Alvaro Redondo Dissertation April 2010

Effects of Land-Use Change on the Conservation of Bird Species in the
Path of the Tapir Biological Corridor, Costa Rica
A Dissertation
Presented to the Faculty of the Graduate School

of
Yale University
in Candidacy for the Degree of
Doctor of Philosophy
by
Alvaro Redondo Brenes
Dissertation Director: Dr. Florencia Montagnini and Dr. Chadwick Oliver
May 2010
Abstract
Effects of Land-Use Change on the Conservation of Bird Species in the Path of the
Tapir Biological Corridor, Costa Rica
By Alvaro Redondo-Brenes
2010
I studied bird species diversity within ten different land-use types in the human-
modified and fragmented Path of the Tapir Biological Corridor, Costa Rica. The ten land-
use types were wildlife refuges, biological reserves, tree plantations, forest fallows, forest
edges, villages, residential tourism projects, homegardens, agrosilvopastoral systems, and
oil palm plantations. I was interested in determining how important the ten different
habitat types are for maintaining the bird diversity of the corridor and how these land
uses can be managed to enhance their conservation value. To address these questions I
selected 20 different sampling points within each habitat type. Bird surveys were carried
out over a two-year period. Each point was surveyed three times over the dry and three
times over the rainy periods of each year, total 12 visits per point. Total observation time
for all surveys was 400 hours. Aside from bird identification, I also recorded bird
activities (e.g. foraging, nesting) and microhabitats where they were observed (e.g. tree,
shrub, ground).
I found a total of 44,917 birds from 48 families, and 334 species. Eighty one
percent of the birds were recorded utilizing forested habitats. However, 77% of the
registered birds were also found in the human-modified land-use types. Moreover,
44.5% of species were classified as forest specialist, 38.9% forest generalist, and 16.6%
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open area specialist. Regarding feeding guilds, 53.8% of species were classified as
insectivores, 21.2% frugivores, 9% nectarivores, and carnivores and granivores 8% each.
I also identified 32 threatened species and 22 endemic species. Eight of the 22 endemic
species were also under threat, and three endangered species were registered: Ara macao,
Amazilia boucardi, and Icterus mesomelas. Overall, 64% and 75% of endemic and
threatened species, respectively, were forest-dependent species. It is suggested managing
the landscape in three units (protected areas, integrated management areas, and tree
plantations) that allow habitat for most species while providing income to local people.
Payment for environmental services, ecotourism, reforestation, environmental education
programs, and private investment, among others, could provide the incentives and
infrastructure that local people need to practice the suggested management effectively.
ii
Effects of Land-Use Change on the Conservation of Bird Species in the Path of the
A Dissertation
Presented to the Faculty of the Graduate School

of
Yale University
in Candidacy for the Degree of
Doctor of Philosophy
by
Alvaro Redondo Brenes
Dissertation Director: Dr. Florencia Montagnini and Dr. Chadwick Oliver
May 2010
iii
2010 © by Alvaro Redondo Brenes
All Rights Reserved
iv
Table of Contents
Abstract……………………………………………………………………………… i
Table and Figures……………………………………………………………………. vii
Dedication…………………………………………………………………………… ix
Acknowledgements………………………………………………………………….. x
Chapter 1: A Review of the Effects of Land-Use Change and Habitat

Fragmentation on Wildlife Species Conservation…………………………………... 1
Chapter 2: Physical, Political, and Socio-Economic Aspects of the Path of the Tapir
Biological Corridor ……………………...………………………………………….. 17
Chapter 3: Avian diversity of ten land-use types in a human-modified biological

corridor in Costa Rica……………………………………………………………….. 58
Abstract……………………………………………………………………………… 58
Introduction………………………………………………………………………….. 59
Methods……………………………………………………………………………… 61
Results……………………………………………………………………………….. 67
Discussion…………………………………………………………………………… 71
Conclusions………………………………………………………………………….. 82
References…………………………………………………………………………… 84
Tables and Figures…………………………………………………………………... 92
Chapter 4: Response of bird guilds to ten land-use types in a human-modified

biological corridor, Costa Rica…………………..………………………………….. 99
Abstract……………………………………………………………………………… 99
Introduction………………………………………………………………………….. 100
Methods……………………………………………………………………………… 102
Results……………………………………………………………………………….. 104
Discussion…………………………………………………………………………… 108
Conclusions………………………………………………………………………….. 116
References…………………………………………………………………………… 118
Chapter 5: Conservation of threatened and endemic bird species in a human-

modified biological corridor: sustainable forestry criteria recommendations............. 131
Abstract……………………………………………………………………………… 131
Introduction………………………………………………………………………….. 132
Methods……………………………………………………………………………… 134
Results……………………………………………………………………………….. 139
Discussion…………………………………………………………………………… 142
v
Conclusions………………………………………………………………………….. 158
References…………………………………………………………………………… 160
Summary and Conclusions………………………………………………………….. 177

APPENDIX 1: Abundance of 334 bird species found in the PTBC...………………. 183
vi
Tables and Figures
Chapter 2: Physical, Political, and Socio-Economic Aspects of the Path of the

Tapir Biological Corridor
Table 2.1 Summary of vegetation data for five land-use types in the Path of the
Tapir Biological Corridor, Costa Rica………………………………………………. 49
Table 2.2 Sorensen Similarity Index for woody species in four land-use types in the
Path of the Tapir Biological Corridor, Costa Rica………………………………….. 49
Figure 2.1 Location of the Path of the Tapir Biological Corridor, Costa Rica……… 50
Figure 2.2 Physical evidence of indigenous people settlements in the PTBC……… 51
Figure 2.3 Land-use change from 1992 to 2000 in the Path of the Tapir Biological
Corridor, Costa Rica. ………………………………………………………………. 52
Figure 2.4 Illegal logging and burning of mangrove forest found on a property in
Hatillo of Aguirre, Puntarenas, Costa Rica…………………………………………. 53
Figure 2.5 Political division of the Path of the Tapir Biological Corridor in three
Conservation Areas (ACOPAC, ACOSA, and ACLA-P) and four counties (Osa,
Aguirre, Perez Zeledon, and Dota)………………………………………………….. 54
Figure 2.6 Land-use types in the Path of the Tapir Biological Corridor in 2000,
Costa Rica…………………………………………………………………………… 55
Figure 2.7. Diameter size class distribution of tree species located in five land-use
types in the Path of the Tapir Biological Corridor, Costa Rica……………………... 56
Figure 2.8. Total height size class distribution of tree species located in five land-
use types in the Path of the Tapir Biological Corridor, Costa Rica…………………. 57
Chapter 3: Avian diversity of ten land-use types in a human-modified biological

corridor in Costa Rica
Table 3.1. Abundance, species richness, and diversity of birds registered in ten
land-use types in the Path of the Tapir Biological Corridor, Costa Rica…………… 92
Table 3.2. Sorensen similarity index and shared species of ten land-use types in the
Path of the Tapir Biological Corridor………………………………………………. 93
Figure 3.1. Location of the Path of the Tapir Biological Corridor, Costa Rica…….. 94
Figure 3.2 Accumulated number of bird species found in ten different habitat types
in the Path of the Tapir Biological Corridor, Costa Rica…………………………… 95
Figure 3.3. Relative abundance of the five more abundant bird families per land-
use type in the Path of the Tapir Biological Corridor, Costa Rica………………….. 96
Figure 3.4. Expected number of bird species in ten habitat types in the Path of the
Tapir Biological Corridor, Costa Rica……………………………………………… 97
Figure 3.5. Similarity of bird assemblages across ten habitat types in the Path of the
Tapir Biological Corridor, Costa Rica………………………………………….. 98
Chapter 4: Response of bird guilds to ten land-use types in a human-modified

biological, Costa Rica
vii
Table 4.1 Abundance and species richness per census (n=12) for habitat bird guilds
in 10 land-use types in the Path of the Tapir Biological Corridor, Costa Rica……... 123
Table 4.2 Abundance and species richness per census (n=12) for bird feeding
guilds located in 10 land-use types in the Path of the Tapir Biological Corridor,
Costa Rica…………………………………………………………………………… 124
Figure 4.1. Relative abundance of species by habitat guild (percentage of species) 125
within 10 land-use types in the Path of the Tapir Biological Corridor, Costa Rica…
Figure 4.2 Non-metric multidimensional scaling (NMDS) ordination of the
distribution of canopy forest specialist species in ten land-use types in the Path of
the Tapir Biological, Corridor ……………………………………………………… 126
Figure 4.3 Non-metric multidimensional scaling (NMDS) odination of the
distribution of understory forest specialist species in ten land-use types in the Path
of the Tapir Biological, Corridor ………………………………………………….. 127
Figure 4.4 Relative abundance of bird species by feeding guild (percentage of
species) within 10 land-use types in the Path of the Tapir Biological Corridor,
Costa Rica …………………………………………………………………………... 128
Figure 4.5 Relative abundance of birds (percentage of birds) for activity within 10
land-use types in the Path of the Tapir Biological Corridor, Costa Rica …………... 129
Figure 4.6 Relative abundance of birds by microhabitat (percentage of birds)
within 10 land-use types in the Path of the Tapir Biological Corridor, Costa Rica… 130
Chapter 5: Conservation of threatened and endemic bird species in a human-

modified biological corridor: sustainable forestry criteria recommendations
Table 5.1 Threatened and Endangered bird species found in the Path of the Tapir
Biological Corridor………………………………………………………………….. 169
Table 5.2 Endemic bird species found in the Path of the Tapir Biological Corridor.. 170
Figure 5.1 Abundance per census (N = 12) of endangered and threatened species 171
found in the Path of the Tapir Biological Corridor………………………………….
Figure 5.2 Abundance per census (N = 12) of endemic species found in the Path of
the Tapir Biological Corridor……………………………………………………….. 172
Figure 5.3. Relative abundance of endemic and threatened bird species by habitat
guilds found in the Path of the Tapir Biological Corridor, Costa Rica……………... 173
Figure 5.4 Relative abundance of endemic and threatened bird species by feeding
guilds found in the Path of the Tapir Biological Corridor, Costa Rica……………... 174
Figure 5.6. Non-Metric Multidimentional scaling (NMDS) ordination of threatened
and endangered species found in ten land-use types in the Path of the Tapir
Biological Corridor …………………………………………………………………. 175
Figure 5.7. Non-Metric Multidimentional scaling (NMDS) ordination for endemic
species found in ten land-use types in the Path of the Tapir Biological Corridor…... 176
viii
To my parents
Maria Eugenia and Alvaro Francisco,
who are always in my heart
ix
Acknowledgments
First of all, I would like to thank mentors, family, friends, and colleagues for their
support throughout my five years at the Yale School of Forestry and Environmental
Studies and at the Path of the Tapir Biological Corridor in Costa Rica.
My gratitude to Florencia Montagnini for the opportunity to work with her for the
last seven years, including two years as a Master of Forest Science Student. I am also
deeply grateful to my other committee members: Chad Oliver, Jonathan Reuning-Scherer
and David Skelly. Thank you all for sharing all your knowledge and for making my
dissertation thesis and time at Yale more meaningful and enjoyable.
My family and friends were always there, supporting me in the good and in the
bad moments. I am lucky to have you all. Without your support this dissertation would
not be possible.
In Costa Rica, especially in the Path of the Tapir Biological Corridor, I am
immensely thankful to all my collaborators and friends for helping me out with all the
data gathering and for the opportunity to learn new things every day I spent with you. All
your time and knowledge is invaluable. Thank you to Cristian Valenciano, my best friend
there, for the introduction to the Path of the Tapir in 2005. Thank you to ASANA, the
local NGO, and all its members, that collaborated throughout the dissertation process. I
also owe gratitude to Jack Ewing and Steve Stroud for allowing me to stay at Hacienda
Baru National Wildlife Refuge for more than two years. Thank you to all staff and friends
of ASANA and Hacienda Baru: Leti & family, Ronald, Pedro, Juan Ramon & family,
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Kika, Danny, Negro, Carlitos, Olman & family, Jose, Papo, among others I was lucky to
meet.
Aside from ASANA and Hacienda Baru, several families/organizations permitted
me to work on their properties and I am thankful to all of them. The Firestone Center for
Restoration Ecology (Pitzer College) not only provided me the opportunity to survey the
Finca Isla del Cielo, but also, Carol Brandt and Don McFarlane gave me the unique
opportunity to teach during my time in the corridor and use their facilities. Moreover,
Isabel, Greddy, Marianela, Francisco, students, and faculty were always supportive. I am
also indebted to the Odio Family (Rancho La Merced), the Duarte family (Oro Verde),
Geiner Guzman & family (La Cusinga), and all friends of La Guapil Reserve, for letting
me onto their properties. Pedro Porras was my mentor in bird identification, but also
Julian Odio, Danny and Janan Duarte, Juan Ramon, Noel Urena, and Cristian Valenciano
provided invaluable bird information for my dissertation.
Several governmental and non-governmental organizations also contributed to my
dissertation work such as INBio, MINAET, SINAC, MEP, ICE, AyA, ASADAS,
Municipalities, local organizations and of course local people.
This project was funded by the School of Forestry and Environmental Studies, the
Tropical Resources Institute, the Council for Latin American and Iberian Studies, the
Center for Field Biology Pilot Grant, and the John F. Enders Fellowships & Research
Grants, all of them at Yale University; and by the Evergreen Fellow Grant Program
through the NGO Friends of Osa and the Firestone Center for Restoration Ecology.
Thank you God for Costa Rica, for the Path of the Tapir, for Dominical and
surrounding areas. These areas and their people are unique.
xi
Chapter 1
A Review of the Effects of Land-Use Change and Habitat Fragmentation on Wildlife

