.-Faboideae
Moraceae
Moraceae
Leg.-Mimo.
71.8
33.7
29.3
41.0
32.2
11.7
11.0
5.2
4.5
6.3
4.9
1.8
2740
1203
1560
615
1126
1968
15.2
6.7
8.7
3.4
6.3
10.8
Flacourtiaceae
Leg.-Caes.
Leg.-Caes.
Sapindaceae
Bignoniaceae
Piperaceae
Sapindaceae
Compositae
Monimiaceae
Apocynaceae
Verbenaceae
Leg.-Faboideae
Lauraceae
Myrtacae
34.4
33.0
27.8
24.9
13.9
23.4
19.0
19.8
8.8
13.9
12.4
8.8
7.3
4.3
5.3
5.0
4.3
3.8
2.1
3.6
2.9
3.0
1.3
2.1
1.9
1.3
1.1
0.6
238
293
324
164
582
115
122
218
660
253
494
217
232
133
1.3
1.6
1.8
0.9
3.2
0.5
0.7
1.2
3.7
1.4
2.7
1.3
1.3
0.7
21. Siparuna
22. Tremamicrantha,
23. Virolaoleifera,
24. Bixa orellana,
25. Cariniana
reginae,
legalis,26. Escheweilera
brauna,
rhodogonoclada,
27. Joanesia
princeps,
28. Cordiasp., 29. Psidium
guayava,30. Tabebuiasp., 31. Melanoxylon
34. Mabeafistulifera,
36. Saratosperma
37. Guapirasp., 38. Ficus
32. Poutenia
sp., 33. Genipaamericana,
leucanthum,
41. Platycyamus
sp., 39. Moraceae, 40. Soroceailicifolia,
43. Virolasp., 45. Bauhiniasp., 46. Zeyhera
regnellii,
47. Guareaguidonia,49. Nectandra
tuberculosa,
sp., 50. Chorisiaspeciosa,51. Nectandra
sp., 52. Vemonia
sp., 53,
Erythrina
54. Eriotheca
crista-galli,
sp., 56. Hymenolobiumjaneirense,
57. Vitexsp., 60. Dimoiphandra
sp., 61. Licania
73. Nectandra
77. Caseariasp., 82. Pseudobombax
sp., 62. Bignoniaceae,66. Cordiatrichotoma,
reticulata,
sp., 84.
Cariniana
86. Amaiouasp., 87. Spondiassp., 89. Houvenia
estrelensis,
dulcis,94. Inga sp., 96. Erythroxylum
sp., 99.
119. Lueheadivaricata,
122.
sp., 107. Machaerium
acutifolium,
120.Xylopiaaromatica,
Cupaniasp., 106.Aspidosperma
Bignoniaceae,123. Leguminosae.
allelopathic effectstowards many native species that colonize its understoreyin
differentregions.
Girth frequency distribution for the total sample shows that 41% of the
individuals were found in the firstclass (5-10 cm). The largest girthwas 91 cm
colubrinavar. cebil.
(dbh = 28.9 cm), belonging to an individual of Anadenanthera
Only 8.8% of the individuals fell into the girth class of 30 cm (dbh = 9.5 cm)
or over. A reversed-Jdistributionwas found (Figure 1) which already indicates
a structureexpected formature communities. Sampled individuals were distributed in 13 height classes. The maximum height recorded was 26.4 m for a
Cecropiasp., a very fast-growingpioneer species. The majority of individuals
(46%) were in the second height class (2.9-4.9 m) (Figure 2).
These results indicate that even after 10 years the presence of Eucalyptus
grandisdid not reduce species diversityin understoreyregenerationand favoured
150
MANOEL
DA
CLAIUDIO
SILVA
ET
JUNIOR
AL.
300
250
?
200
150
0
E
C
Z 100
50
N
E
ia
pq a-
Xn
01-
N~J
1-
N'J
NN
1CN
N
-
c'i
N
r
CNt
cN
fl
N1'
ED
CD
N
N-
N-
mN
co
co
CD
Girthclasses (cm)
Figure 1. Girthdistribution
forthe totalsampling.
the establishmentof a young communitycharacteristicof an advanced stage of
succession. This is demonstrated by a number of characteristics: species richness, girth distribution showing a reversed-J pattern and some individuals
reaching considerable size (to 91 cm gbh and 28.5 m tall).
Stands of E. grandisand many other fast-growingtrees of similarlyhomogeneous, evenly spaced, non-stratifiedartificialforestsallow much higher levels of
light reaching the lower strata and are thus probably more favourable habitats
forregenerationthan the heterogeneous, dense, multi-storeyedstands of native
forest.
Apuleia leiocarpareached the highest density and basal area per ha of the
native species. Almost 87% of its population were less than 10 m tall, while the
maximum was 15.2 m and no individual had yet reached the canopy layer. At
Belo Oriente it was also the dominant tree in the understoreyunder Eucalyptus
grandisand E. paniculata(Calegairio 1992). However, it was not one of the most
frequentspecies among the adult trees and regenerationin the native forestat
Rio Doce State Park, where it was reported as an emergenttree receivingdirect
sunlight (CETEC 1982). Light sensitivityof this species may be related to age.
While Rizzini (1971) and Reitz et al. (1979) cite it as a light-demandingspecies
and Leite & Hay (1989) suggest that is is shade-sensitive, Maixner & Ferreira
(1979) argue that this species does not tolerate high light intensitiesin its early
stages.
Regeneration
of an Atlanticforestformation
in south-eastern
Brazil
300 -
250-
200
151
N
-
~~~~~~~~~~~~~~~~~~~3TotalI
;Apuleialeiocarpa|
C150
0
0
E_
.0
Zoo
50
colb)a
LO
a
C)
U
co)
U)
Classes ofheight(m)
Figure2. Heightdistribution
forthe total samplingand ofApuleialeiocarpa.
In our study the dominance of A. leiocarpain the community indicates the
presence of an intermediate stage of succession that is obviously very different
from that of the native vegetation in the Rio Doce State Park, where the
dominant adult species, typical of climax or near-climax vegetation, are
stronglyrepresentedin the regeneration. It is also differentfromthe community that one would expect to find after 10 years in an abandoned clear cut
area.
In conclusion our study indicates that a stand of Eucalyptus
grandiscan provide
the degree of shade necessary for the regenerationof forestsof natural species.
There appears to be no allelopathy and E. grandisacts as a nurse-treespecies,
playing a role that would probably be taken by Cecropiaspp. and other pioneers
under natural conditions. E. grandisplantations can thus be an effectivemeans
of promotingnatural regenerationand recoveryof tropical forestsin Brazil.
ACKNOWLEDGEMENTS
We thank Cia Agricola e Florestal Santa Barbara for our funding,Dr Walter
Suiter Filho and the staffof Dionisio unit for their support, Dr Dorothy S. D.
Araujo, Dr James A. Ratter for their useful comments on the manuscript.
152
MANOEL
CLAUDIO
DA SILVA
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JUNIOR
ET AL.
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Accepted 6 February 1994