Ecological Restoration and Reforestation of Fragmented Forests in Kianjavato, Madagascar

Abstract

A reforestation effort in Kianjavato Commune in southeast Madagascar is presented that combines native diversity with rapidly growing introduced and native pioneer trees. This work utilizes a three-tiered corridor design that capitalizes on the region’s mountainous terrain. The process of seed selection, transplantation, and survival rate of seedlings over a 16 month period is reported. The uppermost 50% of each mountain is planted with 38 woody species and most closely approximates native forest. This tier was divided into two categories, pioneer and secondary species. Most of the pioneer species were not native; however, results showed that four fast-growing, environmentally-tolerant native species could be suitable alternatives:
Streblus mauritianus, Syzygium bernieri, Treculia madagascariensis
and
Uapaca thouarsii
. More than 70,000 seeds of secondary species were extracted from fecal samples from wild, free-ranging black and white ruffed lemurs; the majority of which germinated significantly better after gut passage. The most effective pretreatment that enhanced germination was to scarify unwashed seeds. Commercially valuable trees, belonging to the community members, were grown on the lower half of each mountain. Lastly, the various contributions of the community are described along with agroforestry development plans designed to reduce pressure on forest resources and generate supplemental income.

Full text

Hindawi Publishing Corporation
International Journal of Ecology
Volume , Article ID ,  pages
http://dx.doi.org/.//
Research Article
Ecological Restoration and Reforestation of Fragmented Forests
in Kianjavato, Madagascar
Christophe Manjaribe,1,2 Cynthia L. Frasier,3
Bakolimalala Rakouth,1and Edward E. Louis Jr.2,3
1D´
epartement de Biologie et Ecologie V´
eg´
etales, Facult´
e des Sciences, Universit´
e d’Antananarivo, BP566,
Antananarivo 101, Madagascar
2Madagascar Biodiversity Partnership, NGO, VO 12 Bis A, Manakambahiny, Antananarivo 101, Madagascar
3Omaha’s Henry Doorly Zoo and Aquarium, Center for Conservation and Research, 3701 South 10th Street, Omaha, NE 68107, USA
Correspondence should be addressed to Edward E. Louis Jr.; genetics@omahazoo.com
Received  January ; Revised  September ; Accepted  September 
Academic Editor: Ram C. Sihag
Copyright ©  Christophe Manjaribe et al. is is an open access article distributed under the Creative Commons Attribution
License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly
cited.
A reforestation eort in Kianjavato Commune in southeast Madagascar is presented that combines native diversity with rapidly
growing introduced and native pioneer trees. is work utilizes a three-tiered corridor design that capitalizes on the region’s
mountainous terrain. e processof seed s election,t ransplantation, and survival rateof seedlings over a  month period is reported.
e uppermost % of each mountain is planted with  woody species and most closely approximates native forest. is tier was
divided into two categories,pioneer and secondary species. Most of the pioneer species were not native; however, resultsshowed that
four fast-growing, environmentally-tolerant native species could be suitable alternatives: Streblus mauritianus, Syzygium bernieri,
Treculia madagascariensis and Uapaca thouarsii. More than , seeds of secondary species were extracted from fecal samples
from wild, free-ranging black and white rued lemurs; the majority of which germinated signicantly better aer gut passage. e
most eective pretreatment that enhanced germination was to scarify unwashed seeds. Commercially valuable trees, belonging to
the community members, were grown on the lower half of each mountain. Lastly, the various contributions of the community are
described along with agroforestry development plans designed to reduce pressure on forest resources and generate supplemental
income.
1. Introduction
Madagascar is world-renowned for its unique biodiversity
with entire families of endemic plants and animals. Ensuring
the survival of these species is complicated by the poorly
regulated use of the country’s natural resources. As of ,
more than % of Madagascar’s  million rural residents
were living below the poverty line []andtheWorldBank
[] reported that % of people living in extreme poverty
depend on forest resources as a direct means of support.
e combination of poverty and a skyrocketing population
has contributed to Maplecro [] ranking Madagascar h
in the world of countries at “Extreme Risk” on the Climate
Change Vulnerability Index reecting the country’s perceived
inability to withstand climate change.
e country’s vulnerability to changing weather patterns
coupled with rapid deforestation and political instability
threaten Madagascar’s citizens and wildlife. More than %
of Madagascar’s forests were estimated to have been lost
between  and  with the deforestation rate escalating
sincethepoliticalcoupin[–]. To ensure the survival
of Madagascar’s wildlife, numerous conservation programs
have concentrated on forest corridor identication, protec-
tion, and restoration as well as reforestation [,].
Two basic methodologies are typically utilized to develop
reforestation programs []. e rst approach uses large
numbers of species associated with various successional
stages that are planted at high densities, and interactions
among this original set of seedlings determine the nal forest
community [,].ismethodissimilartothe“framework

International Journal of Ecology
species method” [] and was implemented in a Brazilian
Amazonia restoration program on a postproduction bauxite
mining site []. e restored forest composition was similar
to that of nearby undisturbed forests aer  years, but struc-
tural recovery had not yet been attained []. A similar eort
was undertaken in northern parcels of Masoala National
Park, Madagascar, using  native tree species preferred by
frugivorous lemurs, and was designed to reconnect forest
fragments by creating corridors [,].
e second, more economical approach described by
Lamb et al. [] uses a small number of fast-growing, short-
lived tree species to create a canopy. e established canopy
shades out competing vegetation and allows for natural
colonization from nearby forests primarily from seed disper-
sal by frugivorous animals [–]. is method is similar
to the protocol followed by the Bureau de D´
eveloppement
d’Ekar Mananjary in Madagascar that created monoculture
plantations of the introduced tree, Acacia mangium (personal
communication).e rationale of planting this re-resistant
tree comes from the expectation, as with other legumes,
that improvements to soil quality through nitrogen xation
and the presence of a forest canopy will promote natural
regeneration [].
is study concentrates on a pilot project conducted
in Kianjavato, Madagascar, that combines native species
and fast growing pioneer trees in a large-scale community-
based reforestation and agroforestry initiative. Within this
commune is the Foibe Fihofanana momba ny Fambolena
(FOFIFA) Kianjavato Coee Reserve on Sangasanga moun-
tain where substantial degradation was noted by Emberton
[]. Sangasanga and nearby isolated forest parcels are signif-
icant as they support two Critically Endangered lemurs, the
greater bamboo lemur (Prolemur simus)andtheblackand
white rued lemur (Varecia variegata).