Species Conservation
Introduction
The establishment of protected areas is one means of counteracting some of the
threats to biodiversity such as habitat loss and fragmentation (Sanchez-Azofeifa et al
2002; Polaski et al. 2005). As a result of this approach, in the last three decades there has
been an exponential growth in the number of protected areas worldwide, particularly in
tropical countries where biodiversity is greatest (Zimmerer et al. 2004; Naughton-Treves
et al. 2005). However, almost 90% of the world’s land surface still remains outside
formal protected areas (Dudley et al. 2005), particularly because of the socio-economic
and political constraints that limit the amount of land allocated to protected status.
Protected areas do not necessarily hold the extent of habitat needed by most living
species, forcing wildlife to seek habitat and refuge beyond the artificial boundaries of
these areas (Powell et al. 2000; Polaski et al. 2005; Dudley et al. 2005; Hilty 2006). In
fact, a recent study has shown that more than 12% of vertebrate species were not
represented in any protected area larger than 1,000 ha and in stricter conservation
classifications such as the IUCN Conservation categories (Rodrigues et al. 2004).
Costa Rica is not an exception, even though it is one of the most diverse countries
of the world (Obando 2002; 2007) and has one of the most significant systems of
protected areas (Groom et al. 2005), fragmentation and isolation of Costa Rican protected
areas are jeopardizing biological conservation (Powell et al. 2000; Sanchez-Azofeifa et
al. 2003). Given the limitations of the protected areas system in fragmented landscapes,
1
several researchers have highlighted that a new conservation paradigm must incorporate
human-modified landscapes in assessments of biodiversity and ecosystem services,
planning of corridors, establishment of buffer zones, and restoration of degraded lands
(Chazdon et al. 2009).
This study was carried out in the fragmented and human-dominated Path of the
Tapir Biological Corridor (PTBC) in Costa Rica, which is one of the most diverse areas
of Central America (TNC-ASANA 2000), but whose biodiversity is at risk because of
deforestation and development. The PTBC’s main objective is to create a network of sites
favorable to fauna (e.g. tapirs) and flora between the forests of the Osa Peninsula and
Golfo Dulce and those located in the Los Santos Forest Reserve in the Talamanca
Mountain Range. The PTBC area covers approximately 82,000 ha, inhabited by
approximately 10,000 people in 55 rural communities.
The main goal of this study was to assess the contribution of ten different habitat
types to the conservation of bird species in the PTBC. The ten land-use types were:
biological reserves, wildlife refuges, homegardens, agrosilvopastoral systems, forest
edges, forest fallows, tree plantations, oil palm plantations, local villages, and residential
tourism projects.
Chapter (1) presents a theoretical background to understand the effects of
fragmentation, land-use change, and human-dominated landscapes on the conservation of
wildlife, focusing on tropical ecosystems and bird species. Chapter (2) presents a review
of physical and socio-economic characteristics of the PTBC: a description of the goals for
the creation the corridor as well as the ten different land-use types selected as part of the
dissertation goals, including vegetation types. Chapter (3) summarizes bird diversity in
2
the PTBC, assessing differences in species composition across the ten land-use types.
Chapter (4) focuses on species composition in the ten land-use types based on bird habitat
and feeding guilds. Chapter (5) evaluates the potential contribution of these habitat types
to species of conservation concern (e.g endemic and threatened species). Furthermore,
based on the PTBC’s management goals, a set of sustainable forestry practices are
recommended.
Fragmentation and Land Use Change effects on wildlife species
Destruction or alteration of natural habitats by people because of land-use change,
represents one of the most serious threats to biodiversity worldwide (Groom et al. 2005),
along with climate change, nitrogen deposition, and invasive species (Sala et al. 2000).
As a consequence of this loss of natural habitats, there is the challenge of maintaining and
conserving biodiversity in fragmented landscapes now dominated by human land use
(Bennett 2003). Habitat fragmentation has two main components: (1) a reduction in the
area covered by a habitat type, or natural habitat generally, in a landscape; and (2) a
change in habitat configuration, with the remaining habitat left in smaller and more
isolated patches (Saunders et al. 1991). As a consequence of these changes at the
landscape level, Anderson and Jenkins (2006) suggested that habitat fragmentation may
lead to
(1) the elimination or dangerous reduction of populations of large, wide-ranging species,
including many top carnivores;
(2) the unraveling of entire biological communities – as, for example, when the decline of
top carnivores in fragmented habitats results in the “release” of smaller predators and
3
herbivores, leading to overpredation or overgrazing that may eventually eliminate
species or destabilize communities;
(3) the destruction or degradation of remaining habitat through the intrusion of edge
effects such as altered microclimates or invasive species; and
(4) the disruption of key ecological processes dependent on increasingly rare animal
agents – such as pollination, seed dispersal, predator-prey interactions, and nutrient
cycling.
The response of wildlife species to habitat destruction and fragmentation is
influenced by the species’ home range area, body size, food resources and foraging
patterns, dispersal capabilities, nesting and shelter requirements, as well as tolerance to
habitat disturbance and sensitivity to altered microclimates (Bennett 2003). Based on
these animal characteristics, the species vulnerable to forest fragmentation can be
organized into six groups (Groom et al. 2005):
(1) Wide ranging species: Species such as large carnivores and migratory ungulates that
roam a large area in the course of their daily or seasonal movements. Also, animals of
heterogeneous landscapes such as amphibians and turtles are vulnerable to
fragmentation because they depend upon distinct habitats for different phases of their
life cycles.
(2) Non-flying species: Species with poor dispersal abilities may not travel far from
where they were born, or may be stopped by barriers such as a road or a clear-cut.
Examples are carrion beetles in Amazonian forests being fragmented by pasture
4
development (Klein 1989), and arboreal mammals, peccaries, and many insect bats
that are unwilling to enter the matrix (Laurance et al. 2002).
(3) Species with specialized requirements: Species with specialized habitat or resource
requirements are often vulnerable to extinction, especially when those resources are
unpredictable in time or space. For example, the great green macaws (Ara ambigua)
nest and forage more than 90% of the time in almond trees (Dipteryx panamensis), so
the decrease of these trees has had a negative impact on the macaw population in
Costa Rica (Powell et al. 1999). Another example is the lotis blue butterfly
(Lycaeides argyrognomon lotis), an endemic species of California, USA that requires
open area. Because of suppression of natural fires in the 1970s, most grasslands
develop into dense forest, and the loss of habitat may have pushed it to extinction
(Oliver, C. pers. comm., January 2010).
(4) Large-Patch or Interior Species: Some species occur only in large patches of forests
or other habitats, and are absent from small patches with little or no true interior
habitat. For example, in Central Amazonia, many forest understory birds--including
army ant followers, solitary species, members of mixed-species flocks, and terrestrial
species--strongly avoid forest edges (Laurance 2004).
(5) Species with low fecundity: A species with low reproductive capacity cannot quickly
rebuild its population after a severe reduction caused by number of factors. For
example, large mammals such as the tapir and jaguar have low fecundity in
comparison with small mammals like rodents.
(6) Species vulnerable to human exploitation or persecution: Some species are actively
sought by people for food, furs, medicine, pets, or other uses; whereas other species,
5
such as snakes and large predators, may be killed on sight. For example, as a result of
hunting, species such as the jaguar and peccaries have decreased populations in
Corcovado National Park in Costa Rica in the last two decades. The jaguar population
dwindled from 150 individuals in 1990 to only 30 individuals in 2004, and peccaries
from 2000 in 2000 to only 400 in 2004 (Carrillo 2004).
As a result of the concerns about the effects of habitat loss and fragmentation on
wildlife conservation, there have been an increasingly large number of studies on single
species or assemblages in fragmented landscapes throughout the world (Laurance and
Bierregaard 1997; see also a large list of papers summarized by Bennett 2003). However,
most of these studies have focused on the forest fragments themselves, ignoring species'
distribution in surrounding sites such as agricultural or other human-dominated areas
(Ricketts et al. 2000). Thus, there is a need to study the importance of the landscape
matrix for biological conservation.
Importance of the Landscape Matrix for Wildlife Conservation
Assessing the conservation potential of human-dominated landscapes requires
investigation of the activities, movement, and persistence of species not only in the
remnants of native habitats but also in the full array of countryside habitats surrounding
them (Sauders et al. 1991; Daily 2001; Hughes et al. 2002), known also as the matrix.
Lindenmayer and Franklin (2002) define matrix as the landscape areas that are not
designated primary for the conservation of natural ecosystems, ecological processes, and
biodiversity regardless of their current condition (i.e., whether natural or developed). The
matrix can also be defined as the most widespread habitat within which other elements
6
are embedded, and it can be either the original habitat type (e.g., a primary forest now
surrounded by a matrix of agriculture) or a modified one (e.g., an agricultural plantation
surrounded by a matrix of pristine forest) (Anderson and Jenkins 2006). In
anthropogenically modified landscapes, some of the matrix or all of it may be human-
modified communities such as agricultural fields of various types, clear-cut forests,
pastures, tree plantations, fallow land, gardens, and towns (Daily et al. 2003; Hilty et al.
2006). In several tropical countries the spread of human-modified land-use types such as
croplands, pastures, and urban areas has been accompanied by biodiversity losses in the
last decades (Foley et al. 2005; Defries et al. 2010).
As part of the landscape, the matrix plays different roles for wildlife conservation:
(1) Matrix regulating the movement of organisms. The matrix has a significant effect
on connectivity in forest landscapes and in determining the quantity of successful
movements among habitat patches (Gascon et al. 1999; Lindenmayer and Franklin 2002;
Hilty et al. 2006), acting as a selective filter (not an absolute barrier) for the movement of
species across the landscape (Gascon et al. 1999). The type of vegetation cover in the
matrix determines the animal movement. What may facilitate the movement of one
species may be a barrier for other species (Anderson and Jenkins 2006). For example, a
matrix maintained in pasture may impede much faunal movement of forest-dependent
species (Gascon et al. 1999), but may facilitate open-area species movement (Sutcliffe
and Thomas 1996).
(2) Matrix as a resource. This is the case when the matrix can provide abundant food
resources that can be exploited by some species. In addition, the matrix may also offer
7
access to some resources that are rarely needed by a species, perhaps seasonally, such as
forage plants rich in a scarce mineral, a favorable hibernation site, or access to a
pollinator (Hilty et al. 2006).
(3) Matrix as secondary habitat. It is possible that the matrix may serve as secondary
habitat for some populations (Lindenmayer and Franklin 2002; Hilty et al. 2006). This
situation is called a source-sink axis (Pulliam and Danielson 1991). The source
population is one in which reproduction is adequate to balance mortality and usually to
export surplus individuals, as well. Such a population supplies the residents with a
secondary habitat. Sink populations are those living in secondary habitat (Hilty et al.
2006). If sink populations produce enough offspring, those may supplement populations
in reserves (Lindenmayer and Franklin 2002; Hilty et al. 2006).
(4) Matrix as a sink or stopper. The matrix may function as a dispersal sink when the
matrix is large relative to the size of patches, when it does not support a resident
population of a target species, and when few other habitat patches are available to
dispersers (Hilty et al. 2006). In addition, the matrix may act as a “stopper” when it is a
complete barrier to dispersal (Hilty et al. 2006).
Several studies addressing the effect of the landscape matrix on wildlife
conservation have been conducted in the tropics (Gascon et al. 1999; Ricketts et al. 2000;
Hughes et al. 2002; Daily et al. 2003; Horner-Devine et al. 2003; Pereira et al. 2004).
Even though the authors mentioned above assert that there is no substitute for native
habitat, they also highlight the importance of human-dominated landscapes (e.g., coffee
plantations, pastures, tree plantations, secondary forests) for the conservation of
butterflies (Horner-Devine et al. 2003), non-flying mammals (Gascon et al. 1999; Daily
8
et al. 2003), frogs (Gascon et al. 1999), moths (Ricketts et al. 2000), and birds (Lawton et
al. 1998; Gascon et al. 1999; Hughes et al. 2002; Lindell et al. 2004). For instance, in a
study of non-flying mammal species in the montane areas of Costa Rica, Daily et al.
(2003) found a total of 26 native species. From these 26 species, 9 (35%) were restricted
to forest habitat, 14 (54%) occurred in both forest and agricultural habitats, and 3 (11%)
were found only in agricultural habitats, showing that the majority of native mammals
use countryside habitats.
In the same region, Hughes et al. (2002) found a total of 144 bird species. For this
study it was estimated that 46% of those native to the region were utilizing the matrix in
some manner. Moreover, it was predicted that bird richness in the matrix would decline
to approximately 40% if tall trees and edge habitats were removed from the landscape.
Results of the Biological Dynamics of Forest Fragments Project (BDFFP) in central
Brazil also present differences in the behavior of wildlife in forest fragments and the
surrounding matrix (Gascon et al. 1999). Overall, for ants, small mammals, birds, and
frogs, it was found that between 8-15% of the nominally primary-forest species were
recorded in matrix habitats. In addition, the three vertebrate groups (mammals, frogs, and
birds) exhibited positive and significant correlations between matrix abundance and
vulnerability to fragmentation, suggesting that species that avoid the matrix tend to
decline or disappear in fragments, while those that tolerate or exploit the matrix often
remain stable or increase (Gascon et al. 1999).
9
Enhancing wildlife conservation through biological corridors
Minimizing the effects of isolation by enhancing landscape connectivity is one
way to counter the adverse effects of fragmentation (Noss 1987; Bennett 2003), in
addition to increasing the effective habitat area (Noss 1987). Connectivity can be defined
as linkages of habitats, communities, and ecological processes at multiple spatial and
temporal scales (Noss 1991). Connectivity is used to describe how the spatial
arrangements and the quality of elements in the landscape affect the movement of
organisms among habitat patches (Forman 1995). Among other alternatives, corridors are
a strategy for maintaining connectivity in the fragmented landscape (Tewsbury et al.
2002). A number of definitions and types of corridors have been proposed over time
(Forman 1995; Bennett 2003; Hilty et al. 2006; Anderson and Jenkins 2006). For
instance, Forman (1995) defined corridors as strips of habitat that differ from the adjacent
habitat on both sides.
A more detailed definition is provided by Anderson and Jenkins (2006) where
corridors are defined as spaces in which connectivity between species, ecosystems, and
ecological processes is maintained or restored at various levels. In general, habitat
corridors are defined as forested or close-forest corridors; however, species that live in
open habitats might also need open-habitat corridors for movement and dispersal (Asking
1994; Haddad and Tewsbury 2005). For instance, a study of butterflies (Aphantopus
hyperantus) in eastern England found that they used grassy tracks as corridors to move
between glades in dense woodlands (Sutcliffe and Thomas 1995). Haddad and Tewsbury
(2005) found similar pattern for two open-habitat butterfly species Junonia coenia and
Euptoieta claudia in the USA.
10
Bennet (2003) classified close-forest corridors as follows:
(1) natural corridors, such as waterways and streams and their associated riparian
vegetations;
(2) remnant corridors, such as strips of unlogged forest within clear-cuts, natural
woodlands along roadsides, and natural habitat retained as links between nature reserves;
(3) regenerated corridors, such as fencerows and hedges; and
(4) planted corridors, such as windbreaks or shelterbreaks and urban greenways.
Regardless the type of corridor, Forman (1995) mentioned that there are six main
functions that corridors provide:
(1) Conduit. A corridor acts as a conduit when it provides for movement between habitat
patches, but organisms do not reside within it (Hess and Fischer 2001).
(2) Habitat. A corridor performs as a habitat when the organisms have enough resources
for survivorship, reproduction, and movement (Hess and Fischer 2001).
(3) Filter. A corridor acts as a filter when there is some level of permeability and some
organisms and material pass through the corridor, whereas others cannot (Hess and
Fischer 2001).
(4) Barrier. A corridor may represent a barrier to animals when it is a complete blockage
and no animals can cross the corridor (Hess and Fischer 2001).
(5) Source. A corridor can work as a source when animals emanate from it (Hess and
Fischer 2001).
(6) Sink. A poorly designed corridor may act as a population sink, because the large
amount of edge exposes animals to predation from matrix dwellers and competition
11
from generalist species (Henein and Merriam 1990; Soulé 1991, cited by Hess and
Fischer 2001).
Although biological corridors can be good tools for biological conservation, there is a
large controversy about their efficacy (Beier and Noss 1998; Tewsbury et al. 2002;
Bennett 2003). Bennett (2003) summarized the critics in three main points:
(1) whether sufficient scientific evidence is available to demonstrate the potential
conservation benefits of corridors;
(2) whether the potential negative effects of corridors may outweigh any conservation
value;
(3) whether corridors are a cost-effective option in comparison with other ways of
using scarce conservation resources (e.g larger reserves).
Even though several disadvantages of corridors have been reported, such as
serving as pathways for fire, predators, and pathogens--which can each undermine
conservation objectives (Simberloff and Cox 1987; Forman 1995; Anderson and Jenkins
2006), and even though it is clear that corridors are not the solution to all of the current
conservation problems (Noss 1987), many of the potential disadvantages could be
avoided or mitigated if the proper design is applied to the specific requirements of the
species, habitats, ecosystems, and ecological processes of concern in each case (Anderson
and Jenkins 2006). However, further research has to be carried out in order to investigate
the positive or negative impacts of corridors, especially at the landscape level.
12
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16
Chapter 2
Physical, Political, and Socio-Economic Aspects of the Path of the Tapir Biological
Corridor
Introduction
The forest cover of Costa Rica has been affected by both natural and human
disturbances throughout its history. Fires, volcano eruptions, and earthquakes have been
documented as major disturbances that have changed vegetation and successional taxa
throughout the country’s history (Horn and Sanford 1992; Anchukaitis and Horn 2005;
Valerio 2006; Meza-Ocampo 2009). Although hurricanes and tropical storms do not
directly strike Costa Rica, indirect effects such as floods and landslides have also
contributed to land-use change in the country, especially on the Pacific coast (Valerio
2006; Meza-Ocampo 2009).
Even though humans have been an important cause of land-use change since they
first appeared, their impact has varied throughout time, and has become more significant
in the last century. For instance, although Costa Rica has been inhabited by humans for
more than 12,000 years (Valerio 2006), the first documented anthropogenic disturbances
on the country’s rainforest have been dated to 3,000 BP (Northrop and Horn 1996;
Clement and Horn 2001; Anchukaitis and Horn 2005).
Results of pollen and charcoal analyses in several locations have depicted that
humans disturbed the landscapes through forest clearance and fires associated with
agriculture (e.g slash and burn agriculture), which were sporadic but destructive
(Northrop and Horn 1996; Clement and Horn 2001; Anchukaitis and Horn 2005). Pre-
Columbian population was estimated to be 27,000 people (Keogh 1984). However, based
on post-Columbian evidence, it is thought that the forest area cleared was less than 2%
17
of the country’s entire area from the sixteenth century (Spanish arrived) through the
eighteenth century (Tosi 1974; Keogh 1984), particularly because of low population
growth (Keogh 1984). Throughout this colonial era, population grew from 15,500 in
1600 to 52,000 in 1800 (Keogh 1984).
In the nineteenth century, coffee production was responsible for most forest
clearance in the central part of the country as well as infrastructure development (e.g
roads – Gomez and Savage 1983). In 1800, cleared agriculture area represented between
3.5 and 5 % of the country and forest cover represented approximately 92 to 93% (Keogh
1984). The country’s forest cover changed most dramatically during the twentieth
century. Until the early 1900s, Costa Rica was still almost entirely covered by rich and
diverse tropical forests (Tosi 1974; Keogh 1984; Harvey et al. 1998; Hall 2000). By
1900, the estimated forest cover in the country was approximately 87 to 90% (Tosi 1974;
Keogh 1984).
Subsequently, as a result of rapid population growth, the forest cover was
estimated to decrease to 76 - 85% of the territory in the 1940s (Sader and Joyce 1988;
Brockett and Gottfried 2001; Kleinn et al. 2002). However, this area decreased further
until loss totaled more than 80 % of the original forest in the 1980s (Brockett and
Gottfried 2001; Kleinn et al. 2002; Joyce 2006). This resulted in part from official
policies that placed a priority on demographic growth and agricultural production
implemented during the 1950s and 1960s (Sanchez-Azofeifa et al. 1999; Brockett and
Gottfried 2001; Pfaff and Sanchez-Azofeifa 2004; Joyce 2006). Population grew
exponentially from 687,000 to 2,044,000 people during the period of 1943 – 1977
(Keogh 1984; INEC 2010).
18
Faced with rapidly disappearing forests, the Costa Rican government responded
with a multifaceted approached. First, the system of national parks was created in the
1970s. Second, a complex regulatory framework governing forestry on privately owned
lands was established through the Forestry Law of 1969 and its revisions of 1973, 1979,
1986, and 1996. Third, financial incentives were provided for reforestation, later for
natural forest management, and most recently for forest preservation (Brockett and
Gottfried 2001).
Over the 1970s and 1980s the greater part of protected areas were created (Kleinn
et al. 2002). Between 1974 and 1978 the areas protected expanded from 3% to 12% of the
national territory, and now stand at around 25-26 % (Pfaff and Sanchez-Azofeifa 2004;
Obando 2007). However, because of the isolation of these protected areas, it was
recommended to implement an appropriate network in order to enhance forest
connectivity (Powell et al. 2000; Sanchez-Azofeifa et al. 2003). This network was
expected to be achieved as part of the Mesoamerican Biological Corridor (MBC), which
aimed to connect most protected areas from Mexico to Panama. The Path of the Tapir
Biological Corridor (PTBC) is a part of the MBC.
This Chapter (2) will present a physical and socio-economic review of the region
as well as an explanation of the goals for the creation of the PTBC. The chapter closes
with a description of the ten different land-use types selected as part of the dissertation
goals, including a description of vegetation types.
19
The Path of the Tapir Biological Corridor
Physical characteristics
The PTBC is located in southwest Costa Rica (Figure 2.1). The corridor’s main
objective is to create a network of sites favorable to fauna and flora between the forests of
the Osa Peninsula and Golfo Dulce, including Corcovado National Park. These forests
connect with those located in the Los Santos Forest Reserve in the Talamanca Mountain
Range. The PTBC area covers approximately 82,000 ha, inhabited by approximately
10,000 people in 55 rural communities. This mosaic of human-dominated land uses
includes primary and secondary forests, native and exotic forest plantations, agriculture,
pastures, bamboo, oil palm plantations, ecotourism projects, and others.
The corridor is composed of terrain ranging in altitude from 0 to 1,100 masl
(TNC-ASANA 2000), and according to the Holdridge System (1967), three life zones are
represented within it: (1) The Tropical Wet Forest that comprises the entire length of the
corridor, especially the lower zones; (2) The Tropical Wet Forest Transition to
Premontane, represented by a small fringe at the intermediate altitudes; and (3) The
Tropical Rainforest Premontane found at the highest elevation in the corridor. The
climate in this region of the country is categorized as Seasonal Tropical Wet with an
annual precipitation of 4,239 mm and average temperature of 27 ºC. There is also a high
precipitation during the rainy season with as much as 685 mm per month (maximum of
1,715 in October 1988), and a pronounced 4 to 5 months dry season with little rain (less
than 100 mm) from January to May (TNC-ASANA 2000). The soils in the corridor are
young, low in fertility, and are generally classified as Ultisols and Inceptisols. There are
more than 30 rivers, among which the main ones are Coronado, Higueron, Uvita, Baru,
Hatillo, Savegre, and Guabo (TNC-ASANA 2000).
20
The PTBC is one of the most diverse regions of Central America because it
represents the transition between the dry forests of the Northeast and the rain forests of
the Osa Peninsula (TNC-ASANA 2000). This part of the country, including the Osa
Peninsula, comprises more than 2.5% of the world’s biodiversity (Obando 2007), with
more than 3000 plant, 173 mammal, 51 amphibian, and 81 reptile species identified in the
corridor (TNC and ASANA 2000, Valenciano 2005). Additionally, 500 avian species
have been registered in the area in the six most recent annual bird counts (Urena, pers.
comm. July 2009). The PTBC provides habitat to some endangered animal species such
as margay (Leopardus wiedii), ocelot (Leopardus pardalis), jaguar (Panthera onca),
spider monkey (Ateles geoffroyi), squirrel monkey (Saimiri oerstedii), scarlet macaw
(Ara macao), great curassow (Crax rubra), and endemic tree species such as quira
(Caryodaphnopsis burgeri). The PTBC region is located in one of the five centers of high
endemism in the country (Obando 2007), with 13 fish, 5 amphibian, 6 plant, 8 reptile, 11
bird, and 1 mammal species enlisted as endemic to Costa Rica.
Land-use in the PTBC
Costa Rican flora and fauna are a fairly recent amalgam resulting from a changing
geography and climate (Rich and Rich 1983; Meza-Ocampo 2009). The continuous
connection between North and South America that it provides has been in existence for
only the past three million years (Meza-Ocampo 2009). Throughout most of the
Pleistocene savanna grassland conditions had a much broader geographic spread in
Central America (Rich and Rich 1983), facilitating species dispersion between North and
South America. However, approximately two million years ago, humid tropical rain
21
forests became widespread in Central America, thus decreasing the effectiveness of the
terrestrial connection (Rich and Rich 1983). Most non-forest species, especially bird
species, persisted in those forested areas utilizing natural light gaps and extensive areas
along rivers that were maintained in early successional stages by periodic floods (Stiles
1983).
In the Holocene (11,300 – 9,600 years), flora was less dense, with some savanna
patches on the Pacific coast as a result of drier and colder weather conditions than at
present (Meza-Ocampo 2009). Subsequently, human disturbances provided more habitats
to non-forest species (Stiles 1983) as described above. The PTBC region had always been
sparsely populated, dating back to the pre-Columbian era (Newcomer 2007), and there is
still physical evidence of these human settlements (Figure 2.2). However, there are no
documented studies of their direct/indirect impacts on forest cover within the PTBC until
more recent post-Columbian times.
The current land use in the PTBC has been the result of the process of local and
national settlement and development policies, as mentioned above. Between 1940 and
1960, Costa Ricans migrated from the northern part of the country toward this region in
the southwest, searching for new lands to clear and cultivate (Newcomer 2007). Over this
period, forest lands were considered worthless on the frontier, and an individual’s
reputation for hard work depended on the amount of land one cleared (Brockett and
Gottfried 2001), thus the principal causes of deforestation were reported to be the
economic demand for land rather than for wood (Hartshorn 1982). On the Costa Rican
frontier, settlers generally occupied land by squatting (Brockett and Gottfried 2001).
22
In the 1970s and 1980s Costa Rica was mainly in the negative environmental
headlines for having one of the highest deforestation rates worldwide. The deforestation
rates consistently averaged two to three times higher than the overall regional average for
Latin America (Wendland and Bawa 1996). Over this period, the forest cover changed
from 49 % in 1969 to only 17 % in 1983 (Sader and Joyce 1988). Between 1960 and
1990, an expanding cattle industry was a major contributing factor to deforestation in the
PTBC (TNC-ASANA 2000).
From the 1990s to date, forest cover has begun to increase as a result of declining
agriculture production in the region, a fall in cattle prices, new environmental laws, and
changing regional demographics (Newcomer 2007, Redondo-Brenes 2007). For instance,
from 1992 to 2000, more than 20,000 ha of pastures were abandoned mainly because of a
decrease in the international beef market. As a result of this land abandonment, there was
an increase in the area under (1) young secondary forests (younger than ten years old), (2)
tree plantations, and (3) the area classified as primary forest (Figure 2.3). From 1997 to
2005, according to Calvo (2010), the PTBC was one of nine biological corridors in the
country that suffered from a high annual deforestation rate (12.5%), however, its net
forest cover and number of forest fragments increased. Deforestation and consequent
land-use change throughout this period was the result of development projects and
changing of forest cover to other agriculture crops such as rice (Figure 2.4). The increase
in net forest cover is likely related to the continued abandonment of pasture lands in the
PTBC, which are naturally regenerating through secondary forest succession.
Aside from anthropogenic disturbances, forest cover in the PTBC has also been
affected by natural events. Earthquakes, floods and landslides are the major natural
23
disturbances in the PTBC. For instance, intense precipitation as a result of hurricane Rita
in September 2005 on the Caribbean coast caused one of the most severe emergencies in
the PTBC in the last 10 years. An area of more than 200 km2 was affected by floods and
landslides (Mendez-Herrera and Esquivel 2006). The large concentration of sediments
along major rivers as well as along the coast exemplifies the indirect effects of hurricanes
and tropical storms during the rainy season on the Pacific coast of the country.
Design, planning, and implementation of the PTBC
Conservation awareness in the PTBC began in the early 1980s, with the creation
of the local NGO ASANA (Friends of Nature from the Central and South Pacific) by
local people being the first step to implement the corridor. ASANA has worked on
conservation issues and developed community-based sustainable projects in the last two
decades. In 1996, the idea of the PTBC was first considered, but it was not until 2000 that
it was officially recognized by Costa Rican authorities. The corridor design encompasses
high elevation lands to protect water sources for local consumption and a matrix of high
and lowlands to protect habitat for hundreds of species. The PTBC is one of the few
projects where a Rapid Ecological Assessment was carried out to justify its importance as
a conservation and development tool in Costa Rica (Ewing, J. pers. comm. July 2006).
The strategy for the creation of the PTBC consisted of three elements:
(1) Encourage land owners to create private biological reserves such as: a) National
Wildlife Refuges like Hacienda Barú and La Merced, b) Informal reserves with
no particular government recognition such as Oro Verde Biological Reserve and
24
Rafiki Safari Lounge, c) Land held in reserve through payment for environmental
services (PES), and d) Land protected by environmental easements.
(2) Encourage and assist landowners in the protection of their reserves. This is being
accomplished through environmental education in surrounding communities, and
by creating covenants through which the National Parks Foundation (NPF) hires
official park rangers, and reserve owners and neighbors reimburse the cost of the
salaries to NPF.
(3) Encourage the creation of natural corridors along rivers, streams and fences in
lands not currently held in reserve, allowing connections between forest patches.
This is being accomplished through environmental education and public
awareness (Ewing, J. pers. comm. 30 October, 2006).
Within their strategy, the PTBC organizations strive to achieve four major goals:
(1) Restore natural habitats to enable the return of terrestrial endangered species and
marine species such as the leatherback sea turtles (Dermochelys coriacea) to the
forests and beaches they inhabited half a century ago, and protect existing species
by allowing local populations to become more robust.
(2) Halt deforestation within the corridor and restore forests in order to complete
connectivity and protect water sources for use by communities, particularly in
watersheds and groundwater recharge zones.
(3) Encourage local communities to value resources and use them sustainably.
(4) Create interconnecting forest reserves between the existing network of forest and
wildlife reserves. Specific conservation programs have been developed by the
25
PTBC organizations to accomplish these goals in the region and they are another
important step in corridor implementation (Anderson and Jenkins 2006).
The main programs implemented by PTBC organizations in the last decade are as
follows:
(1) Environmental education programs in more than 32 schools and workshops for
local adults to build a better understanding of sustainable forest management. In
addition, within the PTBC there are road signs to caution drivers about the
presence of wildlife species and to inform people about the existence of the
corridor.
(2) Promotion of land conservation areas through PES in collaboration with the
government (For more details about PES see Redondo-Brenes and Welsh 2006).
(3) Community-development projects to provide better revenues to local people
through sustainable agricultural practices.
(4) Coordination to develop associations aimed at providing drinkable water to locals
and proper watershed management.
(5) Training and approval of 130 voluntary local game wardens to patrol the regions
ad honorem.
(6) Monitoring and protection of sea turtles and other flora and fauna species. To
date, some of those programs are working well and others still need more funds to
be properly implemented. Despite all these conservation programs, there are
several threats to biodiversity in the region.
26
Threats to biodiversity are development and deforestation, illegal logging, poaching
and hunting, and lack of law enforcement. This is a very attractive area for foreigner
people and tourism projects, and as a result, several real estate developers are buying the
land from locals, cutting the forest to create new roads, and building residential areas,
thus increasing fragmentation. Moreover, even though it is claimed that hunting has
decreased in the last three decades (Ewing, J, pers. comm. June 2006), it is still an issue
and hunters mostly come from communities outside the corridor. Another main constraint
for the PTBC to achieve biological conservation is the lack of funding to support further
projects. The threats to biodiversity grow when we add to the described problems a lack
of law enforcement, interest to control those problems, funds, and in a few cases
corruption from governmental organizations.
Economics
Within the PTBC most local people work in agriculture, and recently ecotourism
has grown to become another important source of income in the region (Newcomer 2007,
Redondo-Brenes 2007). However, as land prices have increased, land ownership is
shifting from local farmers to wealthy foreigners. Costa Ricans have either become
employees of the new owners or migrated to larger cities in Costa Rica or to the United
States and Europe. This poses a new challenge for the region since protected areas are
interspersed with private lands. Costa Ricans have migrated out of the region, and thus
populations have decreased in the four most important districts that share land with the
PTBC (Puerto Cortes, Savegre, Baru, and Bahia Ballena). Even though between 1973 and
2008 there was a 166% increase in population (1.7 to 4.5 million people) in Costa Rica,
27
population decreased 14% from 15,550 to 13,467 people in these four districts, of which
approximately 10,000 inhabit the corridor (INEC 2010).
Agricultural lands in the region are mostly planted with rice, oil palm, and small
scale crops such as maize, cassava, coffee, and beans, and many farmers have
agroforestry combinations of some of these crops with trees and cattle. Pastures for cattle
have different characteristics than cultivated land, as some of them have living fences of
native tree species as well as scattered remnant trees as shade for cattle. Tree plantations
are mostly timber species located in the Caribbean and Pacific lowlands of the country.
The most commonly planted species in this region, respectively, are Tectona grandis
(teak), an exotic species brought to Costa Rica from Asia, Terminalia amazonia
(amarillon), a native species planted in medium or small scale plantations, and Gmelina
arborea (melina), also exotic to Costa Rica. Teak and melina plantations represent more
than 50% of the total area of tree plantations in the country (Chinchilla-Mora and Mora-
Chacon 2002).
Conservation in the region has been undertaken through the establishment of five
National Wildlife Refuges and more than 30 private nature reserves, all of which were
established after 1995 (Ewing, J, pers. comm., June 2006). The participation of local
landowners and their willingness to maintain forest cover will determine the future
success of the PTBC project. The fate of the existing biodiversity in the region depends
on landowners’ management decisions, meaning attractive incentives are necessary to
ensure their engagement in conservation practices (Anderson and Jenkins 2006). The
PTBC organizations help landowners find economic incentives for sustainable land
management (Newcomer 2007; Redondo-Brenes 2007), by providing guidance in
28
obtaining payment for environmental services (PES) from the government (currently
more than 4,500 ha of forest are under protection through PES), funding for community-
development projects, and technical assistance to address water management and other
conservation issues in the region. Additionally, local landowners can benefit from
conserving their lands through ecotourism.
Socio-political issues
In the PTBC almost 100% of the land is privately owned, therefore, decision
making in the region involves many stakeholders (Basso and Newcomer 2009).
According to Newcomer (2002), the PTBC participants can be grouped into three main
in-country clusters: government, community organizations, and landowners. The
government is represented by a local liaison between the PTBC program and Ministry of
Environment (MINAET), and National ministries and municipal government agencies
have direct interests in and influence on the program. The PTBC consists of three
municipalities (Aguirre, Osa, and Perez Zeledon – Figure 2.5), and three conservation
areas (ACOPAC, ACLA-P, and ACOSA – Figure 2.5) led by MINAET. Municipalities
develop zoning plans and provide construction permits within their borders, while the
conservation areas are responsible for the proper management of natural resources in the
country. Currently, only the municipality of Aguirre and ACOPAC (and to a lesser extent
ACLA-P and ACOSA) are supporting the PTBC conservation and development
initiatives. Contrary to the supportive position of these parties, the Mayor of the
municipality of Osa encourages development in the region. In fact, from 2006 to 2007
29
construction permits increased 200% in this county (Redondo-Brenes and Villalobos
2008).
Community organizations include cooperatives, foundations, women’s groups,
environmental groups, agricultural groups, community development associations, and the
PTBC Coordinating Committee. These groups are very active, running most conservation
and community development projects and denouncing any illegal activity against natural
resources in the region. Finally, landowners can be divided into those who participate in
conservation and those who do not. Developers and real estate agencies comprise most of
the non participatory group, which constitutes one of the main threats to biodiversity
conservation in the area. In fact, Basso and Newcomer (2009) reported that more than
71% of people interviewed in the corridor stated their concern about the high rate of real
estate development in the PTBC.
A fourth group not mentioned by Newcomer (2002) consists of individuals and
institutions funding specific conservation and development projects within the PTBC.
Among them are AVINA Foundation, Cedarena Land Trust, Costa Rica Conservation
Trust, the Mesoamerican Biological Corridor Coordinating Committee, and the United
Nations Development Programme (UNDP).
Large-scale conservation projects in private and human-dominated landscapes
require the participation of local people, of which most who necessitate economic
incentives to ensure their support (Anderson and Jenkins 2006). To successfully acquire
local support for the PTBC project, the participants’ needs were taken into consideration
in the corridor’s objectives. For example, since water quality and quantity are important
issues in the region (Welsh 2006), and a water conservation approach has proved to be an
30
effective tool for biodiversity conservation (Redondo-Brenes and Welsh 2006), including
water as a priority in the region was a successful tactic employed by the corridor leaders
to increase local support.
Despite the work that has been done by the PTBC organizations to match
conservation and development in the region, to date, most of the biological data of the
corridor is lacking and further research is necessary to assess the conservation status of
wildlife within this fragmented corridor.
Description of the selected land-use types in the PTBC
The ten land-use types were selected based on several variables. First, available
GIS land-use layers were used to identify the main land-use types in the region, and
second, field work was done to verify the land-use maps (Figure 2.6). Thirdly, interviews
were conducted with local organizations and people to determine what they consider to
be the land-uses of most concern in the region. The fourth factor was the use of Costa
Rican bird guides as well as other studies that have assessed bird diversity in human-
modified landscapes (Stiles and Skutch 1989; Hughes et al. 2002; Peh et al. 2006;
Garrigues and Dean 2007).
At the time of my first visit, the most updated land-use layers were created in
2002 by the National Institute of Biodiversity as part of a project called Ecomapas (INBio
2005). This layer was used as a first step to determine the main land use types in the
region. By 2002, the main land uses in the region were primary and secondary forests
(35,682 ha), pastures (22,243 ha), young secondary forests (13,488 ha), agriculture (4,074
ha), forest tree plantations (2,915 ha), and wetlands (1,160 ha). Based on these land-use
31
types, a few months were spent in 2005, 2006, and 2007, re-classifying each land-use
category in the field and also talking to local organizations. For instance, for the forested
habitats, it was determined that there were four categories of interest: wildlife refuges,
biological reserves, forest edges, and forest fallows. Oil palm plantations and
homegardens were selected as part of the sites classified as agriculture. Two common
human settlements were selected: villages and the new residential tourism projects. For
tree plantations, an exotic (teak) and a native (amarillon) species were selected as will be
described below.
Wetlands, such as major rivers and mangroves, were discarded because of a lack
of accessibility and funding. Access for most rivers needed special permits and in the
case of mangroves the best access was by boat. Therefore, wetlands were considered too
expensive for the dissertation budget. Among the people interviewed were local birders,
staff from ASANA, private reserve’ owners and workers, local tourist guides, and
landowners. In addition, workers from the Minister of Environment, INBio, Neotropical
Foundation, TNC, Universidad Nacional (UNA), and University of Costa Rica were
interviewed.
In summary, based on the four described variables, ten land-use types were
selected as part of this project: wildlife refuges, biological reserves, forest edges, forest
fallows, tree plantations, oil palm plantations, homegardens, residential tourism projects,
villages, and agrosilvopastoral systems.
For the forested habitats (wildlife refuges, biological reserves, forest edges, forest
fallows), seven temporal plots of 15 m-radius were set in order to quantify the vegetation
greater than 5 cm of diameter at breast height (dbh). The seven plots were located at
32
random based on the existing 20 sampling points. Total area sampled by land-use type
was 0.5 ha. In the agrosilvopastoral systems a total of 10 50-m transects were set in order
to determine the trees greater than five cm of diameter planted along living fences and
scattered trees. In the other human-dominated land-use types (homegardens, villages,
residential tourism projects, and oil palm plantations) in ten 50-m radius plots all plants
found were identified. Results of this vegetation inventory is included as part of each
land-use description shown below.
Wildlife refuges
In Costa Rica there are more than 40 legally recognized wildlife refuges that have
been established as part of the national system of protected areas (e.g. national parks).
Wildlife refuges consist of informally protected private nature reserves that, since 1992,
qualify as official national wildlife refuges. Under this program, landowners must
develop a government-approved management plan specifying restrictions of land and
resource use. In exchange, there are three incentives: (1) property tax exemption, (2)
technical assistance for managing the property, and (3) protection against squatter
invasion (Langhloz et al. 2000). In the PTBC there were five wildlife refuges recognized
by 2005: Rancho La Merced, Hacienda Baru, Portalon, Transilvania, and Boracayan.
Rancho La Merced (329 ha) and Baru (330 ha) were selected because of accessibility and
the owner’s willingness to participate in the project. Forested areas (primary or old
secondary forest) were surveyed within the two refuges (10 sampling points at each
refuge for a total of 20 points for this type of habitat).
33
Horizontal and vertical structure of these forests, those in the biological reserves,
and forest edges were characteristic of primary and old-growth secondary forests of Costa
Rica (Figure 2.7; 2.8; Valenciano 2005). The tallest canopy trees were approximately 32
m, and there were trees with diameters of 250 cm. A total of 76 woody species from 31
families with diameters larger than five cm were identified in 0.5 ha sampled plots (Table
2.1). The most abundant plant families were Araliaceae, Tiliaceae, Annonaceae,
Moracea, and Mimosaceae. Moreover, Dendropanax arboreus, Apeiba tiborbou,
Tachigali versicolor, Hyeronima alchorneoides, and Brosimun lactecens were some of
the most common species found in the sampling plots. Brosium utile, Hura crepitans, and
Hyeronima alchorneoides were the species that dominated the canopy, and Apeiba
tiborbou, Tachigalia versicolor, and Dendropanax arboreus dominated the understory.
Tree average basal area at both refuges was 55 m2/ha and volume 620 m3/ha.
Biological reserves
Privately owned reserves continue to attract interest across the tropics for their
ability to blend biodiversity protection with sustainable development (Langholz and
Lassoie 2002). According to Langholz and Lassoie (2002), in Costa Rica there are around
250 private reserves which cover approximately 63,832 ha, or 1.2% of the national
territory. These reserves are protecting ecologically important habitat, particularly
primary rain forest, and they are used for a wide variety of activities, especially
ecotourism and for the owners’ personal enjoyment (Langholz and Lassoie 2002). The
main difference between these private reserves and the wildlife refuges is the legal
34
declaration. Private reserves do not necessarily receive tax exemptions nor receive special
protection from the government.
In the PTBC there are around 30 private reserves. Many of them are used for
ecotourism, but also there are some with restricted access. During my preliminary
assessment of the study region, I visited ten of these reserves. I selected two of these
reserves based on size (the largest ones) and accessibility and in addition, at both
locations ecotourism was practiced as well as in the wildlife refuges. La Cusinga (250 ha)
and Oro Verde (150 ha) were the selected private reserves. Ten sampling points within
each refuge were located. Horizontal and vertical structure of these forests is similar to
the forests of the wildlife refuges (Figure 2.7, 2.8). Sampling points were only located in
areas dominated by primary and/or tall secondary forests. The tallest canopy trees were
approximately 35 m. Ninety nine woody species from 39 families with diameters larger
than five cm were found in 0.5 ha sampled plots.
Tree family composition was also similar to wildlife refuges (Table 2.2).
However, the most abundant families differed between forested habitat types. In the
biological reserves, Mimosaceae, Euphorbiaceae, Sapindaceae, Myristicaceae, and
Lauraceae were the most abundant families. Moreover, tree species composition differed
across wildlife and biological reserves (Table 2.2). Canopy of these forests were
dominated by Hura crepitans, Pterocarpus hayessi, Annona amazonica, Apeiba
membranacea, and Sapranthus palanga. The understory was dominated by Croton
schedianus, Tetrachylachium macrophyllum, Cupania sp, Psycotria sp, and Socratea
exhorriza. Tree average basal area at both reserves was 41 m2/ha and volume 476 m3/ha.
Values were lower than those from the wildlife refuges.
35
Forest edge
The presence and abundance of wildlife species usually differ between forest edge
and interior habitats. Study of forest edges is relevant because there are several forest-
interior species that avoid edges and also there are many open-area species that visit those
habitats (Gascon et al. 1999), making edges highly diverse areas for birds (Stiles and
Skutch 1989). To determine any potential difference in the abundance and composition of
birds across forests and forest edges, twenty sampling points were located in areas along
the edge of primary and old-secondary forests. These forest edges were similar in
structure to the forests sampled in wildlife refuges and biological reserves. The canopy
was slightly shorter than the other forests (28 m) and was dominated by primary forest
species such as Sapranthus palanga, Virola sp, Brosimum utile, Annona sp, and Balizia
elegans. The understory was dominated by primary forest species such as Brosimum sp,
Virola sp, and Ocotea sp, as well as secondary forest species such as Casearia arborea
and Croton schidianus. Also, diameter classes were smaller in forest edges than forest
interior sampling plots (Table 2.1).
Tree species composition was different across the reserves and the edges, but for
tree families similarity index values can be considered high (Table 2.2). Thirty five
different tree families and 93 tree species were identified. The most abundant families
were Euphorbiaceae, Moraceae, Myristicaceae, Mimosaceae, and Tiliaceae. Average
basal area for forest edge stands was 39 m2/ha and volume 386 m3/ha.
Forest fallows (young secondary forest)
For the present study, forest fallows were defined as areas of forest regenerating
in recently abandoned pasture lands (1-5 years after abandonment). These fallows are
36
known as the first stage of tropical secondary succession. Pioneer herbs, shrubs and
climbers invade the site after abandonment (Finegan 1996). Likewise, pioneer tree
species of other ecological guilds colonized the stand (Finegan 1992). For the present
study, 42 woody species from 28 families with diameters larger than five cm were
identified in the seven plots that covered 0.5 ha.
Forest fallows were totally different than the forest interior and edges in tree
structure and composition (Table 2.1, 2.2; Figures 2.7, 2.8). Most of these forest fallows
were dominated by herbaceous bush, shrubs, and a few pioneer trees. The most abundant
tree families were Myrtaceae, Cecropiaceae, Sterculiaceae, Melastomataceae, and
Piperaceae. The understory of these fallows were dominated by Psidium guajaba,
Cecropia obstusifolia, Guazuma ulmifolia, Schefflera actinophylla, and Piper sp. Canopy
trees were mostly E. cyclocarpum, Cecropia obstisifolia, C. insingnis, Guazuma
ulmifolia, and Cordia cymosa. According to the Sorensen index (Table 2.2), similarity
values across fallows and the other three habitats were less than 0.58 for tree families and
0.28 for tree species.
The tallest canopy trees were approximately 13-15 m, with a few remnants that
reach 20 m such as Enterolobium cyclocarpum. Also, even though vertical structure of
forest fallows had a similar inverted J shape to the other forested habitats, forest fallow
size classes were lower (Figure 2.6). Tree basal area for forest fallow stands was 7 m2/ha
and volume 31 m3/ha.
37
Homegardens
The homegardens were traditional Costa Rican, and composed of a combination
of annual/perennial crops, ornamental plants, and timber and fruit trees. The selected
areas had a minimum of a hectare of land under this system. Most sampling points were
located in small rural villages such as San Buenaventura, Coronado, and Silencio where
this type of agroforestry system is still a part of the landscape.
Assessing the area around a radius of 50 meters from the sampling point center, a
total of 65 plant species from 34 families were identified within this habitat. The most
common crops were banana (Mussa sp), maize (zeea mais), papaya (Carica papaya), and
cassava (Manihot sculetum). Tree species were located in three main groups: timber,
ornamental, and fruit species. Among the timber species, Cedrela odorata, Samanea
saman, Enterolobium cyclocarpum, and Tabebuia rosea were some the most common
native species. Nevertheless, some exotic species such as teak (Tectona grandis) and
melina (Gmelina arborea) were seen as part of the homegardens. Fruit trees seemed to be
the most commonly used tree species in these selected sampling areas. The most common
fruit trees were mango (Magnifera indica), guava (Psidium guajava), several species of
citrus, several species of Inga spp, start tree (Averroa carambola), among others. For
ornamental purposes, Costa Ricans used Heliconia spp, palms, as well as tree species
such as beach almond (Terminalia catappa), chumico (Schefflera actinophylla), and poro
(Erythrina poepiggiana).
38
Agrosilvopastoral systems
Most of the pastures in the region were used to raise beef cattle and had mostly
been planted with exotic pasture species. At the time of this study, all of the selected
sampling sites were under used (e.g. cattle grazing). Moreover, all sites had either living
fences or scattered trees that provide perches or food sources for birds. Using line transects
in 10 out of 20 sample points, a total of 41 different woody species (28 families) were
identified with a diameter larger than five cm.
Tree species varied in composition and structure from site to site. Some sampling
points were single-species living fences, however, the majority of pasture lands had more
than one tree species planted. Diameter sizes varied from 3 cm to 76 cm. Tree height also
varied, from 2 m to 25 m. The most common tree species used were Bursera simaruba,
Gliricidia sepium, Erythrina poepiggiana, Tabebuia rosea, Enterolobium cyclocarpum
and Cupania guatemalensis.
Tree plantations
Most tree plantations in the PTBC consist of teak (Tectona grandis) and amarillon
(Terminalia amazonia). Teak is one of the most important tropical hardwood species in
the international high-quality timber market (Bermejo 2004), mostly exported to Europe
and the United States. It is native to India, Myanmar (Burma), Thailand and Laos, but
this broadleaved tree species now grows in the entire inter-tropical region (excluding
desert areas of Africa- Bermejo 2004). In Costa Rica more than 40,000 ha were planted
with teak in the lowlands of both the Pacific and Caribbean regions in the last two
decades (Kanninen et al. 2004).
39
These plantations are the first serious reforestation projects under intensive
silvicultural management. Teak has since acclimatized well and has been widely grown
in both industrial plantations and small community woodlots (Bermejo 2004). In the
PTBC there are both industrial plantations and small-scale reforestation programs,
however, management intensity at both ends of the spectrum is similar. Most plantations
that were visited did not have any understory regeneration. However, in some cases,
remnant trees and living-fences of native species were seen as part of these monocultures.
Amarillon is native to Costa Rica, and is one of the most promising species for
timber production in the country (Streed et al. 2006; Redondo-Brenes 2007; Calvo et al.
2007). Terminalia amazonia can be found naturally throughout Central America, ranging
from Mexico to South America and the Antilles, and throughout Costa Rica and much of
Central America, it has commonly been used within hardwood plantations (Jimenez et
al. 2002). The T. amazonia species is a promising plantation species because it can thrive
in almost all types of soils, it has an average growth of five m in height and 5.8 cm of
diameter after only three years in plantation, and the average survival rate within
plantations in the northern part of Costa Rica is 85% (Jimenez et al. 2002). Previous
research even suggests that T. amazonia is the most adapted native hardwood species for
survival in eroded soils (Carpenter et al. 2004).
In the PTBC, T. amazonia plantations were mostly established with government
incentives, and most were a part of small-scale reforestation programs. Several of them
established with purposes other than timber production such as restoration of degraded
landscapes. Silvicultural management of amarillon plantations differed from teak
plantations in that they did not have a defined management plan like teak plantations did.
40
Most amarillon plantations contain a dense understory dominated by herbaceous and
woody tree species.
Overall, amarillon and teak plantation sizes varied from 2 ha to 30 ha, with teak
plantations being the largest. Tree diameters varied from 9 cm to 30.5 cm and height
varies from 12 m to 22.5 m, with the teak trees having the largest overall sizes. Since
there was not enough T. amazonia to set up 20 sampling points, ten points were located in
teak and ten in amarillon plantations in order to have a total of 20 points for this habitat
type.
Oil palm plantations
In the northern region of the corridor, there were hundreds of ha of oil palm
plantations (Elaeis guineensis), which is an exotic crop to Costa Rica. All plantations
belong to Coopesilencio, a local farmers’ cooperative with more than 600 ha of oil palm
in the region. Most of them have either living fences or a few scattered trees as part of
their land-use type. The most common tree species found were Gliricidia sepium,
Guazuma ulmifolia, Enterolobium cyclocarpum, Cordia alliodora, Cecropia sp, among
others. Aside from the tree species mentioned above, natural grasses and bushes are also
part of the landscape, especially for young plantations before the canopy closes (4 m tall).
However, most grasses are manually eliminated as part of the plantation maintenance.
Many granivore bird species are found visiting young plantations.
In the Silencio community 400 people depend mainly on their palm plantations
located on the community’s 1,100 ha landholdings. The harvest season is from April to
November, and from December to March, the plantation is in reproduction mode to get
41
ready for the new harvest. To maintain their crop, different sections of the plantation are
harvested every year to ensure the sustainability of their crop. One hundred and forty
three palms are planted per ha. The life cycle of these plantations is around 20 years
when plantations reach ten meters in height. Maximum yields are obtained between 5-15
years. The sampling points were located in a range from recently established plantations
(1.5 m tall) to mature plantations (up to ten m height).
Residential tourism projects
Because the PTBC is along the Central and South Pacific Coast of the country,
there has been an exponential growth in the number of foreigners building summer or
retirement houses in the area in the last ten years. Most people live here for one or a few
periods throughout the year, and only a minority of people occupy their houses year
round. The sampling points were located in areas where more than one house was built.
Houses in residential tourism projects were mostly surrounded by forested habitats, but
most keep at least a 30 m distance between forest and the infrastructure. They were
located in different locations along the PTBC such as Ojochal, Escaleras, Lagunas,
Portalon, and Pasito. These houses were two to three times larger than houses in villages
or those with homegardens. Most of them also included exotic plant species, such as
palms, as part of their landscaping.
Regarding plant diversity, a total of 47 plant species from 33 families were
identified within the sampling points. Some of the common tree species were Tabebuia
rosea, Guazuma ulmifolia, Ficus sp, Citrus sp, Cecropia sp, and E. cyclocarpum.
42
Villages
Two main areas were selected for sampling. The first one was the largest village
within the corridor, Puerto Cortez, and here 10 sampling points were selected. The other
10 sampling points were located in the small villages along the main paved highway:
Uvita, Dominical, Hatillo, and Matapalo. Most sampling areas were a combination of
residential and commercial buildings. Construction area and population density in the
villages was greater than in homegardens or residential tourism projects.
Vegetation of these villages was a combination of native and exotic plant species.
A total of 57 plant species from 34 families were identified within the sampling points.
Some of the common plant species found in these habitats were Terminalia cattapa,
Mangifera indica, Coccos nucifera, and Ficus sp.
43
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48
Table 2.1 Summary of vegetation data for five land-use types in the Path of the Tapir
Biological Corridor, Costa Rica.
Wildlife Biological Forest Forest Tree

