256 S. Halz and G.

Placci
Hall. K. 2002. Effect af 5hrubs on tree secdling estab
lishment in an abandoned tropical pasture.Joumal
af Ecalagy 90: 179-187.
Janzen. O.H. 1988. Guanacaste Natianal Park;
tropical ecolagical and biocoltural restaratian.
In: Caims. J.J.. ed. Rehabilitating Damage Ecasys
tems, vol. 2., CRC Press, Boca Ratan, FL. pp.
143-192.
Nepstad. D.CC. Uh\. C. Pereira CA.. and Cardasa
da Silva.J.M.1996. A camparative study af tree of
tree establishment in abandoned pasture and
mature fore51 of eastem Amazonia. Oikos 76;
25-39.
Purata, S.E. 1986. Floristic and structural cbanges
during oldtield succession in Mexican tropies in
relation te site history and species availability.
Joumal oi Tropical EcoJo'gy 2:257-276.
RamirezMarcial, N.. Ganzalcz-Espinoza. M., and
Garca-Maya, E. 1996. Establecimiento de
Pinus spp en matorrales y pastizales de Los
Altos de Chiapas, Mxico. Agrociencia 30(2):
249-257.
ReyBenayas, J.M.. Espigares, T., and Castro-Diez, P
2003. Simulated effect of herb campetitian 00
planted Quercus faginea seedlings in Meditcr
ranean abandoned cropland. Applied Vegetatian
Sciencc 6:213-222.
Slocum, M.G. 2000. Logs and fero patches as recruit
meot siles in a tropical pasture. Restoratian
Ecology 8(4):408--414.
Wunderle, J.M. 1998. The role of anima.! seed disper
sal in accelerating native forest regeneration on
degraded tropicallands. Forcst Ecalagy and Man
agement 99(1-2):223-235.
After regeneration begios on previously for
ested sites, carefully desigoed silvicultural
strategies can accelerate growth, infiuence tbe
direction of succession, increase tbe goods and
services provided, or enhance diversity.m Se
lecting proper treatment options requires an
understandng of the factors Iimitiog succes
sional change and ncreases in desiJed species.
These treatments should be designed to assist
natural processes rather tban figbt them. This is
most likely lo occur when forest restoraton
plans (1) consider and remove tbe underlying
}l1 Lamb and Gilmour. 2001
causes rather tban the symptoms oE degrada
tion; (2) are based on an uoderstaoding of suco
cession and threshold barriers that must be
overcome through desigoed ,interventions; and
(3) stimu1ate the desired successional behav
iour with mnimal interventions.
Many forest restoration prograrrunes fai!
because they do not address the underlying
causes of degradation. A nunrber of social, polit
ical, and economic factors are often the under
Iying cause of forest loss or degradation. [t is
equally important to identify the biophysical
barriers to recovery through natural successional
processes. For example, uvestock may con
tribute to degradation in sorne situations but be
3D important part of the recovery plan in other
circumstances. Forests Iimited by excessive tire
and invasive grasses may benefit from cattle that
reduce fuelloads untilthe tree canopy begins lo
close. [n contrast, forests lirnited by Iivestock
that consume high percentages of developing
seedlings benefit more (rom livcstock exclusion
tban from control of unsustainable harvest of
wood and nonwood forest products,

..
extraction). However, the recovery process can
be extremely slow or inhibited in highJy
degraded ecosyslems. Tht: principal challenges
for lhose working in forest restoralion are to
evaluate a forest's pOlenlial fer recovery and, if
necessary, to "accelerate" this process. Stimu
lating natural regeneration generaUy entails a
lower financial cost than other restoration
strategies, making it an attractive oplion for
restoring large sections of land.
Natural regenerati on can follow difierent
trajectories and velocities according to how the
different variables act in the system that is
undergoing recovery. Variables such as Iight,
hurnidity, temperature, availability of seeds and
young trees, predation, and the structure of
in.itial vegetation detennine the successional
trajectory of each si te. This implics that, in
general, succession in a region does not follow
a linear and unique trajectory, but manifests
itself in a whole range of stable and transitional
states with different likely outcomes.
lII
Thus, a
great variely of restoration altematives-mod
eUed from the spccific characteristies of the
system and the specific objectives of the
restoration project--<:an be proposed for a
given system that are compatible with the likely
outcomes of the natur-a-\ succession that would
otherwise occur.