A majority of Kianjavato’s landscape is classied as
degraded humid secondary lowland rainforest, savoka,that
is, the result of slash and burn agricultural practices [].
Aer several repetitions of slashing and burning, savoka
is transformed into roranga or grassland []. Roranga is
dened as short vegetation (.–. m) dominated by grasses
with scattered small trees or shrubs. Forests that are not
targeted for agriculture are le fragmented and marooned
in a landscape dominated by cropland with a periphery of
abandoned roranga or savoka. is abandoned land provides
an opportunity for the regrowth of a few fast-growing pioneer
speciesbutisoencolonizedbyinvasiveplantssuchas
Lantana sp. [,]. is interrupts normal migration and
foraging behaviors of resident fauna, which can be devastat-
ing for local populations of threatened species [].
To expand habitat and reconnect forest fragments in
Kianjavato, the Education Promoting Reforestation Project
(EPRP) was created. is grassroots program works directly
with local leaders and residents to plant corridors across
the roranga and savoka to connect the remaining vestiges of
forests from the eastern to the western borders of Kianjavato
Commune. ese corridors are intended to accelerate natural
colonization and succession and enable the movement of
previously isolated wildlife. However, a limitation associated
with large-scale reforestation eorts using native species is
the production of sucient quantities of seedlings adapted to
the local environment. To maximize seedling production, it
was predicted that utilizing seeds extracted from the feces of
the black and white rued lemur, the primary seed disperser
in Kianjavato’s forest, would increase the production of
seedlings in comparison to extracting seeds from ripe fruits
as gut passage oen increases germination performance [–
].
Although the conversion of forest to agricultural lands
theoretically provides greater access to food, there is no
mechanism in Kianjavato, Madagascar, to replace the ecolog-
ical services provided by the razed forest. e EPRP model
aims to restore ecosystem services by approximating a native
forest using gut passed seeds and a modied framework
species method []. To address the socioeconomic factors
that led to the initial deforestation, the EPRP will provide
supplemental income through the growth of timber and
nontimber products to provide benets to area residents
greater than those extracted from slash and burn farming.
With these objectives the present study was undertaken.
2. Methods
2.1. Site Description. is research was conducted in the
commune of Kianjavato, located in southeastern Madagascar
between Ranomafana National Park and the coastal city
of Mananjary. Kianjavato is within the Vatovavy-Fitovinany
Region and the province of Fianarantsoa (Figure ). e study
was carried out as a pilot project in the eastern portion
of Kianjavato Commune from Vatovavy Classied Forest
across unprotected, deforested habitat to the Kianjavato
Ahmanson Field Station (KAFS) operated by the Madagascar
Biodiversity Partnership and Omaha’s Henry Doorly Zoo and
Aquarium (Figure ). e reforestation corridor included 
mountains and nine valleys, with % of the area consisting of
roranga and % of savoka. Valleys were intensively farmed,
mostly for rice cultivation, and were not part of the refor-
estation project. In total, the area of the pilot project corridor
was approximately  ha ranging from  to  m wide. All
necessary permits were arranged through national and local
Malagasy Ministry of Environment and Forests oces.
e natural vegetation surrounding Kianjavato is low
elevation humid evergreen forest with a tree ora dominated
by Anthostema and Myr isticaceae []. e local topography
ishillywithelevationsrangingfromtomandthelocal
climate is characterized by an annual rainfall of , mm,
falling mostly between the months of December and March.
e mean annual temperature is .∘C, with monthly means
ranging from .∘C in June to .∘C in January (FOFIFA
weather station, ). During the rainy season, the region can
experience devastating cyclones.
2.2. Community Acceptance and Involvement. Prior to begin-
ning any eld-based work, the community was approached to
gauge their interest and support for a local reforestation eort.
Meetings were arranged with the mayor, representatives from
each local village, Fokontany (the lowest recognized admin-
istrative subdivision) presidents, local traditional authority

International Journal of Ecology 
N
E
W
S
0 50 100
(km)
Commune center
Department center
Mountain forest fragment
Kianjavato Ahmanson Field
Nursery
Corridor 1
River or wetlands
Paved road
Park boundary
Station or KAFS
F : Study site for the Education Promoting Reforestation Project (EPRP) in Kianjavato Commune, Madagascar. Var e c i a v a r i e g ata fecal
samples were collected from Ambatovaky, Sangasanga, and Vatovavy as sources of seeds for the Permanent Tier.
Permanent tier
Timber tier
Non-timber tier
F : e three-tiered corridor design incorporating a Perma-
nent Tier composed of mainly native tree species on the upper %
of hills, that is, not subject to harvest and two commercial layers, the
Timber and Non-Timber Tiers, occupying the lower % of the hill
that can be sustainably harvested. Agriculture on valley oors was
not disrupted by this reforestation program.
gures, and school employees. e objective was to com-
municate the benets of intact forests to the community,
generate enthusiasm, obtain feedback, and reduce possible
future land-use conicts.