Refuge Reserve Edge Fallow Plantations
Families (N/0.5 ha) 31 37 37 27 2
Species (N/0.5 ha) 76 99 93 42 2
Densities (N/ha) 768 868 688 562 787
Diameter (cm) 18.3 16.6 18.7 10.2 22.9
Height (m) 9.2 9.8 10.4 5.0 20.4
Basal Area (m2/ha) 55.2 40.6 38.7 6.8 26.7
Volume (m3/ha) 629.8 476.6 386.5 30.6 272.9
Table 2.2 Sorensen Similarity Index for tree species (bold) and tree families in four land-
use types in the Path of the Tapir Biological Corridor, Costa Rica.
Biological
Reserves Forest Edge Forest Fallow Wildlife Refuges
Species
Biological Reserves 0.552 0.269 0.571
Forest Edge 0.777 0.281 0.532
Forest Fallow 0.593 0.58 0.237
Wildlife Refuges 0.805 0.738 0.596
Families
49
Figure 2.1 Location of the Path of the Tapir Biological Corridor, Costa Rica
50
Figure 2.2 There is physical evidence that indigenous people settled a few locations on
the PTBC. These petroglyphs were found in a farm in the central part of the Corridor. (A)
represents a whale, (B) a turtle, and (C) a more sophisticated map. Photos by Alvaro
Redondo-Brenes.
51
45000 1992
2000
40000
35000
30000
25000
Area (ha)
20000
15000
10000
5000
0
Tree Infraestructure Young Mangrove Agriculture Forest Pastures
plantations Secondary
Forest
Figure 2.3 Land-use change from 1992 to 2000 in the Path of the Tapir Biological
Corridor, Costa Rica. Layer Source: INBio (2005)
52
Figure 2.4 Illegal logging and burning of mangrove forest found on a property in Hatillo
of Aguirre, Puntarenas, Costa Rica. Photo taken in April 2010 by A. Redondo-Brenes.
53
Figure 2.5 Political division of the Path of the Tapir Biological Corridor in three
Conservation Areas (ACOPAC, ACOSA, and ACLA-P) and four counties (Osa, Aguirre,
Perez Zeledon, and Dota).
54
Figure 2.6 Land-use types in the Path of the Tapir Biological Corridor in 2000, Costa
Rica. In this black & white version dark colors represent forested habitats and light colors
human-modified land-use types.
55
250
Biological Reserve
Wildlife Refuge
200 Forest Edge
Forest Fallow
Tree Plantation
150
Number of trees
100
50
0
5-10 15-20 25-30 35-40 45-50 55-60 65-70 75-80 85-90 115-120 130-135 160-165 245-250
Diameter (cm)
Figure 2.7 Diameter size class distribution of tree species located in five land-use types in
the Path of the Tapir Biological Corridor, Costa Rica
56
200 Biological Reserve
Wildlife Refuge
180 Forest Edge
Forest Fallow
160
Tree Plantations
140
Number of Trees
120
100
80
60
40
20
0
0-5 5-10 10-15 15-20 20-25 25-30 30-35
Height (m)
Figure 2.8. Total height size class distribution of tree species located in five land-use
types in the Path of the Tapir Biological Corridor, Costa Rica
57
Chapter 3
Avian Diversity of Ten Different Land-use Types in a Human-Modified Biological