TIte first step in the process of stimulating
forest recovery involves identifying the princi
pal factors that are acting as "barriers" or as
Forests can regenera te in. previously forcsted "facililators" lo regeneratioo. Once these
areas once lbe land ceases to be used for alter
nalive purposes (e.g., grazing, agriculture. Vallejo el al. 2003. )11
factors have beeo identified, they can be manip .
ulated to accelerate forest regeneratioo. Most
studies have identified dispersion, competition
with ht:rbaceous plants, and poor soil condi
tions as being the most important barriers for
tree settlement in abandoned farmlands
l11
(also
see section "Restoring After Disturbance" in
this book). These studies highlight the impor
- tance of physical as well as biological barriers.
On the other hand, trees, bushes, fems, and
fallen trees can also facilitate the natural recov
ery of an area.
J1l
10e remaining vegetation
attracts dispersers; microclimatic conditions
that favour the regeneration of young trees
develop undemeath tltis vegetation, which can
thus serve as "regeneration nuc1ei."lI4 The rela
ti ve infiuence of each factor on regeneration
depends on each system and 00 the temporal
and spatial scale in which the analysis is carried
out. Restoration methods that use natural
regeneration are based on barrier elimination,
stimulation of facilitating factors, or tbe como
bined rnanipulation of both types of factors. In
selecling the best methods for restoring the
forests of a particular area, it is extremely
important to study the forests intensively, in
order to understand their behaviour at difiero
ent scales.
Several factors limit the successful apptica
tion of restoration methodologies based on
stimulating natural regeneratioo:
Lack of seed sources and dispersers: [n many
cases, there are no forest remnants tbat can
bebave as seed sources at restoration sites:
therefore, nahlral regencration possibilities
rernain restricted to the existing soil seed
baok. In otber cases, there are nearby
but no seed dispersers due to the low number
of animals (e.g., birds, mammals) in the arca;
thus, natural regeneration may be largely
confined to species whose secds are dis
persed by wind.
Uncertain directionality: Allowing natural
regeneratioo to occur-without controllillg
the species' pool tbat is allowed to occupy a
restored area-does not guarantee a high
1Il HolI.I999.
m Peterson aod Haioes, 2lXXJ_
'lO Guevara et al. 1986.
36. Stirnulating Natural Regeneration 251
t
diversity of species in afores!. This may limil
tbe success of restoralion efforts in improv
ing ecoDomc value for future wood exploita
lion or other spccific activity.
Difficulty in obtaining a high forest species'
diversity: In additioo lo insufficient seeds, in
areas with scarce or degraded forest rcm
nants, there may be 'lhe added complication
that sorne species will simply not be able to
settle, thereby creating a forest with more
limited diversity of species.
Length of time required: A naturally regen
erating forest goes through more succes
siona] states, and thus rcquires more time to
reach a state similar to a mature forest than
does a plantation cornposed oC diverse
species.
Natural regeneration can be used in very dif
ferent ways when detining a landscape restora
tion strategy. Sorne examples of diJferent
methods are iUustrated below:
The Ijltora! area of southem Brazil, formerly
covered by Serra do Mar (Atlantic forest), is
now severely deforested (Fig. 36.1). Currently,
numerous aetions are being carried out to
preserve the remaining forests, and to restare
the deforested areas.
Ji5
Tree cover restoration
in tbe Environmental Protection Area of
Guaraque;:aba, is being aided by a strategy in
which small stands of piooeer species (i.e.,
generally 1000 to 5000 young trees) are planted
in the surrounding. more diverse stands (i.e..
composed of pioneer species, initial secoodary
species, late secondary ,pecies, and climax
species). The lalter are either rlantcd or are
fragments of already existing roresls in the
area.
ll6
Plantations are carried out cllher in the
whole area being restored or in half of this area,
depending on the size of the area, its proximity
)1' See Sociedade de PesqUlSll cm Vida Sclvagem e
Ambiental (SPVS). www.spvs.org.br.
'" ferrelli.2OO2.
252 S. Holz and G. Placci
FIGURE 36.1. Abandoned pastures in
Antonina Reserve (AlIantic forest,
Brazil), wherc the natural regenera
tion is U.auted by grass competition.