2.3. Corridor Design. In order to develop a community-based
cooperative program and establish permanent corridors for
the biodiversity in this region, the landscape was divided
into three tiers as follows: Permanent, Timber, and Non-
Timber (Figure ). e Permanent Tier occupies % of the
land committed to the reforestation project and is situated on
the upper half of the mountains. It predominantly includes
species consumed by V. v a r i eg a t a and has the greatest
diversity of all the tiers. is portion of the corridor most
closely resembles the natural assemblage found in nearby
intact forest patches with the addition of pioneer species to
provide rapid canopy cover. e Permanent Tier will not
be subject to harvesting and fullls the role of restoring
ecological services; the pilot project focused mostly on this
tier.
e remaining half of the forest corridor is occupied by
the Timber and Non-Timber Tiers. e Timber Tier occupies
% of the corridor starting just below the Permanent Tier. It
includesspeciesmostfrequentlyusedbythelocalcommunity
for fuel and construction. e species repertoire of this tier
overlaps slightly with the Permanent Tier and will be subject
to sustainable harvesting. e Non-Timber Tier provides
fruit and other products of commercial value and was planted
in the lowest % of the corridor nearly abutting the rice
elds.
2.4. Species Selection. ere are two categories of trees within
the Permanent Tier, pioneer and secondary species (Ta b l e  ;
family aliations follow APG III []andtheCatalogueof
the Vascular Plants of Madagascar []). Four pioneer species
were used for the reforestation corridor (Ta b l e  )asthey
are considered to be fast growing and tolerant to both sun
exposure and poor soil conditions. ese include the intro-
duced species, Albizia chinensis,A. lebbeck,andA. saman
oftheFabaceaefamily,whichisknownforitsnitrogen-
xingabilities.enativeHarungana madagascariensis was
also used as a pioneer species. e secondary species of the
Permanent Tier were collected from black and white rued
lemur fecal samples from three sites in Kianjavato Commune:

International Journal of Ecology
T : Species used in the Education Promoting Reforestation Project organized by tier and family.
Tier Family Scientic name Vernacular name
Permanent (Pioneer) Fabaceae Albizia chinensis (Osbeck) Merr. Aliboza
Fabaceae Albizia lebbeck (L.) Benth Bonary
Fabaceae Albizia saman (Roxb.) Benth. Enga
Hypericaceae Harungana madagascariensis Lam. ex Poir. Harongana
Permanent (Secondary) Acanthaceae Mendoncia cowanii (S. Moore) Benoist Vahimpotsy
Anacardiaceae Abrahamia a. sericea (Engl.) Randrian. & Lowry Malambovony lahy
Anacardiaceae Poupartia chapelieri (Guillaumin) H. Perrier Voamainty fotsimaso
Annonaceae Uvaria combretifolia Diels Vahy
Apocynaceae Landolphia a. gummifera (Poir.) K. Schum. Voakilimena
Arecaceae Dypsis brosa (C.H. Wright) Beentje & J. Dransf. Vonitra
Arecaceae Dypsis mananjarensis (Jum. & H. Perrier) Beentje & J. Dransf. Lafa
Burseraceae Canarium boivinii Engl. Sandramy
Burseraceae Canarium madagascariense Engl. Ramy
Burseraceae Protium madagascariense Engl. Rar`
a
Celastraceae Salacia madagascariensis (Lam.) DC. Vahinkakao
Dichapetalaceae Dichapetalum chlorinum (Tul.) Engl. Vahinkafe
Euphorbiaceae Suregada celastroides Radcl.-Sm. & Petra Hom. Hazomby
Fabaceae Dalbergia madagascariensis Vatke Voamboana
Fabaceae Dupuya haraka (Capuron) J.H. Kirkbr. Kajaherotrala
Gentianaceae Anthocleista longifolia (Lam.) Boiteau Lendemy
Lauraceae Aspidostemon a. perrieri (Danguy) Rohwer Sily
Lauraceae Beilschmiedia opposita Kosterm. Hazombato
Lauraceae Cryptocarya dealbata Baker Tavolo lavaravina
Lauraceae Cryptocarya ovalifolia (Danguy) van der Wer Tavolo boribory
Lauraceae Cryptocarya thouvenotii (Danguy) Kosterm. Hazomboanjo
Lauraceae Ocotea nervosa Kosterm. Varongy
Menispermaceae Burasaia madagascariensis DC. Vodihazo teloravina
Moraceae Streblus mauritianus (Jacq.) Blume Ampaliala
Moraceae Treculia madagascarica N.E. Br. Friampaly
Moraceae Trophis monta na (Leandri) C.C. Berg Mahanoro
Myrtaceae Syzygium bernieri (Drake) Labat & G.E. Schatz Voamborizany
Oleaceae Noronhia introversa H. Perrier Tsilaitra beravina
Oleaceae Noronhia urceolata H. Perrier Tsilaitra madinidravina
Phyllanthaceae Uapaca thouarsii Baill. Voapaka
Rhamnaceae Bathiorhamnus louvelii (H. Perrier) Capuron Hazoaraka
Rhamnaceae Bathiorhamnus macrocarpus (Capuron) Callm., Phillipson & Buerki Hazoaraka beravina
Rubiaceae Coea perrieri Drake ex Jum. & H. Perrier Kafeala
Sapindaceae Macphersonia gracilis O. Hom. Voanemba/Sanira
Sapindaceae Tina apiculata (Radlk.) Radlk. ex Choux Letsy ala
Sapotaceae Chrysophyllum boivinianum (Pierre) Baehni Rahiaka
Sapotaceae Chrysophyllum perrieri (Lecomte) G.E. Schatz & L. Gaut. Voantsikidy
Sapotaceae Sideroxylon betsimisarakum Lecomte Tavia
Timber Euphorbiaceae Croton mongue Baill. Molanga
Euphorbiaceae Suregada celastroides Radcl.-Sm. & Petra Hom. Hazomby
Fabaceae Dalbergia madagascariensis Vatke Voamboana
Lauraceae Cryptocarya dealbata Baker Tavolo lavaravina
Meliaceae Khaya madagascariensis Jum. & H. Perrier Hazomena

International Journal of Ecology 
T : Continu ed.