Corridor in Costa Rica: Implications for Land-Use Planning
Abstract
As a result of the limitations of current protected area systems and forested areas
in providing habitat for most wildlife species, in the last two decades efforts have shifted
to studying wildlife conservation in human-modified landscapes. The present study was
carried out in the Path of the Tapir Biological Corridor, Costa Rica. The corridor, an
82,000 ha area of fragmented forests, encompasses 55 rural communities with more than
10,000 people, and is considered one of the most diverse regions on the Pacific Coast of
Central America. However, deforestation and development are threatening this high
biodiversity. The main objective of this chapter is to describe the contribution to bird
biodiversity conservation of ten different habitat types in the corridor: wildlife refuges,
biological reserves, forest edges, forest fallows, tree plantations, agrosilvopastoral
systems, homegardens, oil palm plantations, villages, and residential tourism projects.
Bird data were obtained using point counts during a two-year period. Each habitat type
had 20 sampling points (200 total), and each point was visited six times during dry
seasons and six times during rainy seasons. Overall, 44, 917 birds from 48 families and
334 species were identified in this study. Forested land-use types had the highest number
of bird species (81%). Very important, 77% of the birds registered were also found in the
human-modified land-use types. In general, it is concluded that forested habitats should
be the priority areas for conservation at the landscape level because they provide the main
habitat for the most species. Moreover, in human-modified landscapes forest fallows,
58
agroforestry systems, and native tree plantations can be incorporated in land-use planning
strategies to provide additional habitat to forest-dependent and non-forest depend species,
and connectivity to the otherwise isolated and fragmented forests.
Introduction
Destruction or alteration of natural habitat because of land-use change represents
one of the most serious threats to biodiversity worldwide (Groom et al. 2005) along with
climate change and invasive species (Sala et al. 2000). As a result of this loss of natural
habitats, maintaining and conserving biodiversity in fragmented landscapes now
dominated by human land use is a challenge (Bennett 2003; Chazdon et al. 2009a).
Assessing the conservation potential of human-dominated landscapes requires
investigation of the activities, movement, and persistence of species not only in remnants
of native habitats but also in the full array of countryside habitats (Saunders et al. 1991;
Daily 2001; Hughes et al. 2002; Chazdon et al. 2009a), known also as the matrix. The
matrix can be defined as the most widespread habitat within which other elements are
embedded, and it can be either the original habitat type (e.g., a primary forest now
surrounded by a matrix of agroforestry) or a modified one (e.g., an agricultural plantation
surrounded by a matrix of pristine forest) (Anderson and Jenkins 2006). In
anthropogenically modified landscapes, some or all of the matrix may be human-
modified communities such as agricultural fields of various types, agroforestry systems,
clear-cut forests, pastures, tree plantations, fallow land, and towns (Daily et al. 2003;
Hilty et al. 2006).
59
Conserving biodiversity should incorporate human-modified landscapes in
assessments of biodiversity and ecosystem services, planning of corridors, establishment
of buffer zones, and restoration of degraded lands (Chazdon et al. 2009a). Understanding
the contribution of the different land-use types within a landscape will be important for
land-use planning, especially in areas with high biodiversity concerns and high human
impact.
The present study was carried out in the fragmented and human-modified Path of
the Tapir Biological Corridor in Costa Rica (PTBC), which is one of the most diverse
areas of Central America (TNC and ASANA 2000), but whose biodiversity is at risk
because of deforestation and development. Current land use in the corridor has been the
result of local and national settlement and development policies (Newcomer 2006;
Redondo-Brenes 2007). Between 1940 and 1960, Costa Ricans migrated from the
northern part of the country toward this region searching for new lands to clear and
cultivate. Between 1960 and 1990, an expanding cattle industry was a major contributing
factor to deforestation (TNC and ASANA 2000). Recently, deforestation rates have
decreased, forest cover has increased as a result of pasture abandonment, and
conservation efforts are being undertaken through the establishment of a National Park,
five National Wildlife Refuges, and 27 Informal Private Nature Reserves (Redondo-
Brenes 2007). However, as land prices have increased, land ownership is shifting from
local farmers to wealthy foreigners; and infrastructure development is growing
uncontrolled. These changes pose new challenges for the region, since protected areas are
interspersed with private lands (Redondo-Brenes 2007).
60
The main goal of this study was to assess the contribution of ten different habitat
types to the conservation of bird species in the PTBC. The ten land-use types were:
biological reserves, wildlife refuges, homegardens, agrosilvopastoral systems, forest
edges, forest fallows, tree plantations, oil palm plantations, local villages, and residential
tourism projects. This information will be important for assessing the current status of
bird species in the corridor as well as the conservation value of the different habitat types
for land-use planning strategies.
Methods
Study Area Description: Path of the Tapir Biological Corridor (PTBC)
The PTBC is located in southwestern Costa Rica (Figure 3.1) and is a part of the
multinational Mesoamerican Biological Corridor (MBC). The corridor’s main objective
is to create a network of sites favorable to fauna and flora between the forests of the Osa
Peninsula and Golfo Dulce, including Corcovado National Park. These forests connect
with those located in the Los Santos Forest Reserve in the Talamanca Mountain Range.
The PTBC area covers approximately 82,000 ha and is inhabited by 10,000 people in 55
rural communities. This mosaic of human-dominated land uses includes primary and
secondary forests, native and exotic forest plantations, agriculture, agroforestry systems,
bamboo, oil palm plantations, ecotourism projects, and others.
The corridor is composed of terrain ranging in altitude from 0 to 1100 meters
above sea level (TNC and ASANA 2000). According to the Holdridge System (1967),
three life zones are represented within it: (1) The Tropical Wet Forest that comprises the
entire length of the corridor, especially the lower zones; (2) The Tropical Wet Forest
61
Transition to Premontane, represented by a small fringe at the intermediate altitudes, and
(3) The Tropical Rainforest Premontane found at the highest elevations in the corridor.
The climate is categorized as Seasonal Tropical Wet with an annual precipitation of 4,239
mm and average temperature of 27 ºC. There is high precipitation during the rainy season
with as much as 685 mm per month (maximum of 1,715 in October 1988), and a
pronounced 4- to 5- month dry season with little rain (less than 100 mm) from January to
May (TNC and ASANA 2000). The soils in the corridor are low in fertility, and are
generally classified as Ultisols and Inceptisols. There are more than 30 rivers among
which the main ones are Coronado, Higueron, Uvita, Baru, Hatillo, Savegre, and Guabo
(TNC and ASANA 2000).
PTBC is one of Costa Rica’s most diverse regions. A total of 2700 plant, 173
mammal, 51 amphibian, and 81 reptile species have been identified in the corridor (TNC
and ASANA 2000). Moreover, 500 avian species have been registered in the area in the
six most recent annual bird counts. The PTBC provides habitat to some endangered
animal species such as margay (Leopardus wiedii), ocelot (Leopardus pardalis), jaguar
(Panthera onca), spider monkey (Ateles geoffroyi), squirrel monkey (Saimiri oerstedii),
scarlet macaw (Ara macao), great curassow (Crax rubra), and endemic tree species such
as quira (Caryodaphnopsis burgeri).
Description of the Habitats Used for Evaluation of Bird Abundance and Diversity
Based on the predominant land uses currently present in the PTBC, ten different
habitat types, located up to 500 m.a.s.l., were selected. The studied habitat types were
selected based on accessibility, availability, and importance within the corridor landscape
62
among others. In the case of the wildlife refuges and biological reserves, they were also
chosen based on their importance as core areas for wildlife conservation within the
corridor, as well as owner’s permission to get access to the properties. A complete land-
use type description can be found in Chapter (2).
(a) Wildlife refuges: Forested areas (primary or old secondary forest) were surveyed
within two refuges: Rancho La Merced and Hacienda Baru (10 sampling points at each
refuge for a total of 20 points for this type of habitat). La Merced (329 ha) and Baru (330
ha) are private reserves and have had governmental protection since 1995. Average basal
area at both refuges was 55 m2/ha and volume 620 m3/ha. In addition, 76 woody species
from 31 families with diameters larger than 5 cm were identified in 0.5 ha sampled area.
The tallest canopy trees were approximately 32 m.
(b) Biological reserves: Two biological reserves were surveyed: La Cusinga and Oro
Verde (10 sampling points within each refuge). La Cusinga (250 ha) and Oro Verde (150
ha) are private reserves without governmental incentives or protection. Average basal
area at both reserves was 41 m2/ha and volume 476 m3/ha. Additionally, 99 woody
species from 39 families with diameters larger than 5 cm were found in 0.5 ha sampled
area. The tallest canopy trees were approximately 35 m.
(c) Forest edge: Twenty sampling points were located in areas along the edges of
primary and old-secondary forests. Average basal area for forest edge stands was 39
m2/ha and volume 386 m3/ha; 93 woody species from 35 families with diameters larger
than 5 cm were identified in 0.5 ha sampled. The tallest canopy trees were approximately
28 m.
63
(d) Forest fallows (young secondary forest): These were areas of forest
regenerating in recently abandoned pasture lands (1-5 years after abandonment). Average
basal area for forest fallow stands was 7 m2/ha and volume 31 m3/ha. Moreover, 42
woody species from 28 families with diameters larger than 5 cm were identified in 0.5 ha
sampled. The tallest canopy trees were approximately 13-15 m, with a few remnants that
could reach 20 m.
(e) Homegardens: These were traditional Costa Rican homegardens composed of a
combination of annual/perennial crops, ornamental plants, and timber and fruit trees. By
assessing the area around a radius of 20 meters from the sampling point center, a total of
65 plant species from 34 families were identified within this habitat.
(f) Agrosilvopastoral systems: Most of the pastures in the region are used to raise
beef cattle and are mostly planted with exotic pasture species. All of the selected sites had
either living fences or scattered trees that provided perches or food sources for birds. Using
line transects in 10 out of 20 sample points, a total of 41 different woody species (28
families) larger than 5 cm of diameter were identified, either as living fences or scattered
trees.
(g) Tree plantations: Most tree plantations in the PTBC consist of teak (Tectona
grandis) or amarillon (Terminalia amazonia). Teak is a valuable exotic timber species
(native to Southest Asia) that is exported to Europe and the United States. Amarillon is
native to Costa Rica and is one of the most promising species for timber production in the
country. Since we did not get access to enough T. amazonia plantations to set up 20
sampling points, 10 points were located in teak and 10 in amarillon plantations in order to
have a total of 20 points for this habitat type. Plantation sizes varied from 2 ha to 30 ha
64
with teak plantations being the largest. Tree diameters varied from 9 cm to 30.5 cm and
height from 12 m to 22.5 m, with the teak trees being the largest overall.
(h) Oil palm plantations: In the northern region of the corridor, there were hundreds
of ha of oil palm plantations (Elaeis guineensis), which is an exotic crop to Costa Rica.
The sampling points were located in a range from recently established plantations (1.5 m
tall) to mature plantations (up to 10 m height).
(i) Villages: Two main areas were selected for sampling. The first one was the
largest village within the corridor, Puerto Cortez, where 10 sampling points were
selected. The other 10 sampling points were located in the small villages along the main
highway. A total of 57 plant species from 34 families were identified within the sampling
points.
(j) Residential tourism projects: Because the PTBC runs along the Central and
South Pacific Coast of the country, there has been an exponential growth in the number
of foreigners building summer or retirement houses in the area in the last 10 years. The
sampling points were located in areas where there was more than one house built. A total
of 47 plant species from 33 families were identified within the sampling points.
Bird Surveys
Point counts were used because they allow one to develop relative indices of
abundance and inferences about bird-habitat relations (Buckland et al. 1993; Bibby et al.
2000; Blake and Loiselle 2001; Sutherland 2006; Barlow et al. 2007; Farwing et al.
2008). The primary advantages of point counts are that the relative abundance of many
species can be determined over broad areas at a moderately low cost and that species-
65
habitat relations can be evaluated effectively compared to other methods (Bibby et al.
2000; Sutherland 2006). For the present study, 20 point counts were located within each
habitat site in 16 sampling routes. Each route was selected based on previous visits to the
area, in an attempt to minimize cost. The minimum and maximum number of sampling
points per route were ten in forested habitats and 15 in more open-habitats. Average
minimum distance in between points was 200 m in forested habitats, and 250 m in open
areas to avoid double-counting the same individual (Bibby et al. 2000; Sutherland 2006).
Each point was surveyed three times in the dry and three times during the rainy
seasons over a two year period (12 visits per sampling point total). Dry season surveys
were conducted from mid-January to mid-April and rainy season surveys from mid-June
to mid-August. At each sampling point, every bird species that was seen or heard within a
50m radius plot and during a period of 10 minutes was recorded and identified. All
surveys were conducted from sunrise to 9:15 am during clear days, avoiding windy
situations. The daily order of point sampling in a route was changed so that each point
was visited once in the first hour, once in the second hour, and once in the third hour of
the season sample period. Field identification was conducted using the Manual of Birds
of Costa Rica (Stiles & Skutch 1989) and the Birds of Costa Rica (Garrigues & Dean
2007), as well as an identification of their calls and songs. All bird identifications were
done by A. Redondo-Brenes. Total observation time was 400 hours for the whole study
(2 hours/point).
66
Data Analysis
For each habitat type the total abundance and species richness of birds were
calculated by combining data from the two sampling periods (dry and rainy) as well as by
comparing between the two years considered for this study. Shannon diversity index,
rarefaction curves, Sorensen similarity index, and the number of shared species using
EstimateS 5.0 were computed (Colwell 2005). Rarefaction curves were plotted against
the number of individuals observed, using the mean of the four commonly employed
abundance-based estimators (ACE, CHAO1, JACK1, and BOOSTRAP; Barlow et al.
2007). Abundance, species richness, and diversity among the ten land-use types were
compared, using analysis of variance and/or Kruskal Wallis tests. Cluster analysis was
also used to group habitat types based on species composition. All statistical analyses
were conducted in PAST v 1.95 (Hammer et al. 2001) and SAS v. 9.2.
Results
Bird Diversity
A total of 44,917 birds, 48 families, and 334 bird species were identified over the
two-year period of this study (Appendix 1). From the total number of observed birds,
21,025 individuals (293 species) were recorded in 2008 and 23,902 (310 species) were
recorded in 2009. As expected, there were differences in the accumulated total number of
species found during dry (307 species) and rainy time (271 species) (Figure 3.2).
However, even though the total difference across both seasons was 36 species, there were
63 species that were only recorded over dry surveys and 27 species only recorded during
rainy surveys. From the 27 species found only in the rainy, 26 species are considered rare
or uncommon species. In fact, only the Dendrocygna autumnalis had abundance greater
67
than 5 individuals for the entire study (73 birds). On the other hand, of the 63 species
only found in the dry period, 46 species (73 %) were neotropical migratory species and
17 species were resident species. Migratory species accounted for 13.7% of the total bird
diversity registered in the PTBC.
Comparing overall abundance at the species level, it was found that while 20
species had more than 50% of the entire records for this study, there were 135 species
(40% of all species) whose sightings averaged less than one for each of the 12 surveys
conducted. Moreover, there were 57 species (17%) that had just one or two individuals
recorded in 2 years. The most common species for this study were Ramphocelus
costaricensis, Quiscalus mexicanus, Sporophila americana, Thraupis episcopus,
Troglodytes aedon, Pitangus sulphuratus, Volatinia jacarina, Brotogeris jugularis,
Tyrannus melancholicus, and Thraupis palmarum. Of the total birds recorded for this
study, these species comprised from 4.7% to 2.5% of the whole abundance. Quiscalus
mexicanus had a population that varied from 743 birds in 2008 to 1001 birds in 2009 - a
34% change.
Bird abundance across the 10 habitat types
At the family level, data among the ten habitat types varied from 39 bird families
in wildlife refuges/homegardens to 31 families in the oil palm plantations. Although this
difference across land-use types is only eight families, the significant difference is at the
species composition level. Overall, five species account for 50 % or more of the total
number of individuals per land-use type (Figure 3.3). Twenty two families were
68
observed in all land-use types and the most abundant were Thraupidae (14% of the total),
Tyrannidae (13%), Emberizidae (8%), Psittacidae (7.4%), and Columbidae (7%).
At the landscape level, homegardens, agrosilvopastoral systems, and villages had
the highest bird abundance; tree plantations had the lowest (p < 0.05; Table 3.1). Forested
habitats (i.e. biological reserves, wildlife refuges, forest edges) also had low bird density
values. Moreover, whereas forest edges, forest fallows, homegardens, biological reserves,
and wildlife refuges were the sites with the highest species richness and diversity of
species, tree plantations, residential tourism, oil palm plantations, and villages had the
lowest diversity values within the biological corridor. The latter two were statistically the
least diverse sites of the ten land-use types assessed (p < 0.05).
Based on the 2-year accumulated data, rarefaction curves confirmed some of the
diversity patterns described above (Figure 3.4). In summary, forest edges, forest fallows,
biological reserves, and wildlife refuges are expected to accumulate the largest number of
species, followed by homegardens, tree plantations, and agrosilvopastoral systems. By
contrast, residential tourism projects, oil palm plantations, and villages seemed to have
the lowest expected number of species.
Species composition also varied across the ten habitat types in the corridor.
Biological reserves, wildlife refuges, and forest edges had the most similar assemblages
of species (Table 3.2, Figure 3.5). This may be because these habitats were dominated by
primary and old-secondary forests, having a majority of forest-specialist bird species
(Redondo-Brenes and Montagnini 2010; see also Chapters 2 & 4). Also, tree plantations
seemed to share many species with these three habitat types. Agrosilvopastoral systems,
homegardens, residential tourism projects, and oil palm plantations had similar
69
assemblage of species because they were dominated by generalist and open area bird
species (Chapter 4). In contrast to these patterns of high similarity, more open area
habitats (i.e. villages, oil palm plantations, residential tourism projects) tended to have the
lowest similarity indexes in relation to forested biological reserves, wildlife refuges, and
even forest edges.
At the species level, of the 334 species recorded, only 10% of these species (33
species) were found in all ten habitat types (Appendix 1), confirming the highly distinct
bird assemblages across the land-use types. The majority of these species (26 species –
76%) corresponded to resident species, and only seven were migratory species:
Empidonax flaviventris, Piranga rubra, Legatus leucophaius, Dendroica pensylvanica,
Vermicola peregrina, Vireo philadelphicus, and Catharus ustulatus. Contrary to the
adaptation capability of most of these species to different habitat types, there were 61
species (18% of the total diversity) that were only found in one particular land-use type.
There were nine in biological reserves, eight in forest edges and agrosilvopastoral
systems, seven in forest fallows and homegardens, six in wildlife refuges, villages and oil
plantations, and only two in residential tourism projects and tree plantations.
Finally, there were 270 species (81%) sighted in forested habitats, including
biological reserves, wildlife refuges, forest fallows, and forest edges. Also very
importantly, 77% of the total species observed (257 species) were recorded within the six
human-modified land-use types. Furthermore, 66 species (18%) were only found in
human-modified landscapes and 77 species (23%) were only recorded in forested
habitats.
70
Discussion
Bird species identified as part of this study corresponded to almost 40% of the
avian diversity in the country and more than 66% of the expected bird diversity in the
region. Most of these bird species were located in forested habitats; however, the human-
modified landscapes provided habitat to more than 77% of the species found during the
2-year study. This 77% is a value higher than other studies that estimated the contribution
of countryside land uses to landscape biodiversity (Hughes et al. 2002). Because of a
generalized interest in understanding the contribution of human-modified land-use types
to biodiversity conservation (Daily et al. 2003; Chazdon et al. 2009a; Gardner et al.
2009), the following section will discuss the contribution of these habitats to avian
diversity. It will focus on the conservation value these habitats shared with forested
habitat, but understanding that human-modified land use types are also very important for
the overall biodiversity of a region, as these landscapes provide habitat and food sources
to most open-area specialists and to many forest-generalist species.
Secondary forests
In the present study, forest fallow was the human-modified land-use type with the
highest bird diversity and also the greatest bird species similarity with forested habitats.
The young secondary forests (forest fallows) shared 143, 106, and 109 species with forest
edges, wildlife refuges, and biological reserves, respectively (Table 3.2). Secondary
forests may play an important role in biodiversity conservation by increasing regional-
scale species richness at the landscape level (Hughes et al. 2002; Barlow et al. 2007,
Chazdon et al. 2009b), improving connectivity across the landscape, and buffering
71
existing forest fragments (Fischer et al. 2006; Chazdon et al., 2009a). In the PTBC,
young secondary forests are mostly growing in former pasture and agricultural
landscapes surrounding most of the private reserves.
Secondary forests are rapidly expanding across tropical landscapes (Barlow et al.
2007). FAO (2007) reported that the area of secondary forests throughout the world was
increasing dramatically and that in many tropical countries they exceeded areas covered
by primary vegetation. In Costa Rica, there were approximately 400 000 to 450 000 ha of
secondary forests in different successional stages in the 1990s (CCT 1991, Segura et al.
1997, Müller and Solís 1997). This area is twice as large as the area of primary forests for
production objectives and three times the size of tree plantations in the country (Berti
2001). The area under secondary succession has continued to increase from the 1990’s to
date, with a gain in forest cover from 42% in 1997 to 51% in 2005 (FAO 2007). A similar
pattern was found in the PTBC. At present, around 17% of area in the PTBC is under
forest fallows (Redondo-Brenes 2007).
Even though forest fallows can provide habitats for many bird species, regaining
the complex microhabitats and structures required by primary forest specialists is likely
to take centuries (Chazdon 2003; Montagnini 2008; Chazdon et al. 2009b). Dunn (2004)
found that species richness of many taxa may recover relatively rapidly in secondary
forests, but species composition can take longer, especially for ant and bird species.
When these secondary forests are acting as buffer zones or are located near primary
forests, bird species richness can be even higher (Blake and Louis 2001). Primary forest
bird species usually are not found in young secondary forests, limiting the value of these
forests as a conservation tool (Dunn 2004; Barlow et al. 2007). In the current study, 67
72
forest specialist species were recorded in the forest fallows (See Chapter 4), being the
highest of all the human-modified land uses.
Tree plantations
Tree plantations can also provide suitable habitats for many bird species.
However, as in young secondary forests, there is a limited number of forest-dependent
species that usually inhabit them (Carnus et al. 2006; Barlow et al. 2007; Brockerhoff et
al. 2008). In the present study, tree plantations shared 97, 96, 125 species with wildlife
refuges, biological reserves, and forest edges, respectively (Table 3.2). Tree plantations
had 55 forest-specialist species (Chapter 4). These values are lower than those from forest
fallows, but the second highest for the seven human-modified habitats in this study.
Tree plantations can enhance local biodiversity by promoting the regeneration of
understory species through shading out grasses, increasing nutrient richness of topsoil,
allowing the growth of more sensitive tree species, and creating a microclimate that
attracts seed dispersers (Cusack and Montagnini 2004; Montagnini 2010). In addition,
tree plantations may serve as perches, food sources and habitat for a number of wildlife
species, thus contributing to an increase in overall biodiversity (Slocum and Horvitz
2000; Slocum 2001; Carnus et al. 2006).
Use of native species, especially in mixtures, may increase wildlife populations in
former degraded landscapes (Lamb and Gilmour 2003). Recent studies conducted in
experimental plantations of several native species in Costa Rica have found abundant and
species-diverse understories in these plantations (Orozco and Montagnini 2007; Butler et
al. 2008; Leopold and Salazar 2008). This understory may provide habitat and other
73
resources to bird and other wildlife species (Carnus et al. 2006; Butler et al. 2008). For
the current study, whereas all T. amazonia plantations had understory vegetation, eight
out of ten T. grandis plantations lacked this vegetation strata because of intensive
silviculture management. Bird abundance and species richness was higher in the native T.
amazonia (1501 birds, 123 species) than in the exotic T. grandis (1278 birds, 104
species).
For bird species, several studies have compared bird diversity in natural forests
and tree plantations. In Kenya, it was found that species richness values were statistically
higher in primary forests than in monoculture tree plantations, but these were not
statistically different between primary forests and mixed-tree plantations (Farwig et al.
2008). In the north-eastern Brazilian Amazonia, species richness of primary forests was
higher than Eucalyptus plantations, with understory insectivores and canopy frugivores
being the main primary forest species absent from tree plantations (Barlow et al. 2007). A
similar pattern with the frugivorous species was found when comparing young Tectona
grandis (teak) plantations in Panama with native forest fragments (Perla et al. 2002).
Frugivorous species were found in small and large forest fragments, but were absent in
teak plantations. Despite these differences in species composition between tree
plantations and primary forests, when tree plantations can replace other human-modified
ecosystems (e.g. degraded pastures) they will almost always support a greater native
biodiversity (Brockerhoff et al. 2008).
74
Agrosilvopastoral
According to recent statistics, pastures and agrosilvopastoral land represent 77%
of the total 500 million ha of agricultural land in the Neotropics (Amezquita et al. 2005).
About 30% of these pastures are on poor, acidic soils (Montagnini 2008), as is the case in
the present study. Agrosilvopastoral systems that involve the combination of trees with
pastures and livestock offer an alternative set of cattle production systems. These systems
are more complex than grass monocultures and are classified based on the functions and
configuration or structure of trees within the system; examples are dispersed trees in
pastures, live fences in pastures, fodder banks, tree alley pasture systems, and pastures
with windbreaks (Pezo and Ibrahim 1999). Because of their increased complexity relative
to grass monoculture systems, agrosilvopastoral practices also have important benefits for
biodiversity (Harvey et al. 2005; 2006; 2008; Francesconi et al. 2010).
Many tropical pastures contain isolated trees that may be remnants of the original
forest. They are conserved by farmers for their shade for animals and humans and as
sources of firewood, timber and fruit. These trees and shrubs may be important in
attracting birds, in comparison with treeless grass monocultures. Crown size and shape,
tree height, fruit production and other characteristics may influence the ability of trees to
attract birds (Slocum and Horvitz 2000; Slocum 2001). Generally, it is thought that birds
prefer taller trees as perches to have a better view of the surroundings and to avert
potential predators. On the other hand, results of a recent study in the Pacific region of
Costa Rica suggested that crown diameter was more important than height of trees in
attracting birds (Slocum and Horvitz 2000).
75
As suggested above, a combination of several characteristics may determine the
role of each tree species in attracting birds to trees in agrosilvopastoral systems. Distance
to forest patches and position in the matrix can also have an important influence. For
example, to evaluate the contribution of living fences to improving the connectivity of the
agricultural matrix, Francesconi et al. (2010) examined bird species composition (forest
specialists, open-area specialists, and generalist species) that use living fences as habitat
in Esparza, in the Pacific lowlands of Costa Rica. Bird species composition changed as a
function of distance to the forest patch; yet species richness, number of individual birds,
and Shannon Diversity indices were similar between forest interiors (control) and living
fences. Their results suggest that living fences near forest patches provide habitats to
many bird species including forest specialists. However, fence structure and composition
significantly influenced usage by birds. The authors concluded that the presence of
diverse native tree species in fences and increased vegetative cover may counteract the
effect of distance to the forest patch, promoting greater bird species diversity in living
fences and in the landscape (Francesconi et al. 2010).
As seen, the specific types of agrosilvopastoral practices adopted by farmers are
important for the conservation of biodiversity (Enriquez et al. 2007; Saenz et al. 2007). In
a long term project in landscapes dominated by cattle in Esparza (Costa Rica), Matiguas
(Nicaragua), and Quindio (Colombia), intensive monitoring of birds in different land use
systems over four years showed that the agrosilvopastoral practices with high tree
densities in pastures and multi-strata or permanent live fences had significantly higher
abundance and species richness of birds than degraded and grass monoculture pastures,
with comparable values to those of forested vegetation (Ibrahim et al. 2010). In their
76
study, Ibrahim et al. (2010) reported that 111 bird species were observed in the landscape
of Esparza, Costa Rica, 170 species in Quindio, Colombia, and 154 species in Matiguas,
Nicaragua. Of these, a total of 60.5%, 54% and 64% of the species inventoried were
dependent on the forest in Costa Rica, Colombia, and Nicaragua, respectively. Likewise,
in a fragmented agricultural landscape in Cañas, Costa Rica, Cardenas et al. (2003)
showed that the agrosilvopastoral systems with high tree densities and permanent live
fences (multi-strata fences) had more than 40 bird species, whereas grass monoculture
pastures had 28 bird species. For the current study, agrosilvopastoral systems had one of
the highest species richness values in the corridor, and they also shared 75, 78, and 110
bird species with biological reserves, wildlife refuges, and forest edges, respectively.
However, only 34 forest-dependent species were found in these habitats, representing
20% of the total. This percentage is low in comparison to the values showed above from
other regions of the Neotropics.
Homegardens
The homegardens, which provide the household with a basic food source as well
as high value products to generate cash, are important in the tropics and are often used as
tools in development projects that promote food security, especially in the poorest areas
(FAO 2001; Montagnini 2006). In several regions of Costa Rica, homegardens are
important for supplying food (Zaldivar et al. 2002); they also serve as a buffer in times of
harvest failure or economic depression (Price 1989).
Homegardens may have positive effects on biodiversity because they can serve as
local refuges for plants and animals that otherwise may be threatened by human or
77
natural disturbances (Montagnini 2006). For example, Griffith (2000) reported that
during the 1998 fires in Petén, Guatemala, homegardens and other agroforestry systems
may have served as critical refuges during a habitat bottleneck for many forest species.
Apparently, agroforestry farms attracted birds by virtue of their complex structure –
similar to that of intact forest patches – and they harbor insects, provide nesting sites, and
offer protection from predators (Griffith 2000). Birds were also attracted by the cultivated
fruit trees, which may have provided some of the only food sources in the region after
fire destroyed most of the surrounding vegetation. Homegardens tend to attract many bird
species of different guilds (Scales and Marsden 2008). For the present study,
homegardens were one of the land uses with the highest accumulated number of bird
species (185 species). They shared 91, 92, and 126 species with wildlife refuges,
biological reserves, and forest edges, respectively, having the second largest number of
forest dependent species (51 species). However, they are disappearing as local people are
selling their properties to developers, who usually replace the complex structure of
native/exotic plants with a simpler one based on exotic species. Also, the area under
construction has increased, especially because most of the new residential areas request
commodities such as swimming pools.
Oil palm
As a result of the rising demand for vegetable oils and biofuels, oil palm (Elaeis
guineensis) is one of the world’s most rapidly increasing crops (Fitzherbert et al. 2008),
becoming a major threat to tropical biodiversity (Turner et al. 2008; Koh and Wilcove
2008). At present, oil palm is grown across more than 13.5 million ha of tropical regions
78
(Fitzherbert et al. 2008). In Costa Rica, the oil palm industry is relatively small, but is
one of the oldest in tropical America. The first commercial plantation was established in
1944. In 2006, the industry covered about 47,807 ha, mostly established on the central
and southern parts of the Pacific coast (96%) (Escobar and Peralta 2007).
According to Fitzherbert et al. (2008), there were only 13 published scientific
papers comparing wildlife diversity in oil palm plantations to that in forests from 1970 to
2006. Overall, oil palm plantations support many fewer species than do forests and often
fewer than other tree crops as well (Fitzherbert et al. 2008). Moreover, species
composition is different between these two habitat types. While most species with
specialized diets and microhabitat were absent from oil plantations, plantation
assemblages were dominated by a few abundant generalists, open-area species, invasive
species and pests (Aratrakorn et al. 2006; Peh et al. 2006; Chey et al. 2006; Fitzherbert et
al. 2008). Oil palm plantations also have lower species richness than do other agricultural
types (Fitzherbert et al. 2008). For the present study, and with the exception of the
assessed villages, oil palm plantations had the lowest diversity values, sharing only
approximately 50 species with the forested habitats, and registering only 22 forest-
specialist species.
As a result of the low conservation value found in oil palm plantations, several
authors have suggested two main approaches for enhancing biodiversity conservation at
the landscape level. The most commonly cited is to preserve as much of the remaining
natural forests by either protecting forest in reserves, buffer zones around plantations, or
forest fragments within the plantations and at the landscape level (Fitzherbert et al. 2008;
Koh 2008; Turner et al. 2008). Conversely, other researchers advocate for the
79
implementation of traditional agroforestry practices to create a more heterogeneous
landscape benefiting both biodiversity and rural communities (Bhagwat and Willis 2008;
Koh et al. 2009). Claims of using agroforestry systems focus on the oil palm plantations
lower species richness values for some taxa in comparison with agroforestry systems
based on rubber plantations (Hevea brasiliensis), cocoa (Theobroma cacao), and coffee
(Coffea canephora) in Southeast Asia (Bhagwat and Willis 2008; Koh et al. 2009). Most
of the sampling points for this study had more than just palm vegetation including living
fences, and remnant trees, among others. In fact, more than 43% of birds found in the oil
palm plantations were registered when utilizing such microhabitats (Chapter 4).
Urbanization
Forest loss with accompanying urbanization is often permanent as forests are
replaced by concrete structures or surfaces. The remaining forest patches are usually
isolated and too small to maintain viable populations of forest-dependent wildlife species
(Er et al. 2005). The effect of urbanization on wildlife conservation can be immense;
however, our understanding is still rudimentary according to some reviews (McKinney
2002; Chace and Walsh 2006). For birds, urban development can affect native species in
many ways because human-influenced changes in the environment often result in
alterations to multiple elements of their habitats (McCaffrey 2009). Therefore, because
birds respond to vegetation composition and structure, urban areas that retain native
vegetation features retain more native species than those that do not (Chace and Walsh
2009).
80
How an individual species is affected by these changes depends on both the
specific habitat features they require as well as the scale at which they respond to habitat
changes (McCaffrey 09). Marzluff and Ewing (2001) and Chace and Walsh (2006)
summarize the factors determining which species can coexist with human settlement as
follows: (1) the presence and patch size of remnant (native) vegetation (Catterall et al.
1991; McKinney 2002; Er et al. 2005; Suarez-Rubio and Thomlinson 2009), (2)
competition with exotic species (Major et al. 1996; McKinney 2002; Er et al. 2005), (3)
non-native predators (Churcher and Lawton 1987; Paton 1990), (4) the structure and
floristic attributes of planted vegetation (Suarez-Rubio and Thomlinson 2009), (5)
supplementary feeding by humans (Brittingham 1990; Major et al. 1996), (6) obstruction
of traditional migratory and dispersal routes and increasing the dangers of dispersal
(Marzluff and Ewing 2001), and (7) residual pesticides (Major et al. 1996).
Overall, as the urbanization processes increase, there is an increase in avian
biomass and induced community homogenization, but a reduction in richness (Chance
and Walsh 2006; Devictor et al. 2007). Some studies have shown that the urbanization
matrix is correlated negatively with native species (Er et al 2005; Chace and Walsh 2006;
Suarez-Rubio and Thomlinson 2009), especially endemic species (Suarez-Rubio and
Thomlinson 2009), and forest-specialist species (Er et al. 2005), but positively with
introduced species (Suarez-Rubio and Thomlinson 2009). Urbanization tends to select for
omnivorous, granivorous, and cavity nesting species (Chace and Walsh 2006). Villages
and residential tourism projects were the two land-use types where cement and concrete
were part of the landscape. However, residential tourism projects tended to keep larger
forest cover around the houses in relation to the selected 20 sampling points in villages
81
and that may explain why the former had 33 forest-specialist species and the latter only
19 species (the lowest of all land-use types). Also, shared species with forested habitats
were higher for residential tourism projects (70 - 101 species) than for villages (52 – 78
species). Land-use change from forested habitats to any residential project should be
avoided in areas where infrastructure may limit the connectivity among forest reserves.
Conclusions: Implications for land-use planning
This study was carried out in a high biodiverse human-modified biological
corridor that is under threat because of residential and tourism development, and whose
main goal is to provide connectivity to the forested reserves. Therefore, management of
the different land-use types as well as their components may have positive or negative
biodiversity implications and should be taken into consideration in land-use planning.
It is important to stress that biodiversity conservation should always include
primary and old-secondary forests (e.g. those forests located in wildlife refuges,
biological reserves, or isolated forest fragments) because the majority of bird species
utilize these habitats the most. Moreover, these are the most important habitat types for
forest-dependent wildlife at the landscape level, and these species are usually more
susceptible to fragmentation and land-use change. However, the results of this research
also show that the matrix around these forests, especially young secondary forests,
homegardens, agrosilvopastoral systems, and tree plantations, are important in expanding
the potential habitat for many bird species in a biological corridor, including both forest
and non-forest species.
82
Agroforestry systems such as agrosilvopastoral systems and homegardens are
diverse habitats that, if managed in the proper way, may increase the current contribution
of the PTBC and other fragmented habitats to wildlife conservation. Adding native tree
species, especially fruit trees, in agricultural systems adjacent to forested areas may add
conservation value to these habitat types, attracting endangered bird species such as Ara
macaw and other forest-dependent bird species (Chapter 4 and 5).
According to the data in this study, it may even be possible to implement
agrosilvopastoral systems that are compatible with production objectives and biodiversity
conservation. Good farm planning is needed, including a mosaic of land uses that are
compatible with both the conservation of biodiversity and production objectives (Saenz et
al. 2007; Harvey et al. 2008). For example, cattle farms could incorporate land uses
which include establishment of fodder banks, multi-strata live fences, and pastures with
high tree cover or density, as well as riparian and secondary forest (Ibahim et al. 2010).
Planting native tree species, such as Terminalia amazonia and other native tree
species of interest in Costa Rica and elsewhere in the Neotropics, could be given priority
to serve biodiversity and economic objectives of humans inhabiting the corridor.
Finally, forest fragmentation, unplanned urbanization, monocultures, and the
introduction of exotic plant species could be avoided in biological corridors in order to
avoid jeopardizing native biodiversity. One example of these monocultures that could be
avoided is oil palm plantations. Even though more than 100 species were found in the oil
palm plantations, their species richness value was one of the lowest of all evaluated
habitats. Furthermore, the worldwide expansion of this type of plantation is cited as a
major threat to tropical biodiversity.
83
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Table 3.1. Abundance, species richness, and diversity of birds registered in ten land-
use types in the Path of the Tapir Biological Corridor, Costa Rica
Habitat Type Abundance Species Richness Shannon