(Photo Silvia Holz.)
to forest patches, and the dcgradation of Lhe
systcm. Planted trces function as seed sOUTCes
facilitaling natural regeneration in tbe whole
area. The lreaLment given to the soil (e.g., c\ean
ing of grasses), tbe distance at which young
trees are planted, 2nd Lheir size are se[ccted
according to sile characteristics (e.g., type of
soil, topograpby, and use history).
' .. ,;..

..;,/""l, :;.
Ecologically and socially appropriate methods
for accelerating the forest recovery process
within protecled areas are bcing invesligated in
tbe seasonally dry tropical forest of lhr: moUD
lains of northem Thaand.
J11
The "framework
species" concepl (i.e., lbe use of pioneer and
climax species thal strongly facilita te, more
tban other species, the natural regeneration of
lbe area) has been adapted in this case. The
main characteristies of framework species trees
are: (1) high survival when planted at degraded
siles; (2) rapid growth; (3) dense and spreading
crown cover tbat shades out herbaceous weeds;
(4)fiowering and fruiting,or pnwision-of-other
resources, altractive to wildfe at a young age;
(5) resilienee to burning (in systems witb a dry
season); and (6) reable seed avaiJability, rapid
and synchronous seed gennination and pro
ll1 See Farest Restoration [n[ormation Service. www.UDep
wcrnc.orglrestoration/.
duction of healthy seedlings in containers.
CombinaLions of 20 to 30 species are used for
plantations. These plantations significantly aid
in tbe reeovery of the basie structure of forests
thal grow naturally, resist disturbances., aod
aLtracl seed-dispersing animals. thcreby faci
tating the natural regeneration of forests withio
the resloration area.
. "fn
':nr {tf'l
"': .t " 11 lo'
@
A great part of the Mediterranean Forests of
Guadalajara in Spain has been transformed by
wood extraclion and grazing into scrublands
with few tree speeies. In the Tonda de Tamajn
woodland, nalive speeies are being introduced
to inerease biodiversity and accelerate the
natural regeneration of the forest.
J18
Trce and
shrub species are seleeted using as criteria fmit
type as well as lhe ccologieal niche that each
one occupies. Efforts are made to inerease the
proportioo of speeies that are used as food by
wd boar populations (an important economic
-resmiroe-(}Hhe"area):-The remainingvegetation
in the area is used as "nurse trees," whereby
planling the young trces below the preexisting
individuals protects them from sun exposure
and againsl predation.
'" See World Wide Fund [or N.ture. Espaa. www.wwf.es.
36. Stimulatiag Na[UIlII RegeneratlO" 253
FIGURE 36.2. Cattle pas[Ure (lefl)
and regenerating forest (right) 2
years a.fter calllc was excluded in
experimental plots in Andresito
(Allantic forest, Argentina). In this
place, lree planting and grass clean
ing was necessary during the fir.;
year of exclusion. (Photo Silvia
Holz.)

v.elSiJectesU,i:,'f:t1,
.. r.,., J:f.i" ." '
...... f .... J
The area of Aodresito, in northeast Argentina,
has been identi1ied as a key area for the eon
servation of the Upper Paran Atlantic Forest
(Fig_ 36.2); remaining forests there can guaran
tee the conncctivity of the greal forest masses
of Brazil and Argentina.JI? In the framework of
a project on Forest Landseape Restoration lhat
involves a largc number of people and institu
lions, different restoration methodologies are
herng investigated.
J20
A particular problematic
issue is the invasion of forests degraded by a
native plant species bebaving as an invasive, the
ca/a (Celt, sp.), lhus inhibiling natural regener
ation. The slrategy used io this case consists (lf
the mechaoical emioation of ca/a, followed by
the planlatioo of yerba mate (l/ex paraguarien.
s')-a native tree species used in in.fusions, and
a key product of the regional economy.JlI The
fruit of yerba male altracts birds. facilitating lhe
natural regeneralion of lhe area. Growth of
canopy species is stimulated through selective
cutting, in order to obtain a yerba mate pro
duetion syslem under forest eoyer. Therefore, in
'" Di Bitetti el al. 2003.
.... See Foresl Landscape Restoration (FLR). http://www.
pand.orglforests/restoration.