Tier Family Scientic name Vernacular name
Non-timber Anacardiaceae Anacardium occidentale L. Voambarika
Annonaceae Annona muricata L. Voantsokina
Annonaceae Annona squamosa L. Zaty
Fabaceae Tam a r i n d u s i n d i c a L. Kily
Lauraceae Cinnamomum camphora (L.) J. Presl Ravintsara
Lauraceae Persea americana Mill. Zavoka
Malvaceae eobroma cacao L. Kakao
Moringaceae Moringa oleifera Lam. Ananambo
Sapindaceae Nephelium litchi Cambess. Litchi
Ambatovaky, Sangasanga, and Vatovavy (Figure ). ese
species are typical of mature forests and represent the fruit-
based portion of the lemurs’ diets and were predicted to grow
more slowly than the pioneer species.
Suregada celastroides,Cryptocarya dealbata,andDalber-
gia madagascariensis areincludedinthePermanentTieras
well as the Timber Tier as they are valued for their wood. e
leaves,notthefruits,ofthelatterspeciesareconsumedby
V. v a r i eg a t a . e Timber Tier also includes Croton mongue
and Khaya madagascariensis that are frequently used by local
residents for construction and as handles for tools. Additional
species of economic importance were selected for the Non-
Timber Tier (Tabl e  ) that do not require harvesting of
the tree itself but provide fruits and essential oils. ese
include Anacardium occidentale (cashew), Annona squamosa
(custard apple, sweetsop, or zaty), Annona muricata (beef
heart or soursop), Cinnamomum camphora (ravintsara),
Moringa oleifera (ananambo), Persea americana (avocado),
Tamarindus indica (tamarind), and eobroma cacao (choco-
late). Nephelium litchi (litchi)wasaddedatalaterdateasitis
a proven income generator and performs well.
2.5. Seed Collection, Germination, and Statistical Analysis.
Seeds of pioneer, timber, and non-timber trees were pur-
chased from regional and local seed vendors. Seeds of
secondary species were obtained by following black and white
rued lemur groups and collecting fecal samples. Vare ci a
variegata individuals were immobilized following the pro-
tocol described by Louis et al. []. Selected individuals
from dierent family groups were tted with radio collars
to facilitate monitoring and collection of fecal samples by
local eld assistants [,]. Fecal samples were bagged
individually and collection was done throughout the year
to ensure that all dominant food species were represented
in the reforested corridors. Each plant species that was seen
consumed by the black and white rued lemur was noted and
marked and voucher specimens were identied by botanists
at the national herbarium (TAN) and the rst author.
Seeds were germinated in a nursery setting where
seedlings were raised to be transplanted into the corridor.
Seeds for pioneer, timber, and non-timber species were
presoaked for  hours before sowing into cold frames as
recommended by Comtet and Rabevohitra [], Manjaribe
[], and the Silo Nationale des Graines Foresti`
eres (SNGF;
unpublished data). Cold frames are simple structures that
sit on the ground with enclosed sides and have lids made
of light wooden frames with clear plastic stretched across
them. e growing medium within the cold frames consisted
of a mixture of local soil, compost, and sand. ese were
positioned under a nursery structure partially shaded by
ravenala fronds.
Seeds for the Permanent Tier collected from black and
white rued lemur fecal samples were subjected to six
dierent treatments to identify the method with the high-
est germination success to maximize eciency in seedling
production. As seeds from the same fecal sample are not
independent, seeds for each treatment per replicate for this
study were collected from multiple fecal samples to reduce
the likelihood of pseudoreplications []. Each treatment
in a replication had  seeds for a total of  seeds per
species per replication. Seeds extracted from fecal samples
were subjected to four treatments: scaried and washed
(SW), scaried not washed (SNW), not scaried and washed
(NSW), and not scaried and not washed (NSNW). Seeds
were also extracted directly from fruit and scaried (CS) or
not scaried (CNS).
Signicant dierences in germination percentages of
treatments were detected using an ANOVA. Although 𝐹-
tests can be applied to balanced designs as in this work,
the heteroscedastic nature of germination trials increases the
probability of type  errors [,]. erefore, ANOVAs were
followed by the Tukey HSD test [], which is more tolerant
to data with unequal variance, but with balanced sample sizes.
Two species that were incorporated into the Permanent
Tierwerenotpartofthegerminationtrials:Streblus mari-
tianus and Dupuya haraka. e seeds of Streblus maritianus
were too small to be scaried, and the seeds of Dupuya haraka
were too large to be swallowed by the black and white rued
lemur. e lemurs eat the eshy portion of Dupuya haraka
fruits and then drop the seeds to the ground. us, only data
were reported for these species aer transplanting seedlings
to the eld.
Aer germination, all seedlings were transplanted to
polyethylene growing bags with drainage holes. Seedlings
were maintained at the nursery for – days until they
reached a height of – cm. Before planting into the corri-
dor, seedlings were hardened o to better withstand full sun
exposure for – days depending on the planting schedule.

International Journal of Ecology
2.6. Corridor Planting and Evaluation. Bamboo stakes were
spaced four meters apart following the contour of the refor-
estation corridor such that they were parallel to the steepest
slope. Seedlings were planted at the base of each bamboo
stake with a nal density of  seedlings per hectare. Holes
for the seedlings, approximately  cm3in size, were dug by
hand by local residents participating in the pilot program.
Grasses and ferns that were cut in the process of digging the
holes were used as mulch around the seedlings to capture
humidity and provide temporary shade and future biological
compost. e area characterized as savoka was near the
edge of the Vatovavy forest, which has shade provided by
shrubs and trees indicative of this vegetation formation [].
As the existing canopy was sucient to provide shade for
seedlings and the soil is of a higher quality than in roranga,
only Permanent Tier species were planted here. In areas of
roranga, species from all tiers were planted following the
corridor design. Pioneer species were planted rst, and the
other species were interplanted among them.
Each seedling was assigned a unique ID number stamped
on a metal tag nailed to a eucalyptus stake placed next to it.
Seedling survival was evaluated at ve and  months aer
planting. Stem basal diameter was measured using calipers
at ve cm above the root collar and height was measured
from the base to the apex. Additionally, GPS coordinates were
recorded for each seedling planted.