(n/census) (species/census) Index
Homegardens 574 (13.2)a 89 (3.3)a 3.88 (0.04)ab
Agrosilvopastoral Systems 533 (13.8)a 82 (2.9)ab 3.84 (0.04)b
Villages 500 (15.9)a 57 (1.9)c 3.35 (0.02)d
Oil Palm 361 (9.2)b 55 (1.6)c 3.45 (0.05)d
Forest Fallows 380 (15.2)c 84 (3.2)a 3.93 (0.05)ab
Residential Tourism 349 (16.1)bc 64 (2.6)bc 3.70 (0.04)c
Forest Edges 296 (17.1)c 83 (3.2)ab 3.96 (0.04)a
Wildlife Refuges 269 (14.2)cd 71 (2.6)b 3.83 (0.02)b
Biological Reserves 250 (12.8)cd 69 (2.4)b 3.84 (0.03)ab
Tree Plantations 231 (15.5)d 62 (2.6)bc 3.77 (0.05)bc
Data represents means per census (N = 12) and standard errors. Different letters within same column
indicate differences across land use types
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Table 3.2. Sorensen similarity index (top values) and shared species (bottom values)
of ten land-use types in the Path of the Tapir Biological Corridor
Sorensen Index
BR FE FF HG OP ASP WR RT TP VI
BR 0.756 0.628 0.524 0.373 0.444 0.825 0.505 0.619 0.385
FE 135 0.753 0.663 0.486 0.593 0.754 0.62 0.729 0.489
FF 109 143 0.762 0.58 0.704 0.609 0.715 0.775 0.583
HG 91 126 141 0.671 0.792 0.523 0.772 0.757 0.699
OP 53 77 89 103 0.711 0.358 0.735 0.652 0.74
ASP 75 110 127 143 106 0.46 0.736 0.691 0.7
WR 134 135 106 91 51 78 0.476 0.624 0.362
RT 74 101 113 122 93 113 70 0.76 0.729
TP 96 125 129 126 88 112 97 106 0.647
VI 55 78 90 108 91 105 52 93 88
Number of Shared Species
BR = Biological Reserves, FE = Forest Edges, FF = Forest Fallows, HG = Homegardens, OP = Oil palm,
ASP = Agrosilvopastoral systems, WR = Wildlife Refuges, RT = Residential Tourism Projects, TP = Tree
Plantations, VI = Villages.
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Figure 3.1. Location of the Path of the Tapir Biological Corridor, Costa Rica.
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Figure 3.2. Accumulated number of bird species found in ten different habitat types
in the Path of the Tapir Biological Corridor, Costa Rica.
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Figure 3.3. Relative abundance of the five more abundant bird families per land-use
type in the Path of the Tapir Biological Corridor, Costa Rica
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Figure 3.4. Accumulated expected number of bird species (2-year data) in ten
habitat types in the Path of the Tapir Biological Corridor, Costa Rica.
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R P P G T
VI W BR FE TP O AS H FF R
-400
Similarity
-800
1200
0 1 2 3 4 5 6 7 8 9 10
Land-use Type
Figure 3.5. Similarity of bird assemblages across ten habitat types in the Path of the
FE = Forest Edges, FF = Forest Fallows, BR = Biological Reserves, WR = Wildlife Refuges, HG =

Homegardens, TP = Tree Plantations, ASP = Agrosilvopastoral systems, RT = Residential Tourism
Projects, OP = Oil palm, VI = Villages.
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Chapter 4
Response of Bird Guilds to Ten Different Land-use Types in a Human-Modified