JlI Holl. 2003.
addition to restoring a degradcd area, an effort
is also made to improve the financial opportu
nities of local farmcrs. This increases Lhe likeli
bood that they will implemcnt the restoration
stralegy, aod that these restored areas will be
preserved in the long termo
There is a wide variety of lools that can be used
to stimulate natural regeneration. The art of
restoring a forest landseape depends heavily on
the seleetion, eombinatiOll. and appropriate use
of different lools for eaeh stage and for each
particular case.
o Management of early stages of natural regen
eration in secondary forests: Natural regen
eralion is the most effectivc and economieal
way of restoring slightly degraded areas, wilh
a good seed han!< in the soil and forest rem
nants nearby. However, eyen lhese relatively
intaet systems should be mon.itored periodi
cally to evaluate the need to carry out enrich
menl planlations.
o Cosures: At silCS with high numbers of her
bivores, natural regeneration can be stimu
lated by limiting animal grazing, thereby
allowing the growth of woody planls.
o Elirnination of barriers using cattle and other
animals: Caltle grazing can be an effeetive,
easy. and LOexpensiye way to decrease the
254 S. Holz and G. Placci
bomass of grasses that compete with young (e.g., birds, marnmals) in an area.This can be
trees-in cases where the tree specics are acbieved by decreasing bunting activities and
themselves not palatable to cattle.
JU
pesticide use, reintroducing species, and cre.
Elimination of barriers through mechanical ating wildlife corridors.
andJor chemical metho.ds: The soil com
paction that retards the settlement of
young trees can be eliminated tluough. for
example, plougbing. Grasses can be elimi
nated tbrough herbicide application, manual
weeding (e.g., using a cane knife). or
1t is crucial to continue studying tbe Eollowing
mechanical weeding (e.g., with weeding
issues in order lo be able to develop restoration
machines).
actions based on natural forest regeneration:
In.stallation of racks to facilitate regenera
tion: Where existing vegetation does not Species' ecology: Little is kno\.\ol1 about the
present a significant barrier to natural regen phenology, reproductive biology, interactions
eration, artificial racks (e.g'
l
crosses, sticks, or with other species (e.g., polliDation, seed dis
wires) on which birds can perch can be used persion, berbivory/predation) of many plant
to increase the seed rain in an area and, species.
thereEore, help accelerate site regeneration. Dynarnics oE ecological succession: Restora
in systems with grasses that retard regenera tion involves the manipulation oE the natural
tion, natural racks (e.g., trees, bushes) are succession process; thereEore, it is necessary
often more effeetive, since tbey i.ncrease the to know the Eactors involved in the natural
seed rain aS weU as acting as shaders, decreas regeneration oE the system and the mecha
m
ing grass coverage. nisms through wbicb they function.
Planting a Eew species to stimulate regenera Bebaviour oE the system at different scales:
tion: The selectve planting oE a Eew tree For many systerns, there is [ittle inforrnation
species can help stimulate natural regenera on patterns and processes operating at dif
tion by (1) offering additional percbes tor ferent scales.
seed dispersers such as birds, and (2) sbading
out competing
Spatial distribution oE species witbio thc
restoration landscape: The presence of
species oE different ecological gmups-
strategically located within the laadscape
can belp accelerate natural regeneration at
Monitoring systems, as weU as statistical
this scale, as well as lowering significantly tbe
methods Eor comparing difEerent types of data
costs that would be incurred by planting
at different scales, are tools that need to be
young trees throughout an entire restoratian
developed Eor adjusting current restoration
arca. Planted stands with high species' diver
methodologies. Detailed records oE the bistory
sity, as we!! as rernnant forests in the land
oE site use and implernented restaration
scape, can function as "diversity islanlis,"
practices, as we!! as the use oE standardised
providing seeds to the arca througbout the
monitorlng protocols, would facilitate 5uch
restoration process
JH
comparisons.
m
The use oE nontraditional sta
Disperscr management: Another possible
tistical methods (e.g., Bayesian methods) can
tool Eor stimulating natural regeneratioo is
allow Eor more efficient evaluation oE restora
to try to ncrease the number of disperscrs
tion metbods, beca use they are more robust
when working with small samples, with DO
III Posada et al. 2000.
)lJ HolI et al. 2000.
n' Kageyama and Gandara. 2000. l1l Hall el al. 2003.