3. Results
3.1. Seed Germination. For all taxa, seeds germinated an
average of  to  days aer sowing except for Albizia
chinensis,A. saman,andAnnona squamosa which took four
to six days. Forty-ve to ninety days aer sowing, the number
of seeds that produced shoots that broke the soil surface was
recorded and used to calculate the percentage of seeds that
successfully germinated per treatment.
irty-eight woody species representing  families were
collected from V. v a r i e g at a fecal samples and incorporated
into the Permanent Tier (Table ). ese species included
shrubs,trees,andlianas.Nosignicantnegativeeectsof
gut passage by V. v a r i e g a ta were observed on the secondary
species subjected to the germination treatments except for
Cryptocarya thouvenotii (Ta b l e ). Generally, gut passage
enhanced germination as all variations of treatments for seeds
extracted from feces of  species had signicantly greater
germination than those extracted from fruits (𝑃 < 0.05).
Seeds can be collected from fruit or fecal matter for Crypto-
carya and Noronhia species, as dierences in the germination
percentages for either sets of treatments were not signicant.
Other than these two genera, there were no clear treatment
preferences favored between closely related species.
Seeds extracted from fruits of the following species
did not germinate or had nominal germination (% or
less): Anthocleista longifolia,Beilschmiedia opposita,Burasaia
madagascariensis,Dypsis brosa,D. mananjarensis,Macpher-
sonia gracilis,Ocotea nervosa,Poupartia chapelieri,Protium
madagascariensis,Sideroxylon betsimisarakum,andUvaria
combretifolia. Species-specic protocols can be derived from
Table , but the treatment with the highest overall germi-
nation percentage for Permanent Tier species was to scarify
unwashed seeds. is treatment performed signicantly bet-
ter than all other treatments for seven species (𝑃 < 0.05)
and had the highest germination percentages for  species
(Table ).
3.2. Planting. Transplanting of seedlings into the corridor
occurred mostly during the rainy season from January to
June. e pioneer species were planted between February and
March , and the secondary species were planted in the
Permanent Tier between May and June . e Timber and
Non-Timber Tiers were planted in mid-December  and
April through May . Overall,  species were represented
in the corridor with more than , seedlings planted from
February  to May .
3.3. Seedling Survival and Growth. Aer one year, manual
weeding was required around the seedlings to improve
seedling survival and encourage noncompetitive growth. e
rst evaluation was performed in June  and focused on
the survival of pioneer seedlings. e second evaluation was
carriedoutinJuneandAugustandincludedallspecies.
e survival and growth of the seedlings in the corridor were
used as measures of their performance. e proportion of
seedlings that survived six to  months varied among site,
category, and species.
e pioneer Albizia saman had the highest survival rate
of Permanent Tier seedlings at .%, followed by Harun-
gana madagascariensis (.%) and Albizia chinensis (.%)
oneyearaerbeingplantedintothereforestationcorridor
(Table ). Albizia lebbeck had the lowest survivorship of the
pioneer species during the same time period. Predation by
herbivores, especially on Albizia saman,wasthegreatest
cause of death for the pioneer species.
Albizia chinensis had the highest relative growth rate in
height and stem diameter with some individuals producing
owers and fruits aer one year. is species had a mean
relative growth rate of . cm in height per month and
. cm in stem basal diameter. Six native secondary species
outperformed the remaining pioneer species in growth rate
(Table ).
Survival rates varied widely for secondary species. Poor
performance was noted in Tina apiculata,bothChrysophyl-
lum species, Dichapetalum chlorinum,andOcotea nervosa.
e death of the majority of seedlings for all of the aforemen-
tioned species occurred mostly in roranga,wheresurvival
rates ranged from . to .%. However, rates improved
when these seedlings were planted near the edge of the
existing forest in savoka (.–.%).enativespecieswith
the highest survival rates in roranga and savoka were Antho-
cleista longifolia,Beilschmiedia opposita,Canarium madagas-
cariense,C. boivinii,Cryptocarya dealbata,Dalbergia mada-
gascariensis,Protium madagascariensis,Streblus maritianus,
Suregada celastroides,Syzygium bernieri,Trecul i a madag a s-
carica,andUapaca thouarsii (.–.%), demonstrating a
tolerance for harsher environmental conditions (Table ).

International Journal of Ecology 
T : Results of the germination study comparing treatments with standard deviations in parentheses. Four treatments were performed
on seeds extracted from feces: SW (scaried and washed), SNW (scaried not washed), NSW (not scaried and washed), and NSNW (not
scaried and not washed). Two treatments were performed on seeds extracted from fruit: CS (control scaried), CNS (control not scaried).
Treatments in bold text performed signicantly better than other treatments. Means sharing the same superscript within a species do not
signicantly dier from each other (Tukey’s HSD, 𝑃 < 0.05); means determined to be signicantly dierent from all other means do not have
asuperscript.
Scientic name Seeds
planted
Number of
Replicates
Treatment (average % germination)
SW SNW NSW NSNW CS CNS
Abrahamia a. sericea   100.0 (0.0) . (.)a. (.)a. (.)a. (.)b. (.)b
Anthocleista longifolia   . (.)a43.0 (1.00) . (.)ab . (.)b. (.)c. (.)c
Aspidostemon a. perrieri   . (.)a. (.)a45.0 (3.83)b50.8 (1.71)b. (.)a45.0 (4.97)b
Bathiorhamnus louvelii   . (.) 63.0 (2.58)a60.0 (2.94)a. (.) . (.) . (.)
Bathiorhamnus macrocarpus   . (.)b69.0 (2.00)a65.0 (1.00)a. (.)b. (.)c. (.)c
Beilschmiedia opposita   . (.)a58.0 (1.53) . (.)a. (.) . (.)b. (.)b
Burasaia madagascariensis   . (.) 90.8a(2.99) . (.) 88.0a(2.58) .b(.) .b(.)