Biological Corridor, Costa Rica: Conservation Implications
Abstract
Many bird species are important because they provide important ecosystem
services such as pest control, pollination, and seed dispersal. This chapter aims to
describe the main bird guilds found in the Path of the Tapir Biological Corridor (PTBC)
and how they contribute to the corridor’s conservation goals. A total of 200 points were
monitored for a period of 400 hours along a two-year period, including ten land-use
types: wildlife refuges, biological reserves, forest edges, forest fallows, residential
tourism projects, villages, tree plantations, homegardens, oil palm plantations, and
agrosilvopastoral systems.
At each point, aside from identifying each bird seen, the microhabitat where they
were seen was recorded (e.g. tree, shrub/bush, palm, human infrastructure, etc) as well as
bird activity (e.g. foraging, perching, just moving). Relative to feeding guilds, of the 301
species and 43,911 birds included in this assessment, 53.8% of species were classified as
insectivores, 21.2% frugivores, 9% nectarivores, and carnivores and granivores 8% each
one. On the other hand, classification of birds by habitat preferences depicted that 44.5%
of total species were forest specialist, 38.9% forest generalist, and 16.6% open area
species. Relative abundance of species did not present statistical differences across the
ten land-use types for feeding guilds (G = 5.14, df = 5, p = 0.398), but it did for habitat
guilds (G = 8.372, df = 3, p = 0.039). However, statistical differences were found for
average number of species and abundance for both guild assemblages across the ten land-
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use types (p < 0.05). As anticipated, for forest-dependent species, wildlife refuges,
biological reserves, and forest edges were the best habitat. However, tree plantations,
homegardens, and forest fallows were also important habitats for this group. The lowest
number of forest specialist species was found in oil palm plantations. Open-area species
were more commonly found in agrosilvopastoral systems, villages, homegardens, and oil
palm plantations. For the bird assemblages that provide specific environmental services,
the understory insectivores and large-seed disperers are the groups that may be of further
conservation priority because they are more sensitive to disturbances. Moreover, aside
from protection of forests, management of more complex homegardens and silvopastoral
systems, forest fallows, and native tree plantations can contribute to the protection of bird
species of concern and therefore the environmental services they provide to the
environment.
Introduction
Because of the concerns about the effects of habitat loss and fragmentation on
wildlife conservation, there have been an increasing number of studies on single species
or assemblages in fragmented landscapes throughout the world (Laurance and
Bierregaard 1997; Bennett 2003). However, most of these studies have focused on the
fragments themselves, ignoring species' distribution in surrounding sites such as
agricultural or other human-dominated areas (Ricketts et al. 2000, Chazdon et al. 2009).
Overall, it has been stated that bird species respond differently to disturbances and that
species’ ecological traits (body size, local population size, and geographic range size) and
evolutionary relationships may influence responses in some guilds (Gray et al. 2007).
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For instance, Gray et al. (2007), based on 25 studies and 1178 bird species in Asia and
the Neotropics, found that after disturbances granivores population increased and
frugivores and insectivores declined in both Asia and Neotropical regions.
Aside from the impact of disturbances on wildlife species, there is a general
concern regarding the effect that disturbances can have on functional diversity, food web
interactions, and ecosystems functioning, especially in human-modified landscapes in the
tropics where there is little known (Petit and Petit 2003; Farwig et al. 2008; Tscharntke et
al. 2008). Therefore, for conservation planning of buffer zones, biological corridors, and
other human-modified landscapes, studies of habitat utilization by wildlife species
assemblages are needed to identify key habitats, foraging and nesting sites, and dispersal
routes of animal guilds in planning conservation units (Chazdon et al. 2009). At the
landscape level, it is also important to determine the presence of species in these habitats
that provide ecological services (Tscharntke et al. 2008; Chazdon et al. 2009). For
instance, many bird species are important because they often provide important
ecosystem services such as pest control, pollination, and seed dispersal (Sekercioglu
2006; Tscharntke et al. 2008).
This chapter aims to describe the main bird assemblages found in the human-
modified Path of the Tapir Biological Corridor and how these assemblages affect
conservation goals. These bird assemblages are based on bird diet as well as preferred
habitat. First, there will be a description of bird species distributions based on their
different feeding and habitat guilds in the PTBC. Second, how bird guilds utilized the ten
land-use types is explored in detail, and finally how presence or absent of specific guilds
may affect ecosystem functions and vegetation composition is explained.
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Methods
Study site description, land-use types assessed, and data collection
For this chapter, the same ten habitat types and their corresponding 20 point
counts per land-use type described in Chapter 3 were used: forest edges, forest fallows,
wildlife refuges, biological refuges, tree plantations, villages, oil palm plantations,
residential tourism projects, agrosilvopastoral systems, and homegardens.
When conducting bird surveys, the bird activity at the time of observation was
recorded (e.g. foraging, nesting, perching, or just moving), as well as the microhabitat
where they were found. Microhabitat categories varied from trees, shrub, bush, palm,
grass/ground, to human infrastructure (e.g building, house, fence, and utility wire). The
microhabitat’s height where birds were observed was also estimated.
Bird guilds
For the present analysis, birds were classified by habitat and feeding preferences.
Bird species were classified in three habitat guilds: forest specialist species, forest
generalist species, and open area species (Stiles and Skutch 1989; Garrigues and Dean
2007). Forest specialists are the species that mostly utilize closed-canopy forests. Forest
species were also classified as canopy and understory forest species. Canopy birds were
defined as the species that frequent middle and upper levels of close-canopy forests
whereas understory birds were defined as the species that spend most of their time in
understory vegetation or on the ground. Both understory and canopy birds can be found
in closed-canopy, primary and old-secondary forests. Forest generalist species can
occupy closed-canopy forests as well as forest edges and clearings. Open-area species are
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not found in forests; they usually frequent areas where trees may be present, but not
necessarily.
Bird species were also classified based on their primary diet preference as
frugivores (e.g. fleshy fruits), nectarivores (nectar), carnivores (large arthropods and
vertebrate prey), granivores (seed and nuts), and insectivores (small arthropods),
following Stiles and Skutch (1989) and Garrigues and Dean (2007).
Data analysis
This chapter’s analysis did not include nocturnal species, or species commonly
found in wetland ecosystems, or swifts and swallows because I did not sample wetlands
and or conduct nocturnal surveys. Moreover, most literature excludes these species when
guild analysis is included (e.g. Hughes et al. 2002)
A one-way ANOVA was used to compare mean number of species and mean
abundance across land-use types. The means were estimated based on values for each of
the 12 censuses conducted. For habitat and feeding guilds, a log-likelihood G statistic
was computed to compare the overall guild structure of bird communities among all
habitats, represented as the number of species in different guilds (Petit and Petit 2003).
The nonparametric test Kruskal-Wallis was used to compare the incidence (percentage of
species) and relative abundance (percentage of birds) for each guild among habitat types.
Krustal-Wallis was also performed to compare bird activity when recorded (the
percentage of birds per activity) as well as to compare the structure where birds were
observed (the percentage of birds per tree/shrub/bush) among the ten different land-use
types. Non-metric multidimensional scaling (NMDS) were performed to see the
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relationship between canopy and understory forest species and land-use types. All
statistical analyses were conducted in PAST v 1.95 (Hammer et al. 2001) and R statistical
software.
Results
Habitat guilds
For the current analysis, 301 species and 43,911 individuals were taken into
account. Of these, 44.5 % were classified as forest specialists, 38.9% as forest generalists,
and 16.6 % as open area specialists. Comparing the total accumulated number of bird
species, the distribution of habitat guild members differed significantly across the ten
land-use types (G = 8.372, df = 3, p = 0.039; Figure 4.1) consistent with the predicted
habitat requirements of guild members. Guild assemblages shifted gradually from a
predominance of forest specialists and just a few open area specialists in forested habitats
to a higher proportion of open area species and few forests specialist in open area land-
use types such as villages and oil palm plantations (Figure 4.1). Analysis of variance also
depicted statistical differences for species richness and abundance across the ten habitat
types (p < 0.05; Table 4.1). Wildlife refuges, biological reserves, and forest edges had the
highest abundance and species richness values for forest specialist species and also the
lowest for open area specialists, as anticipated. Concentration of species and abundance
of understory and canopy forest specialist were also different across the ten habitat types
(Figure 4.2; 4.3). The differences among forested habitats and open area habitats were
better identified by understory specialist than by canopy specialist. Many canopy species
can move more easily to land-use types other than forests in search of food because of
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dispersal abilities (e.g. parrots & parakeets). Forest generalist species were more
commonly found in homegardens, followed by forest fallows and agrosilvopastoral
systems. Open area species were more commonly found in agrosilvopastoral systems,
villages, homegardens, and oil palm plantations.
Feeding Guilds
In general, of the 301 bird species included in this analysis, 53.8 % corresponded
to insectivorous species, 21.2 % to frugivores, 9% nectarivores, and 8% each to
carnivores and granivores. Contrary to the findings of habitat guilds, when comparing the
total accumulated number of bird species, distribution of feeding guild members did not
differ significantly across the ten land-use types (G = 5.14, df = 5, p = 0.398; Figure 4.4).
However, there were statistical differences across the ten habitat types for species
richness and abundance when compared at the census level (p < 0.05; Table 4.4).
Insectivorous species not only had the most species, but also had the most
individuals registered (40.4%). The majority of insectivorous species were recorded at
forest edges and forest fallows, followed by wildlife refuges, biological reserves, and
agrosilvopastoral systems. The most important species in terms of abundance were house
wren (Troglodytes aedon) great kiskadee (Pitangus sulphuratus), tropical kingbird
(Tyrannus melancholicus), red-crowned woodpecker (Melanerpes rubricapillus), and
chestnut-backed antbird (Myrmeciza exsul), all together representing 32.5 % of the total
birds for this guild. The former four species were observed more in open habitats while
the antbird was recorded mostly in forested habitats.
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Although frugivores had only 21.2% of the total number of species, they
encompassed 36.9 % of the total number of registered birds. They were more commonly
found in homegardens and forest edges and the most common species were cherries
tanager (Ramphocelus costaricensis), blue-gray tanager (Thraupis episcopus), orange-
chinned parakeet (Brotegeris jugularis), palm tanager (Thraupis palmarum), and
chestnut-mandibled toucan (Ramphastos swainsonii). These five species represent almost
36% of total frugivore sightings. Aside from the toucan, all species did not visit closed-
canopy forests. Oil palm plantations had a statistically lower species richness and
abundance values across the 10 land-use types (p < 0.05).
Granivores represented 17% of total records for this study. As expected, average
abundance and species richness were highest for open habitats such as agrosilvopastoral
systems, oil palm plantations (e.g. young plantations), homegardens, and villages. On the
contrary, the lowest numbers were found in the forested land-use types. The most
commonly found species were great-tailed grackle (Quiscalus mexicanus), variable
seedeater (Sporophila americana), blue-black grassquit (Volantinia jacarina), ruddy-
ground dove (Columbina talpacoti), and white-tipped dove (Leptotila verreauxi),
representing 81.7% of the registered granivores. Quiscalus mexicanus is considered an
invasive species of open areas.
Nectarivores and carnivores comprised only 3.8% and 1.9% of the birds,
respectively. While nectarivores tended to be more commonly found in homegardens and
forest edges, carnivores were more commonly encountered in agrosilvopastoral systems,
villages, and oil palm plantations. Blue-throated goldentail (Hylocharis eliciae), rufous-
tailed hummingbird (Amazilia tzacatl), bananaquit (Coereba flaveola), long-billed hermit
106
(Phaethornis logirostris), and purple-crowned fairy (Heliothryx barroti) were the most
abundant nectarivorous species (64.0% of all nectarivores); and roadside hawk (Buteo
magnirostris), black vulture (Coragyps atratus), turkey vulture (Cathartes aura), yellow-
headed caracara (Milvago chimachima), and laughing falcon (Herpetotheres cachinnans)
were the most commonly recorded carnivores, representing 83.9% of all birds in this
guild.
Resource use
Overall, it was estimated that the activity of a particular bird was not independent
of the land-use type where found (X2 = 1594.1; df = 27; p < 0.001). Moreover, birds were
recorded foraging more than any of the other activities (perching, moving, or nesting) in
the ten different land-use types (Figure 4.5). Of all species, 263 species (87.4%) were
registered foraging at least once, with forest fallows (154 species), homegardens (145
species), and forest edges (144 species) being the most visited habitats. In contrast, only
51 species were registered nesting, preferring homegardens (21 out of 51 species) and
villages (16 out of 51 species). Great kiskadee was observed nesting 41 times, social
flycatcher (Myiozetetes similis) 21 times, gray-capped flycatcher (Myiozetetes
granadensis) 13 times, and the yellow-throated euphonia (Euphonia hirundinacea),
golden-hooded tanager (Tangara larvata), and streaked flycatcher (Myiodynastes
maculates) 10 times each.
Finally, the bird preference by microhabitat (e.g. trees, shrubs/bush, ground,
human infrastructure) was assessed. It was found that microhabitat used by birds and
land-use are not independent (X2 = 21095; df = 36; p = 0.000; Figure 4.6). In general,
107
with the exception of oil palm plantations, more than 65% of the individuals were
observed in trees, a small percentage were also recorded in shrub/bush combinations and
the least were observed in the ground/grass and human-infrastructure (e.g. utility wires,
fences, buildings). It is important to highlight that more than 43 % of sightings in oil
palm plantations were recorded in plants other than the palms, highlighting the
importance of other vegetation in large-scale monocultures. Remnant trees and living
fences were a part of these monocultures, as well as grassy/bushy areas in younger stages
of the palm plantations.
Discussion
Habitat guilds
Conservation of bird guilds in human-dominated landscapes depends on many
factors, including the configuration of the matrix around forested areas as well as the
adaptability of many of these species assemblages to changes at the micro and macro
level (Sodhi et al. 2004; 2008; Brockerhoff et al. 2008). The general concept that the
human-modified landscapes are very important for the conservation of open-area and
forest-generalist bird species was confirmed in this study. However, it was found that 70
out of 134 forest-dependent species were also utilizing these habitats. Understanding the
persistence mechanism of tropical forest species in human-modified landscapes is
therefore a fundamental challenge of tropical ecology and conservation, especially
because of the low adaptability of many of these species to disturbances (Sodhi et al.
2004; Sekercioglu et al. 2006).
108
Several studies have shown that some of these human-modified land-use types
can contribute to the conservation of a portion of forest specialists (Petit and Petit 2003;
Farwing et al. 2008). For instance, in western Kenya, Farwing et al. (2008) did not find
statistical differences in the number of forest specialist species between natural forests
and native mixed and pure plantations. All of these land-use types had a higher number of
forest specialists than exotic monocultures in the same region (Farwing et al. 2008). In
the current study, for forest bird species, forested habitats were statistically more diverse
and more abundant than any other human-modified land-use type. However, 67, 55, and
51 forest specialist species utilized forest fallows, tree plantations, and homegardens as
alternative habitat, respectively. From the conservation point of view, this is positive
because it indicates that not all forest species are totally restricted to forested habitats,
increasing availability of suitable habitat, especially for connectivity purposes.
Forest fallows or secondary forests have been mentioned as an alternative habitat
for forest species (Dunn 2004; Waltert et al. 2004; 2005; Chazdon et al. 2009). In
Uganda, songbird community composition and richness of forest bird species were
similar between intact, mature forest and 13-year old regeneration forest (Naidoo 2004).
However, some studies have found that secondary forests and exotic tree plantations can
only hold a few forest species compared to primary forests (Zurita et al. 2006; Barlow et
al. 2007), contradicting the conservation values of secondary forests described in other
studies. Regardless of these numbers, both tree plantations and secondary forests may be
more suitable habitat than degraded traditional pasturelands. For instance, cattle pastures
left fallow for less than two years supported more than twice as many total species as
actively grazed pastures in Panama (Petit et al. 1999). In Colombia, tree plantations also
109
had higher bird species richness than other more intense production systems such as
pasture lands (Mendoza et al. 2007).
For the present study, tree plantations also had statistical differences from the
native and the exotic species assessed as a part of this study. For instance, twice as many
forest specialist species utilized T. amazonia plantations (native - 52 species and 684
individuals) than T. grandis plantations (exotic - 26 species and 100 individuals).
According to Brockerhoff et al. (2008), the extent of the difference between natural forest
and tree plantations varies considerably across the range of management intensities and
the degree to which plantations deviate from the tree species composition and structure of
natural forests in the same region. Natural regeneration growing beneath the canopy of
most T. amazonia plantations may provide different micro-habitat conditions than the
absence of understory under T. grandis plantations because of different management
regimes. Only three out of 55 forest specialist species found in the tree plantations were
solely recorded in T. grandis and not in T. amazonia: olivaceous piculet (Picummus
olivaceus -1 bird), yellow-bellied flycatcher (Empidonax flaviventris - 3 birds), and
scarlet macaw (Scarlet macao - 2 birds).
As for native tree plantations, complex agroforestry systems such as homegardens
and some types of agrosilvopastoral systems are more supportive of forest species than
mono-crop systems (McNeeley and Schroth 2006; Scales and Marsden 2008). However,
as for secondary forests and tree plantations, the presence of forest-dependent species in
agroforestry systems depends on several factors; and most of the time agroforestry
systems have been cited as having a low number of species from this guild compared to
primary forests (Scales and Marsden 2008).
110
For the current study, it is interesting to note that a few forest specialists were
found even in highly human-impacted habitats such as villages and residential tourism
projects as well as large-scale monocultures such as the oil palm plantations. However,
because development is an unplanned ongoing process in the region (Redondo-Brenes
and Villalobos 2008), response of bird species composition and abundance may change
in the coming years as urbanization spreads along the corridor. For now, even the
endangered scarlet macaw can be seen in a few of the villages and homegardens,
especially southern ones where macaws feed on several tree species around human
settlements (See Chapter 5). Contrary to these findings in the PTBC in Panama, Petit and
Petit (2003) did not report any forest specialist species utilizing fallow pastures, pine
plantations, sugar cane, grazed pastures, villages, and rice paddies.
Overall, data from the corridor landscape matrix for bird species seems to present
promising numbers in comparison to some other studies in the tropics, especially with
regard to the utilization of these human-modified habitats by forest species. However,
because distance from primary forest was not taken into account and many of the
sampling points were close to them, this factor may favor the presence of more forest-
dependent species in the human-modified matrix (Barlow et al. 2007). Some of the
forest-specialist species recorded in this study frequently utilizing the human-modified
landscape were the chestnut-mandibled toucan, red-lored parrot (Amazona autumnalis),
fiery-billed aracari (Pteroglossus frantzii), short-billed pigeon, and cocoa woodcreeper
(Xiphorhynchus susurrans), great crested flycatcher (Myisrchus crinitus), mealy parrot
(Amazona farinose), and scarlet macaw. Each species was observed more than 100 times
over the 12 censuses in the 2-year period of this study. With the exception of the fiery-
111
billed aracari in oil palm plantations, cocoa woodcreeper in villages, and scarlet macaws
in oil palm, agrosilvopastoral systems, and residential tourism projects, all other highly
abundant species were found in all six human-modified habitats.
Bird feeding guilds and environmental services
In most rain forests, birds are the single most important group of seed dispersal
agents (Primack and Corlett 2005), and the fate of the seed – and thus the success of seed
dispersal - depends critically on how the frugivorous animals select and process the fruits,
and what it does afterwards (Primack and Corlett 2005). For instance, it was estimated
that 90% of seed dispersal agents for 336 tree species in ten tropical rain forests of
northern Costa Rica were animals (Chazdon et al. 2003). Guindon (1997), studying seed
dispersal of cloud forests in Costa Rica, also reported that 80% of 225 tree species in his
study site depended on vertebrates for their dispersal with most (63%) primarily by birds.
Several studies in the tropics have shown that the loss of frugivores in forests can have
significant impacts on forest structure and composition (Guindon 1997; Gross-Camp et
al. 2009; Moran et al. 2009). For instance, change in the composition of the frugivores
assemblage in fragmented subtropical Australian forests has decreased the dispersal of
12% of the native plants species; and populations of these plants have substantially
reduced in the region studied (Moran et al. 2009). In addition, lack of bird dispersers for
the endemic tree Leptonychia usambarensis (Sterculiaceae) was the main factor
associated with seedlings and saplings being more aggregated around mother trees in
fragmented forests than in continuous forests in Tanzania (Cordeiro and Howe 2003),
increasing predation and therefore mortality.
112
In the PTBC, frugivorous species composition changed across the ten different
habitat types, but abundance of frugivores was similar across most land-use types, except
oil palm plantations where it was lowest (Figure 4.4, Table 4.2). For conservation
purposes, this similarity is important because bird abundance rather than species richness
has been mentioned to be the main factor that drives seed dispersal in human-modified
landscapes (Pejchar et al. 2008). Moreover, in fragmented forests small-seeded plants
may be more resilient to forest fragmentation while large-seeded species may be more
susceptible because large-seeded species rely on a few, extinction-prone dispersers
(Guindon 1997; Cramer et al. 2007), and should be a priority for conservation. In the
PTBC, important small-seed dispersal families such as Frigillidae (euphonia), Thraupidae
(tanagers), and Pipridae (manakins) were recorded in large numbers in all habitat types.
The latter family was found mostly in forested areas, but the former two were
encountered in different land-use types.
Some general, large-seed dispersal birds such as chestnut-mandibled toucan were
registered in all ten land-use types. The other member of the Ramphastidae family, the
fiery-billed aracari, was also found in all habitats except oil palm plantations. Other
large-seed dispersers such as great curassow (Crux rubra) and crested guan (Penelope
purpurascens - both Cracidae) were totally restricted to forested habitats and both are
enlisted as endangered species (See Chapter 5). Of greater concern is the crested guan
whose observations were lower than great curassow. The Three-wattled bellbird
(Procnias tricarunculatus), an altitudinal migrant species, is another large-seed disperser
that was also restricted to forested habitats. This species is considered an important
dispersal agent, especially of the Lauraceae family in montane forests of Costa Rica
113
(Guindon 1997). The trogons (Trogodinae), also known as important middle-size seed
dispersal agents in the tropics (Primack and Corlett 2005), were found in most habitat
types, especially forests, homegardens, silvopastoral systems, and tree plantations.
Like frugivores, nectarivores have important consequences for plant population
and community dynamics (Sekercioglu et al. 2004). However, the proportion of bird
species that provide pollination services to plants in the Neotropics is lower than those
that disperse seeds (Primack and Corlett 2005). In fact, Chazdon et al. (2003) found that
only 24 out of 336 tree species in Northern Costa Rica were primarily pollinated by
hummingbirds, the main bird pollinator group in the Neotropics (Primack and Corlett
2005). Hummingbirds usually pollinated herbaceous plant species such as Heliconias and
other forest understory plants (Primack and Corlett 2005). For the current study, 27
hummingbird species were identified, favoring homegardens, forest edges and forest
fallows, where abundant numbers of flowering herbaceous plants were available.
However, it was also interesting than 47% of the time, hummingbirds were registered
foraging on trees. Flowers of tree species such as Erythrina poepiggiana, Cedrela
odorata, Bombacopsis quinatum, several species of the genus Inga sp, Cecropia sp,
several fruit trees, among others, were some of the trees hummingbirds commonly
visited.
In the tropics, the most common guild of bird species is the insectivores (Primack
and Corlett 2005). However, because of their high ecological specialization, they are
highly sensitive to fragmentation and therefore are extinction-prone (Sekercioglu et al.
2004), especially forest understory species (Rendijo 2001; Laurance et al. 2002; Lens et
al. 2002; Siegel et al. 2006; Stouffer et al. 2006; Peters et al. 2008). For the present study,
114
insectivores accounted for 53.8 % of the species registered. They were found in the ten
land-use types, with forest edges, forest fallows, biological reserves, wildlife refuges, and
homegardens being the most important land-use types for this guild. The lowest numbers
for both frugivores and nectarivores were in oil palm plantations. As mentioned above,
understory insectivores have been cited as the guild that is most affected by
fragmentation and land-use change. For the ten land-use types in this study, numbers
confirm that pattern, as more understory insectivores were restricted to forested areas (13
out of 33 - 40% of total members for this guild) than canopy insectivores (22 out of 129 -
only 17% of all members for this guild).
Insectivores feeding not only benefits natural forests (Van Bael et al. 2003), but
also important crops in tropical countries such as coffee (Kellernmann et al., 2008;
Johnson et al. 2009), cocoa (Van Bael et al. 2007), and oil palm plantations (Koh 2008)
by decreasing foliage damage by arthropods. Furthermore, insectivorous birds can benefit
human welfare by increasing total revenues from healthier crops. For instance, in
Jamaica, Kellermann et al. (2008) estimated a benefit of US $4018 (118% of the per
capita GNI for the 2004-2005 season) to four farmers from bird insectivores, which led to
healthier coffee fruits , and consequently increased value on international markets.
Another bird group that also provides benefits to the environment and humanity
are the carnivores. They are important in maintaining low rodent populations and also
have important decomposition functions (e.g. scavengers) (Primack and Corlett 2005). In
the PTBC they seem to be equally distributed, but are not abundant in species and
individuals compared to frugivores or insectivores. The largest raptors that depend on
forested areas are the ones that seemed to be under more serious threats as a result of
115
forest disappearance. However, whereas at present they can be seen in various locations
throughout the corridor, according to local residents most of them were gone because of
losing of forest cover in the 1960s and 1970s. White hawks (Leucopternis albicollis) and
King Vultures (Sarcoramphus papa) exemplify two of the species that are more
commonly found in the corridor now than 10-20 years ago.
Conclusions
In the Path of Tapir Biological Corridor, land use has changed from dense
primary forests in the 1900s to extended and degraded pasture lands in the 1960s and
1970s (Redondo-Brenes and Montagnini 2010). This was followed by the recovery of at
least 50-60% of the forest cover throughout 1990s and 2000s through natural
regeneration and establishment of tree plantations (Redondo-Brenes 2007; Redondo-
Brenes and Montagnini 2010). However, as land value has increased, a new and
uncontrolled tourism and residential development boom is jeopardizing the connectivity
of large forest fragments in most of the Pacific coast of Costa Rica (Redondo-Brenes and
Villalobos 2008). The disappearance of natural forests and consequent avian composition
shift may also affect several of the environmental services that birds provide to the
system, including pest control, seed dispersal, and pollination.
Conservation planning should focus on protecting main forest fragments within
biological corridors, limiting residential development in these areas, especially to protect
some of the most sensitive guilds such as understory insectivores and large-seed
dispersals that are only found in closed-canopy primary and old-secondary forests.
116
In addition, land-use practices that include management of more diverse
agroforestry systems, native tree plantations, and secondary forests as buffer zones or as
connectors of the already existing forest fragments should be promoted in order to
provide more suitable habitat for the forest-dependent species and open-area species.
Moreover, inclusion of more fruiting trees as part of the landscape may attract more birds
that may facilitate the regeneration of bird-dispersed rain forest plants (Luck and Daily
2003).
Connectivity of forest fragments through suitable habitat (e.g secondary forests)
may aid several sensitive species to survive in the short to middle term, allowing them to
move in between fragments (Stouffer et al. 2006), but there is no guarantee that all
species will survive in the long term (Stouffer et al. 2006; Sigel et al. 2006). For instance,
even a large reserve (La Selva in Costa Rica – 1611 ha), despite a forested connection to
Braulio Carrillo National Park (45,000 ha), is apparently too small to even maintain all
species of one major guild (understory insectivores) (Sigel et al. 2006). Therefore, since
the PTBC (80,000 ha) has less than 50,000 ha under forest cover and struggles with large
residential projects, large efforts are required to maintain and enhance connectivity in
order to achieve effective diversity conservation, especially for the species that are more
susceptible to fragmentation or those that need large habitat areas to survive in the long
term.
117
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Table 4.1 Bird abundance and species richness by habitat guilds in 10 land-use types in the Path of the Tapir Biological
Corridor, Costa Rica. Data are averages per census (N=12).
Forest Specialist Forest Generalist Open Area Species
Species Species Species