36. Slimulaling Natural Regenerllion 25S
Hall. K. 1999. Faetors IilIlltUlg tropIcal rain rorest
replica tes, or witb much noise in the system,and
regcneratiolllll abandQlIed pasture: secd rain, seed
they also allow Eor the combination of difierent
germination, microclimate and soil. Biotropica 31:
types oE data.
1l6
229-242.
Holl, K.O., Crone, E.E., and ScblJJtz, CH.B. 2003.
Landscape restoration: moving from generalitics
10 methodologies. BioSciencc 53(5):491-502.
HolI, K.O., Loik, M.E., Lin, E.H.. and Samucls. LA.
2000. Tropial montane forest restorarion in Costa.
ln general, recovery areas are perceived as non Rica: Qvercoming barriers to dispersal and estab
productive areas. If people can identify and lishment. Restoration Ecology 8(4)339-349.
appreciate the multiple functions oE these areas,
Holz, S. 2003. Atlantic Forest restoration in the
buffer zone of 19uaz National Park (Argentina).
the potential Eor preserving the forest will
Tcchnical Report (not pubJished).
rncrease, as will possibilities of implementing
Kageyama. P., and Gandara F. 2000. de
restoration projects in which natural regenera
areas ciliares. Captulo: 15. In: Rodriguez, R., and
tion will playa key role. This issue is particu
Filho, L., eds. Matas Ciliares: e Recu
larIy important lO the development of
Edusp, Sao Paulo. Brazil.
educational programmes.
Marcol, B.G., Halthausen, R.S.. Raphael. M.G..
Rowland, M.M . and Wisdom. M.J. 2001. Using
Bayesian belief networks to evaluate fish and
wildlife population viability under land manage
ment alterna ti ves from an environmental ,impact
The development oE strategies for decreasing
statement. Forest Ecology and Management 153:
operating and increasing incentives Eor
29-42
Peterson, eJ.. and Haines. B.L. 2000. Early succes
stimulating natural regeneration is essential in
sional patterns and potential fa.cilitation of woody
applying tbe restoration methods developed at
plant colonizatian by rotting logs in premontane
the experimental scale to tbe restoration oE
Costa Rica pastures. Restoration Ecology 8(4):
large areas. For example, it is important to con
361-370.
sider the increase in the production capacity of
Posada, lM., Alde, T.M., aod Cavetier. l 2000. Cattle
tbe restored area, compensation for the oppor
and weedy shrubs as restoration tools of tropical
tunity cost Eor landowners, payrnent Eor envi montane rainforest. Restoration Ecology 8(4):
ronmental services, and the implementation oE 37G-379.
tax incentives. Vallejo, R., Cortina. l, Vilagrosa, A., Seva, H" and
Alloza. J.A. 2003. Problemas y perspectivas de la
utilizacin de leosas autctonas en la restau
racin forestal. In: Rey, lM.. Espigares, T.. and
Nicolau. lM.. eds. Restauracin de Ecosistemas
Mediterrneos. Universidad de Alcal, Alcal de
Di Bitetti. M.S.. Placci, G.. and Dietz, L.A. 2003. A Henares, pp. 11-42.
biodiversity vision for the Upper Paran Atlantic
Forest ecoregion: dcsigning a biodiversity conser
I _ . I il 1'." . ',\ ; .
vatian landscape and setting priorities for conser
.
vation action. V,TWF, Washington, De.
"1",1. ,'h.: ,:
Ferre!li, A.R. 2002. Modelos de plantio para a re
Guariguata, M.R., and Ostertag R. 2001. Neotropical
In: A da Mata Atlantica
secondary forest sllccesion: changes in structural
em reas de sua Primitiva Ocorrencia Natural.
and functionai characteristics. Forest Ecology and
Embrapa Florestas, Colombo, pp. 35-43.
Management 148:185-206.
Guevara, S., Purata. S.. and Van dcr Maaler. E. 1986.
Guimaraes Vieira, Le.. llhl. C.. and Nepstand. D.
The role of remnant forest trees in tropical seco
1.994. The role of sbrllb Cordia mulrispicata Cham.
ondary suceession. Vegetatio 66:77..,s4.
as a asuccession facilitator" in an abandoned
pasture. Paragominas. Amazonia. Vegctatio 115:
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IJl Marcol el al. 200!.