Canarium boivinii   74.3 (1.15)a74.0 (1.73)a75.0 (2.00)a71.0 (2.65)a. (.) . (.)
Canarium madagascariense   . (.)b77.0 (2.71) . (.)a. (.)a. (.)b. (.)
Chrysophyllum boivinianum   . (.)a. (.)a72.0 (3.24)b72.0 (4.30)b. (.)c. (.)c
Chrysophyllum perrieri   73.0 (0.58)a77.5 (2.00)a82.0 (2.52)a80.0 (2.65)a. (.)b. (.)b
Coea perrieri   . (.) 75.0 (2.16) . (.)a. (.)a. (.)b. (.)b
Cryptocarya dealbata   . (.)ac . (.)c53.0 (2.65)ab 59.0 (3.61)b. (.)c. (.)c
Cryptocarya ovalifolia   . (.)ac . (.)a66.0 (2.65) . (.)bd . (.)b. (.)cd
Cryptocarya thouvenotii   . (.)a. (.)b. (.)b. (.)b. (.)ab 59.3 (4.16)
Dichapetalum chlorinum   70.0 (2.65)a73.0 (2.65)a72.3 (3.51)a67.0 (3.61)a. (.)b. (.)b
Dypsis brosa   . (.)a. (.) 48.0 (1.73) . (.)a.  (  .   ) b. (.)b
Dypsis mananjarensis   22.0 (1.83)a21.0 (2.94)a. (.)b. (.)b. (.)c. (.)c
Landolphia a. gummifera   . (.)a89.0 (2.65) . (.)a. (.)b. (.) . (.)b
Macphersonia gracilis   . (.) 92.0 (1.73) . (.)a. (.)a. (.)b. (.)b
Mendoncia cowanii   60.0 (2.65)a57.0 (3.00)a61.0 (3.46)a53.0 (6.08)a. (.)b. (.)b
Noronhia introversa   100.0 (0.00)a. (.)b100.0 (0.00)a100.0 (0.00)a100.0 (0.00)a. (.)b
Noronhia urceolata   96.0 (2.65)abc 100.0 (0.00)a98.3 (2.08)ab 100.0 (0.00)a. (.)bc . (.)c
Ocotea nervosa   . (.) 72.0 (2.65)a72.0 (2.65)a. (.) . (.)b. (.)b
Poupartia chapelieri   67.0 (3.46)ab 73.0 (2.00)a. (.)b69.0 (3.00)ab . (.)c. (.)c
Protium madagascariensis   . (.)b73.3 (1.15)a. (.)b69.3 (2.08)ab . (.)c. (.)c
Salacia madagascariensis   . (.)a66.0 (1.00)b. (.)ac 64.0 (4.36)bc . (.)d. (.)d
Sideroxylon betsimisarakum   . (.)a66.0 (1.41)b56.0 (4.24)ab 64.5 (3.54)b. (.)c. (.)c
Suregada celastroides   85.3 (2.52)a90.7 (3.06)a. (.)b. (.)b. (.)c. (.)c
Syzygium bernieri   94.0 (2.00)a94.0 (2.65)a90.7 (2.52)a90.0 (4.00)a. (.) . (.)
Tina api culata   78.0 (3.00)a. (.) . (.)b70.0 (4.36)ab . (.)c. (.)c
Treculia madagascarica   . (.) 62.0 (3.51) . (.) . (.) . (.)a. (.)a
Tro phi s montan a   78.0 (3.00)a81.7 (2.08)a. (.)b. (.)b. (.) . (.)
Uapaca thouarsii   . (.)a73.0 (2.65) . (.)a. (.)a. (.) . (.)
Uvaria combretifolia   . (.) 88.0 (2.16) . (.)a. (.)a.  (  .  ) b. (.)b
e timber species Khaya madagascariensis (.%),
Cryptocarya dealbata (.%), and Suregada celastroides
(.%) had the highest seedling survival from nursery to
corridor in this tier, which decreased to .% for Croton
mongue (Table ). e seedling survival data for C. dealbata
and S. celastroides for the Timber Tier is independent of
the Permanent Tier. All Dalbergia madagascariensis seedlings
survived through the evaluation period, although very few
wereplantedincomparisontomostotherspecies.is
sought-aer rosewood species attained the least amount of
height amongst the timber species, but it expanded its girth
quicker than others.
Eight species were selected for the Non-Timber Tier. e
survival rates of Tamarindus indica and Persea americana

International Journal of Ecology
T : Percent survival and growth rates of seedlings in the corridor.
Tier Scientic name Seedlings planted Seedling survival (%) Average growth rate (cm/month)
Basal diameter Height
Permanent Tier (Pioneer) Albizia chinensis  . . .
Albizia lebbeck  . . .
Albizia saman  . . .
Harungana madagascariensis  . . .
Permanent Tier (Secondary) Abrahamia a. sericea  . . .
Anthocleista longifolia  . . .
Aspidostemon a. perrieri  . . .
Bathiorhamnus louvelii  . . .
Bathiorhamnus macrocarpus  . . .
Beilschmiedia opposita  . . .
Burasaia madagascariensis  . . .
Canarium boivinii  . . .
Canarium madagascariense  . . .
Chrysophyllum boivinianum  . . .
Chrysophyllum perrieri  . . .
Coea perrieri  . . .
Cryptocarya dealbata  . . .
Cryptocarya ovalifolia  . . .
Cryptocarya thouvenotii  . . .
Dichapetalum chlorinum  . . .
Dupuya haraka  . . .
Dypsis brosa  . . .
Dypsis mananjarensis  . . .
Landolphia a. gummifera  . . .
Macphersonia gracilis  . . .
Mendoncia cowanii  . . .
Noronhia introversa  . . .
Noronhia urceolata  . . .
Ocotea nervosa  . . .
Poupartia chapelieri  . . .
Protium madagascariensis  . . .
Salacia madagascariensis  . . .