Land Use Richness Abundance Richness Abundance Richness Abundance
W Refuge 56.2 (0.8)a 231.3 (8.9)a 13.6 (1.7)e 34.9 (5.6)d 0.7 (0.3)f 2.1 (1.1)e
B Reserve 55.4 (1.4)ab 210.6 (9.4)a 12.6 (1.1)e 36.8 (4.4)d 1.1 (0.4)f 2.3 (0.9)e
F Edge 50.4 (1.4)b 204.5 (7.6)a 27.8 (1.9)cd 79.8 (9.8)c 3.9 (0.5)f 10.9 (1.8)bc
F Fallow 30.8 (1.2)c 112.5 (7.1)b 39.1 (2.3)ab 188.3 (10.3)ab 14.3 (0.7)d 79.4 (5.2)ab
T Plantation 24.3 (1.0)d 54.3 (5.6)c 27.2 (1.6)cd 103.8 (8.5)bc 10.3 (0.6)e 61.8 (6.3)b
Homegarden 21.3 (1.6)d 70.8 (4.7)bc 43.3 (1.8)a 234.4 (9.1)a 22.0 (0.7)b 264.3 (8.6)a
R Tourism 15.8 (1.2)e 52.7 (3.9)c 29.5 (1.4)bcd 139.1 (8.93)b 17.8 (0.6)c 153.7 (7.8)a
Silvopastoral 12.1 (1.2)e 32.9 (4.1)d 35.9 (2.2)b 188.9 (10.8)ab 26.7 (0.6)a 268.4 (6.4)a
Oil Palm 5.2 (0.7)f 9.8 (1.3)f 24.7 (1.1)d 111.6 (5.3)bc 21.3 (0.6)b 221.3 (7.9)a
Village 5.1 (0.8)f 17.5 (1.8)e 26.9 (1.1)cd 146.3 (10.9)b 22.1 (0.6)b 324.9 (9.1)a
Values represent accumulated average values for each of the 12 censuses conducted and they are followed by standard
errors in parentheses. Different letters mean statistical differences with an α < 0.05.
123
Table 4.2 Bird abundance and species richness by feeding guild in 10 land-use types in the Path of the Tapir Biological
Corridor, Costa Rica. Data are averages per census (N=12).
Frugivores Insectivores Nectarivores Granivores Carnivores

Land
Use Species Abundance Species Abundance Species Abundance Species Abundance Species Abundance
ASP 20.7 (1.0)cd 152.3 (10.3)bc 34.3 (2.0)abc 182.0 (9.1)ab 4.8 (0.4)bc 9.8 (0.7)de 9.3 (0.4)a 116.9 (9.2)a 4.6 (0.3)a 13.9 (1.9)a
BR 22.5 (0.9)bc 95.5 (6.0)d 36.8 (1.4)ab 134.3 (7.5)bc 6.1 (.3)b 14.0 (1.0)cd 2.3 (0.3)c 3.7 (0.5)d 1.5 (0.4)c 1.8 (0.5)e
FE 26.1 (1.1)ab 120.8 (9.0)cd 40.5 (2.3)a 142.3 (8.2)bc 9.4 (0.7)a 20.5 (1.7)ab 3.2 (0.2)c 7.3 (0.9)c 2.7 (0.3)bc 3.8 (0.5)de
FF 25.8 (0.8)abc 157.9 (7.9)b 40.3 (2.4)a 144.1 (9.0)bc 8.8 (0.5)a 19.3 (1.6)bc 7.3 (0.4)b 53.4 (4.8)b 1.8 (0.2)c 5.2 (0.7)cd
HG 28.2 (0.9)ab 247.7 (6.2)a 35.9 (2.5)ab 191.7 (9.7)a 10.1 (0.6)a 25.5 (1.6)a 8.3 (0.4)ab 68.9 (6.0)ab 3.0 (0.2)bc 6.1 (0.7)bcd
OP 11.2 (0.4)e 63.3 (5.4)e 23.6 (1.2)cd 148.3 (6.9)ab 2.0 (0.4)c 3.2 (0.7)e 8.8 (0.4)ab 104.4 (7.2)a 3.8 (0.3)ab 10.2 (1.0)ab
RT 20.3 (0.9)c 146.8 (5.8)bc 26.6 (1.5)cd 123.6 (7.9)bc 5.8 (0.5)b 13.6 (2.1)cd 7.3 (0.3)b 51.5 (4.6)b 2.4 (0.3)bc 5.6 (0.8)bcd
TP 17.5 (0.9)d 84.2 (6.2)d 31.8 (1.9)bc 114.7 (10.3)c 5.8 (0.5)b 11.8 (1.0)d 3.6 (0.4)c 10.9 (1.2)c 2.7 (0.4)bc 8.3 (0.9)abc
VI 17.1 (0.6)d 184.8 (8.5)b 19.8 (1.1)d 147.1 (7.5)ab 3.4 (0.3)c 7.2 (0.7)e 8.1 (0.5)ab 55.8 (5.3)b 3.8 (0.4)ab 11.8 (2.0)ab
WR 20.8 (1.0)cd 98.5 (4.9)d 38.3 (1.7)ab 149.8 (9.2)ab 6.3 (0.4)b 12.4 (0.8)d 2.1 (0.1)c 4.0 (0.5)d 3.0 (0.5)bc 3.6 (0.6)de
Values represent accumulated average values for each of the 12 censuses conducted and they are followed by standard errors in parentheses. Different
letters mean statistical differences with an α < 0.05.
ASP = Agrosilvopastoral systems; BR = Biological Reserves; FE = Forest Edges; FF = Forest Fallows; HG = Homegardens; OP = Oil Palm; RT =
Residential Tourism Projects; TP = Tree Plantations; VI = Village; WR = Wildlife Refuges
124
Figure 4.1. Relative abundance of species by habitat guild (percentage of species)
within 10 land-use types in the Path of the Tapir Biological Corridor, Costa Rica
125
sp29
1.0
sp19
0.5
TP OP
sp12
NMDS2
sp23
sp21
sp52 sp45
sp38
sp5 sp49 sp17
sp1
sp55
sp6 sp24
sp11
sp39
0.0
sp36 sp41
sp28 sp48
BR sp31
WR sp8 RT
FE HG sp27
sp2
sp40
sp20
sp22
sp14
sp30
sp53
sp25
sp35
sp13 FF ASP
sp34
sp15
sp26
sp46
sp54 sp9 sp16
sp50
sp4 sp51 sp44
sp10
sp42 sp47 sp33
sp43 sp3
sp32 sp56
sp7
sp18 VI
-0.5
sp37
-1.5 -1.0 -0.5 0.0 0.5 1.0 1.5
NMDS1
Figure 4.2 Non-metric multidimensional scaling (NMDS) ordination of the
distribution of canopy forest specialist species in ten land-use types in the Path of
the Tapir Biological, Corridor.
BR = Biological Reserve; WR = Wildlife Refuge; FE = Forest Edge; FF = Forest Fallow; TP = Tree

Plantation; HG = Homegarden; ASP = Agrosilvopastoral System; RT = Residential Tourism; OP =
Oil Palm Plantation; VI = Villages. Numbers represent species (See Appendix 1)
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1.0
sp34
sp78
0.5
OP
sp38
sp26
sp72
sp59 sp66
NMDS2
FF
sp57
sp16 sp49sp22 sp17

sp31
sp32
sp5
sp53
sp45 sp40 sp28
sp55
sp70
sp20 sp18 WR
sp60 sp64 sp9 sp69
sp56 sp25
sp21
sp63sp50 sp14
0.0
sp8
sp65sp3 sp7 sp15
sp76 sp43 sp46 VI
BR sp19
sp13sp36 sp29 HG sp12
sp71
sp10
sp67
sp54 sp30
sp73
sp77
sp74
sp75
sp6
sp61
sp62
sp44 sp35 sp33 sp42
FE sp2 RT sp4
sp51
sp27
sp37
sp23 TP
sp58
sp52
sp48
sp39 sp68
sp11
sp47 sp24 sp41
ASP
-0.5
sp1
-1.5 -1.0 -0.5 0.0 0.5 1.0
NMDS1
Figure 4.3 Non-metric multidimensional scaling (NMDS) odination of the
distribution of understory forest specialist species in ten land-use types in the Path
of the Tapir Biological, Corridor.
BR = Biological Reserve; WR = Wildlife Refuge; FE = Forest Edge; FF = Forest Fallow; TP = Tree

Plantation; HG = Homegarden; ASP = Agrosilvopastoral System; RT = Residential Tourism; OP =
Oil Palm Plantation; VI = Villages. Numbers represent species (See Appendix 1)
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Figure 4.4 Relative abundance of bird species by feeding guild (percentage of
species) within 10 land-use types in the Path of the Tapir Biological Corridor, Costa
Rica
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Figure 4.5 Relative abundance of birds (percentage of birds) for activity within 10
land-use types in the Path of the Tapir Biological Corridor, Costa Rica
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Figure 4.6 Relative abundance of birds by microhabitat (percentage of birds) within
10 land-use types in the Path of the Tapir Biological Corridor, Costa Rica
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Chapter 5
Conservation of Threatened and Endemic Bird Species in a Human-Modified