Sideroxylon betsimisarakum  . . .
Streblus mauritianus  . . .
Suregada celastroides  . . .
Syzygium bernieri  . . .
Tina api culata  . . .
Treculia madagascarica  . . .
Tro phi s montan a  . . .
Uapaca thouarsii  . . .
Uvaria combretifolia  . . .
Timber Tier Croton mongue  . . .
Cryptocarya dealbata  . . .
Dalbergia madagascariensis  . . .
Khaya madagascariensis  . . .
Suregada celastroides  . . .
Non-Timber Tier Anacardium occidentale  . . .
Annona muricata  . . .
Annona squamosa  . . .
Cinnamomum camphora  . . .
Moringa oleifera  . . .
Persea americana  . . .
Tam a r i n d u s i n d i c a  . . .
eobroma cacao  . . .

International Journal of Ecology 
trees were % during the evaluation period, although only a
small number was planted. e survival rate of Cinnamomum
camphora seedlings in the reforestation corridor was also
high at %. Moringa oleifera and eobroma cacao had the
lowest survival rates, .% and .%, respectively. Both
species of Annona had similar seedling survival rates (.–
.%), which were close to that of Anacardium occidentale at
.% (Table ). Moringa oleifera is known for its medicinal
properties and high protein foliage []aswellasitswater-
clearing characteristics []. is tree had the highest growth
rate of the Non-Timber Tier and expanded its girth faster than
the quickest growing pioneer species, Albizia chinensis.
3.4. Community Participation. Fiy-eight people from the
Kianjavato Commune attended the rst of the reforesta-
tion village meetings. is meeting resulted in a pledge by
Fokontany and villages of four sites for the nurseries and
the enrollment of  private landholders in the reforestation
pilot project. To prevent future disputes on land enrolled
in the reforestation program, the MBP prepared contracts
that were signed by community and government ocials.
ese contracts stipulate that nurseries are donated in per-
petuity to the fokonolona (the people of the Fokontany)
and that participation in the program is contingent upon
leaving the Permanent Tier intact. Participants are able to
sustainably harvest from the lower % of the reforested
land that represents the Timber and Non-Timber Tiers. e
consequences of violating this agreement will be determined
by a local environmental code and enforcement body that was
reinvigorated to protect remnant forests as well as reforested
areas. In total, approximately , seedlings were voluntarily
planted by community members on three separate occasions
in .
4. Discussion
Forest corridors increase the potential of maintaining viable
populations of biota in fragmented landscapes by enhancing
connectivity [–]. To improve the resilience of the cor-
ridor planted as part of the EPRP pilot project, a parallel
corridor will be installed. is will create a zone between the
corridors that is protected from anthropogenic disturbance
thus promoting natural successional processes and expand-
ing the width, a key factor expected to inuence the vertebrate
assemblage [].
e high visibility tactic of linking the EPRP to the
Critically Endangered black and white rued lemur main-
tains this species as a prominent subject of education events.
Additionally, constant monitoring of groups reduces the
opportunity for illegal hunting. Awareness eorts commu-
nicate the ecological services provided by V. v a r ie g a t a ,akey
seed disperser for Madagascar’s lowland eastern forests [],
especially for large seeds that exceed the gape of smaller
frugivorous lemurs. Overdor and Strait []conrmedthe
eciency of this lemur as a seed disperser based on its
home range size, daily path lengths, and seed intactness aer
digestion. is study demonstrated that seeds subjected to
gut passage generally have a higher germination percentage
than those extracted from the fruit, as has been shown in
other studies [,]. erefore, these seeds are especially
desirable for the quick and ecient production of seedlings
for the reforestation eort. Five native tree species are pos-
sibly dependent on endozoochory for germination (Tab l e  );
thereby restricting their use in more traditional reforestation
programs. For the minority of species in this study whose
germination was not aected by gut passage, the lemurs may
still be important for dispersal away from the parent tree.
By extracting seeds from black and white rued lemur
fecal samples from various collection sites, the EPRP built
an inventory of genetically diverse large and small-seeded
native tree species that were adapted to the environment.
is provided high quality seedlings that are currently
persisting through the establishment phase, which typically
lasts between three and ve years, while seedlings are in
intense competition with weeds and especially vulnerable to
environmental stresses [].
A typical indicator of success during the establishment
phaseisthesurvivalrateoftrees[]. For this study, seedling
survival was generally higher in savoka near the forest edge
than in adjacent areas of roranga.ismaybeattributedto
dierences in microclimate conditions such as soil nutriment,
shade, and humidity. Dichapetalum chlorinum,Ocotea ner-
vosa,Chrysophyllum boivinianum,C. perrieri,andTinopsi s
apiculata were observed to be sensitive to sun exposure as
these seedlings survived well under a moderate canopy but
didpoorlyinopengrasslands.Ahighersurvivalrateof
seedlings planted in sites with remnant trees compared to
those planted in sites without a tree canopy was also reported
from a Mexican cloud forest []andinamountainforestin
Costa Rica [].
Trends associated with poor seedling performance in
exposed areas prompted the use of pioneers, especially
Fabaceae species, to facilitate the growth and survival of
young trees by ameliorating local microclimate conditions
[,], improving soil chemical and physical properties [,
], and suppressing aggressive grasses that could compete
with forest seedlings [,]. ree of the four pioneer
species in this pilot program performed well with survival
rates exceeding % in the corridor; however, two of the
most successful species are not native, Albizia chinensis and
A. saman (Tab l e  ). e performance of Streblus maritianus,
Syzygium bernieri,Treculia madagascariensis,andUapaca
thouarsii suggests that they could also be used as pioneers
in reforestation eorts based on their survival rates in the
nursery and the corridor as well as their high growth
rates (Tab l e  ). ese species can shi the dependency on
using introduced species as pioneers to native species in
Madagascar’s humid forest.
Forest trees growing in open areas oen fruit years earlier
than when in a closed forest as reproductive maturity for
many canopy tree species may be triggered by exposure to full
sun, inducing fruiting as early as three years aer germination
[,]. In this study, Albizia chinensis and Syzygium bernieri
had fruit aer  and three months, respectively. Establish-
ment of a second generation of Malagasy native trees will
be the rst indication that the EPRP yielded a reproductive
population of a target species. e growth and reproduction

 International Journal of Ecology
of these two fast-growing trees will result in stands stocked at
 individuals/ha using the design of the pilot project and
will facilitate natural regeneration by native woody plants in
their understories [].
Seedling predation was clearly a factor inuencing
seedling establishment as some trees were subject to high
levels of herbivory. It is widely recognized that high tree
mortality can result from the eects of insect herbivores [].
However, the average survival rate of .% for secondary
species in this study is comparable to that of the restoration
of Ambatotsirongorongo, a newly dened protected area in
the coastal forest of southern Madagascar, which utilized
native species with a survival rate of % []. is study also
exceeds the mean survival rate of .% aer one year for
seedlings planted in the perimeter of a forest on Madagascar’s
central high plateau []. is may be due to the use of
herbaceous plant material removed in the process of planting
the trees as mulch around the seedlings and manual weeding
aer one year.
Drayton and Primack [,] stressed that the success of
programs that aim to establish new populations of perennials
should be measured by the presence of subsequent repro-
ductive generations. e existence of these young individuals
would be an indicator that the reforestation program has
transitioned from the establishment to the building phase,
the period during which trees grow, reproduce, and die [].
erefore, an eective evaluation strategy for the EPRP will
be a long-term eort reecting the time frame of the trees’ life
cycles.
Another measure of success will be the EPRP’s long-
term eect on the rate of conversion of forest to agricultural
land. e slash and burn cropping system in Madagascar
is deeply rooted in the country’s culture, although it was
only sustainable when the population was low. Farmers
rst cultivated the lower parts of valleys and moved slowly
uphill, allowing elds to go fallow for years at a time to
naturally rebuild their soil nutrient stocks []. However, the
country’s population is rapidly increasing, so fallow periods
are becoming shorter and productivity is declining forcing
farmers to deforest hills at alarming rates. Styger et al. []
estimate that at the current fallow use frequency a rainforest
can transition to roranga within – years and depending
on the number of fallow cycles may require a minimum of 
years before it is again capable of rice production.
is pilot project’s reforestation area consisted of %
roranga, a vegetation category that is unsuitable for the
production of many staple crops. If roranga is burned, it
transforms into “impoverished grasslands” that are relegated
to cattle grazing and managed with an annual re regime
as only the youngest growth is palatable to livestock [].
ese grazing areas do not regenerate and are referred to
asdeadlandsinsomeareasofthecountry[]. erefore,
thepresenceoflargeareasofroranga should be warning
ags for communities that survive as subsistence farmers. e
EPRP may prevent that nal transitional step of roranga to
dead lands and is a model that could be introduced to other
communities that are also on the cusp.
To maximize the probability of obtaining both physical
(i.e., increasing forest cover and protecting biodiversity) and
nonphysical (i.e., boost income, improve health, and raise
environmental awareness) objectives, the EPRP addressed
numerous drivers of reforestation success summarized by
Le et al. []. ese range from the reliable production of
high quality seedlings adapted to the environment to the
careful consideration of socioeconomic factors that inuence
the long-term maintenance of forested areas. Other refor-
estation projects in Madagascar [,,] demonstrated
that community participation increased the probability of
success. Recognizing the need for sustained community
support, the EPRP drew local residents into the planning
and implementation phases and included landholders and
nonlandholders.
e signing of contracts by individuals and government
ocials delineating the boundaries of reforested areas and
stating the ownership of forest products assists with address-
ing future concerns that may arise related to tenure security.
e development of business plans addressing harvesting,
marketing, and distribution of prots intends to keep this
reforestation model economically and nancially viable for
participating communities. A checks and balances system
willbedevelopedincollaborationwiththelocalenviron-
mental enforcement body that outlines appropriate penalties
for violation of land use laws. e Madagascar government
is transferring management of a portion of its protected
lands throughout the country to these small environmental
enforcement organizations.
Madagascar’s declining ability to eectively respond to
natural disasters coupled with political instability highlights
theneedforgrassrootseortsthatcombatapathyand
empower individuals and communities. With the highest
rate of poverty in Africa [], Malagasy communities are
unable to rely on governmental assistance and therefore must
leverage their own resources to sustain themselves. e EPRP
promotes a model that protects those resources, provides
income via responsible long-term agricultural practices, and
has the possibility of functioning without the continued
presence of an NGO.
Acknowledgments
isresearchwassupportedbythegenerosityoftheAhman-
son Foundation, Association of Zoological Horticulture,
Association of Zoos and Aquariums Conservation Endow-
ment Fund, Conservation International, Denver Zoo, Gog-
gio Family Foundation, Irwin Andrew Porter Foundation,
Margot Marsh Biodiversity Foundation, Mohamed bin Zayed
Species Conservation Fund, Primate Action Fund, and an
anonymous donor. is work would not have been possible
without contributions from the Omaha Zoo Foundation,
Omaha chapter of the American Association of Zoo Keepers,
and the chapter of the Association of Zoo and Aquarium
Docents, as well as V. J. and Angela Skutt Catholic High
School in Omaha, NE, USA. e authors are especially
grateful to Conservation Fusion Inc. and S. McGuire for their
dedication to environmental education and for the support
and advice of Kianjavato’s mayor, S. Lan-Tu-Hin. e authors
extend their thanks to the dedicated MBP nursery team of

International Journal of Ecology 
R. J. Razakatiana, V. T. ´
eoslas,L.Andr
´
e, and V. T. B. Nivo,
as well as to J. Andrianasolo and MBP’s committed drivers
and oce sta. e authors also wish to acknowledge the
contributions of J. Taylor, A. Ginter, and S. Holmes.
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