Biological Corridor: Sustainable Forestry Criteria Recommendations
Abstract
Conservation of threatened and endemic species is a priority at the local, national,
and global levels, especially in the current human-modified landscapes. These groups of
species are relevant because they are intolerant of forest fragmentation and land-use
change and identification of their habitat preferences is important for their conservation.
This chapter will describe the status of endangered, threatened, and endemic bird species
found in the Path of the Tapir Biological Corridor, Costa Rica. In addition, based on
species habitat preferences and following sustainable forestry criteria, a series of
recommendations are described for conservation of bird species, and also for other
species of conservation concern. Using 200 sampling counts at ten different land-use
types (wildlife refuges, biological reserves, forest edges, forest fallows, tree plantations,
residential tourism projects, villages, oil palm plantations, homegardens, and
agrosilvopastoral systems), 32 threatened species and 22 endemic species were identified.
Eight of the 22 endemic species were also under threat, and three endangered species
were registered: Ara macao, Amazilia boucardi, and Icterus mesomelas. Overall, 64%
and 75% of endemic and threatened species, respectively, were forest-dependent species.
Moreover, frugivores and insectivores were the most abundant guilds across endemic and
threatened species. When the habitat dependency of bird species and the socio-political
and economic situation in the PTBC were considered, based on concepts of sustainable
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forestry, it was assessed that the corridor could be sustainable if the natural resources are
managed in three units that includes protected areas, integrated management areas, and
tree plantations. Using payment for environmental services, ecotourism, reforestation,
environmental education programs, and promoting national and foreign investment in the
region, among other strategies, could provide the rural infrastructure that local people
need to practice integrated management effectively.
Introduction
Fragmentation and isolation of Costa Rican protected areas are jeopardizing
biological conservation (Powell et al. 2000; Sanchez-Azofeifa et al. 2003). Therefore,
given these limitations of the protected area system, there is a pressing need to study
biodiversity and improve management not only within protected areas’ boundaries, but
also in the fragmented landscape matrices that surround them. However, because of the
complexity of human-modified landscapes, especially in the tropics, there are no generic
recipes that can be applied appropriately everywhere for conservation planning
(Lindenmayer and Franklin 2002; Lindenmayer 2009).
Threatened species (Gaston and Blackburn 1996), endemic species (Myers et al.
2000), and small geographic range species (Rendijo 2001) should be considered as
priorities for conservation planning. These groups of species are relevant because they
are intolerant of forest fragmentation and land-use change (Rendijo 2001) and
identification of their habitat preferences is important for their conservation (Gaston and
Blackburn 1996). Moreover, although it may take a century for all sensitive species to be
extirpated from a site following habitat loss, species with larger or heavier bodies and
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those foraging on insects, fruits, or both are particularly extinction prone, and therefore
should also be considered within sustainable forest management practices (Sodhi et al.
2004).
This chapter will describe the status of endangered, threatened, and endemic bird
species found in the PTBC. In addition to assessing the status of species of concern, some
alternatives for their conservation following sustainable forestry criteria developed by
Oliver (2003) and Oliver and Deal (2007) will be described. The goals of sustainable
forest management are to provide a steady flow of resources and income, while
preserving forest cover, biodiversity and ecosystem integrity (Oliver 2003). According to
Oliver and Deal (2007) two ways of discussing sustainable forestry have been proposed.
The first approach (Venn diagram approach) considers sustainable forestry as
overlapping acceptable ecological, economic, and social activities. The second approach,
known as the matrix approach, ensures that one ecosystem is neither hoarding nor overly
exploiting forest values to the detriment of itself or other ecosystems (Oliver and Deal
2007).
Based on the PTBC’s conservation objectives, recommendations are described for
conservation of endangered and endemic species while managing existing resources. The
recommendations may be applicable to other human-modified landscapes with similar
ecological, political, and socio-economic features. The criteria include biodiversity
analysis and the status of the threatened and endemic species found in the PTBC,
socioeconomic benefits, productive capacity, forest health, soil and water quality, and
carbon sequestration in the region (Oliver 2003).
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Methods
Study area: Path of the Tapir Biological Corridor
Conservation objectives
The corridor’s main objective is to create a network of sites favorable to fauna
and flora between the forests of the Osa Peninsula and Golfo Dulce, including Corcovado
National Park. These forests connect with those located in the Los Santos Forest Reserve
in the Talamanca Mountain Range. In addition, the PTBC was created as a means to
achieve four major goals (TNC-ASANA 2000; Redondo-Brenes 2007a): (1) Restore
natural habitats to enable the return of terrestrial endangered species and marine species
to the forests and beaches they inhabited half a century ago, and protect existing species
by allowing local populations to become more robust; (2) Halt deforestation within the
corridor and restore forests in order to complete the corridor and protect water sources for
use by communities, particularly in watersheds and groundwater recharge zones; (3)
Encourage local communities to value resources and use them sustainably; and (4)
Reduce and prevent soil erosion to benefit local farmers and for biodiversity conservation
in rivers, mangroves, and coral reefs within the region, especially in Marino Ballena
National Park.
Corridor justification
According to TNC-ASANA (2000), there are four main elements that justify the
creation of the PTBC: (a) providing connectivity between protected areas, (b) protecting
endemic species, (c) protecting endangered species, and (d) promoting sustainable
management of natural resources.
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Providing connectivity between protected areas
One of the main problems with the protected areas system in Costa Rica is the
isolation of most of these areas (Sanchez-Azofeifa et al. 2002a, 2003). The PTBC, which
is a part of the multinational Mesoamerican Biological Corridor, was proposed with the
intent of diminishing the negative effects of fragmentation and habitat loss in the Central
and South Pacific regions of the country by providing connectivity through several forest
remnants (e.g., privately-owned protected areas as stepping-stone corridors) to protect
areas such as Corcovado National Park, Piedras Blancas National Park, the Terraba-
Sierpe Wetlands, and Golfo Dulce and the Los Santos Forest Reserves.
Protecting endemic species
According to a rapid ecological assessment (TNC-ASANA 2000), the PTBC area
has a moderate level of endemism in comparison with other areas of the country such as
Corcovado National Park. For example, the PTBC provides habitat to 6 plant species
endemic to Costa Rica. From the 26 amphibian species found in the PTBC, four species
are endemic to the country and another four are found only in the area between Nicaragua
and Panama. In addition, one mammal, eight reptiles, and 11 bird species endemic either
to Costa Rica alone or jointly to Costa Rica and Panama are also found in the corridor
area. Finally, special attention needs to be placed on fish species since the corridor has 13
endemic species, representing 68.4% of the freshwater endemic species of the country.
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Protecting threatened and endangered species
As a result of habitat loss, fragmentation, and illegal and legal hunting and
poaching, a total of 38 species of flora and fauna are considered either threatened or
endangered species in the region (IUCN 2009). Within the 38 species, there are six trees,
three amphibians, seven reptiles, 10 birds, and 12 mammals. According to the 2009
IUCN Red List of Threatened Species, the study area provides habitat to several
endangered bird and mammal species such as the squirrel monkey (Saimiri oerstedii),
yellow-billed cotinga (Carpodectes antoniae), and mangrove hummingbird (Amazilia
boucardi). In addition, the turquoise cotinga (Cotinga ridwayi) and the three-wattled
bellbird (Procnias tricarunculata) are considered vulnerable species, and the black guan
(Chamaepetes unicolor), the great curassow (Crax rubra), baird’s trogon (Trogon
bairdii), the harpy eagle (Harpia harpyja), and the jaguar (Panthera onca) are listed as
species near threatened (IUCN 2009). Other species of concern in the IUCN list found in
the region and considered priority species for conservation in Costa Rica are margay
(Leopardus wiedii), ocelot (Leopardus pardalis), scarlet macaw (Ara macao), and spider
monkey (Ateles geoffroyi).
Promoting sustainable management of natural resources
Aside from the ecological justifications such as biodiversity protection, there were
at least two socio-economic aspects relevant for the creation of PTBC: water-soil
protection and ecotourism. This area has more than 30 rivers, all of which provide
potable water for domestic consumption and water used in agriculture practices,
especially in the corridor lowlands. Thus, it is justifiable to protect the corridor highlands
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in order to secure water quality for its inhabitants, especially because to date local people
see water scarcity and quality as an issue in the region (Welsh 2006). Secondly,
ecotourism is growing in the region and this is very important because it provides new
job opportunities for local people. Even when not considering rivers, this area still has
more than 50% of the landscape under forest cover. It also provides the visitors with
waterfalls and more than 70 km of coastline.
Land-use types part of this study
For this chapter, I used the same ten habitat types and their corresponding 20
point counts per land-use type described in Chapter (3): forest edges, forest fallows,
wildlife refuges, biological refuges, tree plantations, villages, oil palm plantations,
residential tourism projects, agrosilvopastoral systems, and homegardens.
Identification of bird species of concern
Species of conservation concern
The list of species of concern described above is based on the IUCN Red List.
However, at the regional level, this list gets larger when the list of species that are
considered under any threat at the country level is included, especially for bird species.
For instance, Obando (2008) recently cited that 9.6% and 1.8% of total bird species in
Costa Rica are threatened and endangered species in the country, respectively. The
Ministry of Environment 2005-Decree lists 81 bird species under the category of
vulnerable or threatened species, and 17 bird species under endangered status. From this
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list of species under category of conservation concern, a total of 32 were identified in the
Path of Tapir Biological Corridor (Table 5.1).
Endemic species
In terms of endemism, Costa Rica is considered a restrained country (Obando
2008). Only 1.3% of all species are endemic to the country, birds being the group that
encompasses the lowest proportion (0.7%). However, about 10% of the bird species that
inhabit Costa Rica have very limited ranges and are considered regionally endemic –
most are only found in the southern part of Central America and nowhere else in the
world (Garrigues and Dean 2007). In the PTBC only one species endemic to the country
was identified (mangrove hummingbird – Amazilia boucardi), but 21 regionally endemic
species were registered for this study (Table 5.2).
Data analysis
When analyzing bird relative abundance, the one way ANOVA test was used to
determine if there were statistical differences across the 32 species of conservation
concern and across the 21 endemic species. Abundance was estimated at the census level,
which means the cumulative number of birds per species per the 20 points within each
land-use type was used. Non-metric multidimensional scaling (NMDS) analyses were
conducted to assess the bird species preferences within the ten land-use types.
In addition, a series of graphs were created in order to compare the relative
abundance of total threatened and endemic species by feeding and habitat guilds. Bird
species were classified in three habitat guilds: forest specialist species, forest generalist
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species, and open area species. Forest specialist species mostly utilize closed-canopy
forests. Forest generalist species can occupy closed-canopy forests as well as forest edges
and clearings. Open-area species are not found in forests, but frequent areas where trees
are present. Classification of bird habitat preference was based on Stiles and Skutch
(1989) and Garrigues and Dean (2007). On the other hand, bird species were classified
based on their primary diet preference as frugivores (e.g. fleshy fruits), nectarivores
(nectar), carnivores (large arthropods and vertebrate prey), granivores (seeds and nuts),
and insectivores (small arthropods), following Stiles and Skutch (1989) and Garrigues
and Dean (2007).
All statistical analyses were conducted in PAST v 1.95 (Hammer et al. 2001) and
the NMDS were performed in R statistical software.
Results
Species of conservation concern
Overall, of the 334 bird species found in the PTBC (Chapter 3), 32 species (9.6%)
were classified in at least two categories of conservation concern (threatened or
endangered species, Table 5.1), and 22 species (6.6%) were classified as regionally
endemic (Table 5.2). Only eight endemic species were also considered under threat:
baird’s trogon (Trogon bairdii), crimson-fronted parakeet (Aratinga finshi), golden-naped
woodpecker (Melanerpes chrysauchen), mangrove hummingbird (Amazilia boucardi),
three-wattled bellbird (Procnias trinarunculatus), white-throated shrike-tanager (Lanio
leucothorax), yellow-billed cotinga (Carpodectes antoniae), and turquoise cotinga
(Cotinga ridwayi).
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Of the 32 species of conservation concern, only three were considered endangered
species at the national level: mangrove hummingbird, scarlet macaw (Ara macao), and
yellow-tailed oriole (Icterus mesomelas). Bird abundance across species was statistically
different (p < 0.05; Figure 5.1). For the species of conservation concern, orange-chinned
parakeet (Brotegeris jugularis) was the species with the highest abundance, followed by
white-crowned parrot (Pionus senilis), red-lored parrot (Amazona autumnalis), crimson-
fronted parakeet (Aratinga finschi), mealy parrot (Amazona farinosa) (all Psittacidae),
and golden-naped woodpecker (Melanerpes chrysauchen). The other 26 species did not
have statistically significant abundance differences and 16 of these species had less than
1 bird registered per census. This means less than 1 individual was observed in 33.3
hours (200 points).
As for endangered species, one endemic species had a statistically large difference
in abundance with respect to the other species in this category (Figure 5.2). Cherries
tanager (Ramphocelus costarricensis) was not only the most abundant endemic species,
but also the most common species for the region (Chapter 3). Moreover, there was a
group of 16 species with no statistical differences in their abundance, and nine of these
had less than 1 individual observed per census. Other endemic species with intermediate
population sizes were yellow-crowned euphonia (Euphonia luteicapilla), black-hooded
antshrike (Thamnophilus bridgesi), and riverside wren (Thryothotus semibadius).
When species compositions were compared by guilds, relative species abundance
in different guilds was more consistent by habitat guilds than by feeding guilds (Figure
5.3; 5.4). For instance, the majority of endemic (64%) and threatened species (75%) were
forest specialists, and consequently, the minority was classified as forest generalist or
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open area species. In contrast, when feeding guilds were compared, endemic species had
similar relative species abundances across categories, but the threatened species were
primarily frugivores (53%), followed by insectivores (25%), and then carnivores (15.6%).
However, as anticipated, frugivores and insectivores together made up the majority for
both endemic and threatened species. Furthermore, no carnivore or granivore species was
endemic to this region; and although there were six endemic nectarivorous species, only
the mangrove hummingbird was also considered an endangered species. Of the other
seven endemic species classified as threatened species, only the crimson-fronted parakeet
(frugivore) was an open area specialist, while the other six species were forest-dependent
species. Four of them were also frugivores (yellow-billed cotinga, turquoise cotinga,
bellbird, and baird’s trogon) and two (golden-naped woodpecker and white-throated
shrike-tanager) were insectivores.
Despite the theoretical habitat classification for endemic species and species of
conservation concern described above, several species utilized more than one habitat
(beyond expected) across the ten habitat types assessed. While in forested habitats, 24
species of conservation and 19 endemic species were observed, in human-modified
landscapes 20 species of conservation concern and 18 endemic species were registered.
Similarly, utilization of the different land-use types varied across bird species. For
instance, even though there were 28 forest species (specialist and generalist together) of
conservation concern, three of these species (turquoise cotinga, yellow-billed cotinga,
and scarlet macaw) were not recorded in forested habitats, but were found in human-
modified landscapes. In fact, 20 forest species from this group utilized at least one
human-modified land-use type, and the multivariate ordination depicted that at least 10
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out of 32 threatened species were associated with six of the human-modified land-use
types (Figure 5.5).
Land utilization of endemic species was similar to the pattern of threatened
species. Several forest-dependent and generalist species were using human-modified
habitats. At least nine out of 22 endemic species could be associated more with human-
modified habitats than forested habitats (Figure 5.6).
Discussion
Use of Sustainable Forestry Concepts in the PTBC
Many frameworks and approaches are found in the literature concerning the best
landscape management practices (i.e. sustainable forestry) to simultaneously maintain
forest biodiversity, ecological functions, and human livelihoods (Hunter and Calhoun
1996; Campos et al. 2001; Noss 2001; Putz et al. 2001; Lindenmayer and Franklin 2002;
Oliver 2001, 2003; Dudley et al. 2005; Higman et al. 2005; Hagan and Whitman 2006).
Regardless of the authors’ background, affiliation, standpoint, or perspective, a common
goal within sustainable forest management is to protect biodiversity. According to
Higman et al. (2005), sustainable forestry can be defined as the forest management
practices that provide goods and services from a forest ecosystem without degradation of
site quality, and without a decline in the yield of goods and services over time. Similarly,
Oliver (2003) mentioned that the definition of sustainable forestry has its roots in the
concept of environmental justice. In other words, people living in one place and time
should provide their “fair share” of values –neither unfairly exploiting nor depriving
themselves of certain values to the detriment or benefit of people in another place or time.
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Achieving a sustainable forestry goal will probably take time as people agree on what
sustainability means at the global, sub continental, national, and regional scales (Oliver
2001).
Aside from timber and biodiversity, forested lands provide other environmental
goods and services such as carbon sequestration, soil and water protection, scenic beauty,
food provision (Groot et al. 2002), and jobs for rural people (Oliver 2003), among others.
However, most of these goods and services as well as biodiversity are being threatened
by habitat loss and the fragmentation of natural habitat by humans (Groom et al. 2005),
along with climate change, nitrogen deposition, and invasive species (Sala et al. 2000).
And as a result of these threats, the conservation community is working with other
stakeholders including government agencies, industry groups, forestry schools, and
consumers of forest products to develop conservation-planning strategies for biodiversity
conservation (Gordon et al. 2005). The main goal is to protect habitats, to maintain
biological diversity, to preserve ecological functions, and to ensure sustainable forest
management in managed areas (Oliver 2003; Gordon et al. 2005). Unfortunately, even
though many of these concepts have been implemented in temperate regions, little is
known about how tropical forests can be best managed to enhance biodiversity (Putz et
al. 2001). But, some recent forest sustainable management projects implemented in the
tropics have sent a positive message about its feasibility (Frost et al. 2006, Dennis et al.
2006, Peña-Claros et al. 2008; Fonseca et al. 2009).
This discussion focuses on the status and habitat requirements of the threatened
and endemic species in the PTBC, by providing an analysis of socio-political and
economic factors that may affect the implementation of sustainable forestry following
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Oliver (2003). The six criteria for achieving sustainable forestry described by Oliver
(2003) are evaluated: biodiversity (protecting habitats for endangered/endemic species),
socioeconomic benefits, productive capacity (wood and non-wood), forest health, soil
and water quality, and carbon sequestration.
Biodiversity conservation of endangered and endemic species
Biodiversity conservation has become the stated objective of national
governments, state agencies, local communities, and scientific organizations (Redford
and Richter 1999). According to Dirzo and Raven (2003), the global state of biodiversity
is important because biodiversity is a central component of Earth’s life support systems
and directly relevant to human societies. Any attempt to defend a social cause, such as
biodiversity, should rest on the best facts available; and the loss of biodiversity is the only
truly irreversible global environmental change the Earth faces today. Moreover, Powell et
al. (2002) mention that biodiversity has intrinsic values and provides practical benefits
such as (1) medicines, plant derivatives and other timber and non-timber products, (2) a
basis for medicinal bioprospecting, (3) pollination and resilience to agricultural systems,
and (4) soil building and nutrient cycling processes. Moreover, since biodiversity has
social value, from recreation to spiritual and cultural benefits, it is increasingly
recognized as fundamental to human health (Powell et al. 2002), and this is a reason to
protect forested lands.
In order to evaluate the role of human-modified land-use types for biodiversity
conservation, it is important to know not only how many species are present within these
systems, but also which species are present and whether any of these species are of
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conservation concern (Petit et al. 1999; Harvey and Gonzalez Villalobos 2008). For the
PTBC, a total of 46 out of 334 species were classified as either threatened/endangered or
regionally endemic species, including eight species that are endemic and under threat.
Among the 46 species, they differed in diet and habitat preference as well as their
conservation priority within the corridor’s conservation goals. For instance, conservation
efforts may not focus on species like the cherries tanager, which despite being an
endemic, forest generalist, and frugivorous species, was the most abundant species in this
study (Chapter 3), and was found in eight land-use types, excluding the interior of
wildlife refuges and biological reserves. However, their role as seed dispersers of small
seeds may be relevant to catalyze secondary succession in the region.
In addition, the eleven species of the Psittacidae family registered in this study are
categorized as being of conservation concern or endangered (e.g. Ara macao) as a result
of habitat loss, direct hunting, and poaching for the pet trade market (Stiles and Skutch
1989; Garrigues and Dean 2007). However, most of these species were observed in more
than one habitat type and the majority of them had high populations when compared not
only to endemic and endangered species, but also across all species in the corridor
(Chapter 3). For most of these species, decreased hunting pressure was mentioned by
local people as one of the main reasons for such high populations compared to the last
10-20 years. Within this family, of greater concern are Pionopsitta haematotis, mostly
found in private reserves, and Ara macao, which once resided on both the Pacific and
Caribbean slopes, and now is only located in a few spots along the Pacific Coast (Stiles
and Skutch 1989; Vaughan et al. 2006; Garrigues and Dean 2007). In addition, there have
been a few reports of this species in the Caribbean lowlands recently (Henderson 2010;
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Guindon, personal communication, April 2010). In the corridor, Ara macao populations
were located foraging and perching mostly in homegardens, villages, and in forest edges
in the southern part of the region. A few individuals were also registered in homegardens
of the PTBC’s northern limit; however, because the forests around these villages were not
surveyed, this species was not included as resident of forested habitats. Nevertheless, as
part of this study, macaws were observed in a few forest patches north of where they
were recorded.
Reintroduction of this species is being assessed in the Hacienda Baru Wildlife
Refuge as part of a national reintroduction program run by Amigos de las Aves.
Hacienda Baru, along with forested areas around this refuge, provide the required food
sources as well as large trees (e.g. Ceiba pentandra) for nesting by scarlet macaws
(Castles et al. 2008). This program will increase the likelihood of connecting macaw
populations along the whole Central and South Pacific Coast of the country, thereby
increasing the probability of gene exchanges, and therefore contributing to the
survivorship of this species in the long term.
The other two endangered species Icterus mesomelas and Amazilia boucardi were
registered only once and twice, respectively, over the duration of this project. Icterus
mesomelas is a forest-generalist, insectivore species that forages in thickets and second
growth near water, but is considered a rare resident of the Caribbean lowlands of Costa
Rica (Garrigues and Dean 2007). However, this is not the only record of this species in
the PTBC, it has been reported as part of the Annual Christmas Count, suggesting that
this species as well as several others (e.g. hoffmann’s woodpecker (Melanerpes
hoffmanii), rusty-margined flycatcher (Myiozetetes cayanensis), blue-headed parrot
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(Pionus menstruus), white-throated magpie-jay (Calocitta formosa)) may be spreading in
the Central and South Pacific lowlands from the Caribbean and other regions of the
country (Ureña, 2005; Garrigues and Dean 2007).
On the other hand, A. boucardi is an endemic species of the Pacific coast of Costa
Rica, specifically mangroves and adjacent forests (Garrigues and Dean 2007).
Unfortunately for this species and many others that are mangrove-dependent
(Nagelkerken et al. 2008), these habitats are illegally disappearing for agricultural
expansion and residential development, despite mangrove forests being considered
protected areas in Costa Rica (Redondo-Brenes and Villalobos 2008). Throughout the
two years I spent in the PTBC, I participated in at least four inspections where damages
to mangroves were found, including drainage, canalization, land-use change, etc. Where
mangroves are native, similar patterns are happening everywhere in the tropics,
jeopardizing the ecological functions these ecosystems provide to people and the
environment (Nagelkerken et al. 2008).
Other bird species whose conservation is also at risk include those that occur only
in large patches of forests and that are absent from small patches with little or no true
interior habitat, as well as non-flying species with poor dispersal abilities (Laurance
2004). Some of these species found in the PTBC were Procnias tricarunculatus,
Spizaetus ornatus, Buteogallus urubitinga, Sarcoramphus papa, Cotinga ridgwayi,
Carpodectes antoniae, Penelope purpurascens, Crax rubra, Micastur semitorquatus,
Deconychura longicauda, Laniocera rufescens, Odontophorus gujanensis, Pionopsitta
haematotis, Lophostrix cristata, Tinamus major, and Trogon bairdii. From this group of
species, more attention should be placed on P. purpurascens, C. rubra, Procnias
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tricarunculatus, and T. bairdii because of their role as large-seed dispersers (Chapter 4;
Guindon 1997; Jimenez et al. 2003; Primack and Corlett 2005). Furthermore, forested
habitats are not only relevant because they provide habitat for these endangered/endemic
species listed above, but also for several understory species that do not cross the matrix,
including army ant followers and members of mixed-species flocks that strongly avoid
forest edges (Rendijo 2001; Laurance 2004). Some of these species, with small
populations in this study, were bare-crowned antbird (Gymnocichla nudiceps), bicolored
antbird (Gymnopithys leucaspis), black-faced antthrush (Formicarius analis), buff-
throated foliage-gleaner (Automolus ochrolaemus), russet antshrike (Thamnistes
anabatinus), streak-chested antpitta (Hylopezus perspicillatus), among many other
species.
Unfortunately, urban and tourism development are jeopardizing the conservation
of many of these species in the region, especially because forest cover is either
disappearing or being fragmented around most private reserves (Redondo-Brenes 2007a;
Redondo-Brenes and Villalobos 2008). Therefore, implementation of sustainable forest
management may provide not only a larger chance for these species to survive, but also
more incentives to local stakeholders to protect natural habitats.
Socioeconomic Benefits
According to Oliver (2003), socioeconomic benefits and biodiversity are two
main criteria used in sustainable forestry approaches such as the Montreal Process
Criteria. The main goal is to integrate socioeconomic benefits and biodiversity from a
local and a global perspective, especially because many scales need to be considered to
148
ensure that actions at one place or scale create positive consequences elsewhere (Oliver
2003). In other words, the goal here is to protect forested reserves for biodiversity while
obtaining economic benefits from these reserves as well as other managed landscapes
such as agricultural fields and tree plantations. Accomplishing these sustainable forestry
practices in a tropical region will need to consider several points within the agenda.
First, the potential conflicts between conservation and human welfare that have
alienated potential allies in the past have to be avoided, especially in human-dominated
landscapes in tropical regions (Brandon and O’Herron 2004; Anderson and Jenkins 2006;
Hilty et al. 2006; Chang et al. 2007). Second, it has to be stated that management for
some goods and services necessarily involves management against some others (Putz et
al. 2001). Therefore, several scenarios can be endeavored in the PTBC to achieve the
biodiversity and socioeconomic benefits criteria.
In a human-dominated landscape like PTBC, protected areas are interspersed
amongst other land-use scenarios such as residential development, pasture lands, tree
plantations, annual and perennial crops, and tourism projects, among others. One of the
first steps is to analyze how to create or to ensure a network connects the more than 30
privately-owned protected areas and Marino Ballena National Park within the PTBC
(Reserves concept in Oliver 2003), and also to provide buffer zones for adjustment of
reserves boundaries (Noss 2001). The reserves can be connected using either natural
regeneration of abandoned agricultural lands or reforestation of those degraded lands
with native species through payments of environmental services (PES). At present,
FONAFIFO, the Costa Rican government institution in charge of the PES program, is
paying a total of US$64/ha/year towards natural forest protection, US$41/ha/year towards
149
natural regeneration of abandoned lands, US$816/ha/5-year period towards the
establishment of tree plantations, and US$1.30/tree towards agroforestry systems
(FONAFIFO 2009). In the PTBC, the PES implementation has secured protection of
more than 5,000 ha from 1997 to 2008. From 1997 to 2002, payments to participants in
the region totaled nearly US$175,000 per year (Newcomer 2007). Although PES
programs are not designated for poverty reduction (Pagiola et al. 2005), for small local
farmers in the corridor, the earnings from the PES represent an important extra income,
especially when they have had a poor agricultural season because of the market prices or
weather conditions.
Despite the fact that PES has been a good source of income for several program
participants, and has helped to protect forests in the corridor, the PES scheme currently
used in Costa Rica needs some re-structuring. Poor funding (e.g., the demand for PES is
much higher than the funding currently available), delays in payments, medium-term
time frames (e.g., PES contracts range from 5 to 10 years), disproportionate
representation of program participants by large farmers and forest owners, and some
mismatch between conservation areas defined by FONAFIFO and conservation priorities
within each region, are some of the flaws of the program (Zbinden and Lee 2005;
Newcomer 2007; Redondo-Brenes 2007a). Thus, either the government should create a
better framework for the PES, or the PTBC organizations should look for alternative
funding sources to provide economic incentives to owners willing to preserve their land
(Anderson and Jenkins 2006; Redondo-Brenes 2007a). When a PES program design is
well thought out and local conditions are favorable, there can be important synergies
between economic, social, and ecological benefits (Pagiola et al. 2006).
150
One of the main approaches to promote sustainable forestry in the region through
economic development (i.e. PES) could be to reduce the need for subsistence farming and
grazing (Oliver 2001) that ultimately would promote better land-use practices. Aside
from PES, alternatives seen as important to promote both human welfare and
conservation in the region include ecotourism, establishment of tree plantations and
agroforestry systems, and low impact logging.
Because of all the natural resources found in the PTBC, ecotourism has grown
very quickly in the region. Most of the privately-owned protected areas integrate
protection and ecotourism, generating an extra income for the protected areas owners and
the people depending on tourism. Many local people who previously depended on
agriculture now are working directly or indirectly in ecotourism. Overall, it is suggested
that ecotourism may potentially make a positive contribution to resource conservation at
the local, national (Weinberg 2002; Rivera 2004; Kruger 2005), and global scales (Hunter
and Shaw 2007). However, unsustainable urbanization can negatively affect the region if
it surpasses the carrying capacity of the land (TNC-ASANA 2000; Newcomer 2007;
Basso and Newcomer 2009). Therefore, proper land-use planning is necessary for the
region.
Reforestation programs can also provide economic benefits and contribute to
biodiversity conservation (Tiarks et al. 1998; Hartley 2002; Lamb et al. 2005; Carnus et
al. 2006), and in fact, within the Costa Rican Plan of Sustainable Forestry, reforestation
and agroforestry systems are a priority (Calvo 2009). Within the PTBC there are more
than 20,000 ha under pastures that could be used for reforestation programs. Although
most of the reforestation programs in Costa Rica have used exotic species, there are
151
several good examples of successful reforestation trials (Redondo-Brenes and
Montagnini 2006) and reforestation programs in small- and medium-sized farms with
native tree species (Redondo-Brenes 2007b; Streed et al. 2006). For example, Streed et
al. (2006), using native species within the PTBC, show that small-scale reforestation
(18.6 ha) with mixtures of native species can be financially profitable for both an investor
and a farmer/landowner, and increases biodiversity in the region through reduced-impact
management (Leopold and Salazar 2008).
Niewenhuyse et al. (2000) demonstrated the profitability of tree plantations in the
Caribbean lowlands of Costa Rica, as they found that melina (Gmelina arborea) and teak
(Tectona grandis) plantations were more profitable than pasture for beef production.
Aside from the economic benefits tree plantations can provide to landowners and the
increase in timber supply in Costa Rica, we also have to consider that tree plantations, if
not intensively managed, can provide habitat for wildlife (Carnus et al. 2006).
Furthermore, all tree plantations, including those of exotic species and those intensively
managed, are better for biodiversity conservation than degraded pasture lands (Gascon et
al. 1999; Ricketts et al. 2001; Lindenmayer and Franklin 2002; Lamb et al. 2005; Carnus
et al. 2006).
The last alternative for integrating biodiversity conservation and economic
benefits in the corridor is reduced-impact management of primary and old secondary
forests. At the landscape level, disturbance in some areas (i.e., forest logging) is
important for biodiversity management because it can provide habitat for some species,
especially wildlife species, that only inhabit open areas (Oliver et al. 1998). In addition,
Dunn (2004) proved empirically that reduced-impact logging appears to have much less
152
impact on faunal biodiversity (i.e., ants, birds, and lepidoptera) than conventional logging
or forest conversion to agriculture does. In Costa Rica, Campos-Arce et al. (2001) and
Campos-Arce (2006) mention that certified reduced-impact managed stands have
minimal effect on the habitat function that natural forests provide. This is based on
harvesting 4-6 trees/ha in a 15-year cycle, according to a 10-year study from researchers
of the Tropical Agricultural Research and Higher Education Center (CATIE) (Campos-
Arce et al. 2001; Campos-Arce 2006). Therefore, it would be possible to identify some
areas of primary forest and old-secondary forests in which local people can practice
reduced-impact logging, following the recommendations by Campos-Arce et al. (2001) to
mitigate the direct and indirect effects of harvesting on biodiversity. However, these areas
have to be outside protected areas and those connecting protected areas to ameliorate
negative effects (Noss 2001; Sekercioglu 2002).
Productive capacity
Even though Oliver (2003) states, (1) the world is consuming much less wood
than it is growing, (2) there seems to be a surplus of low quality temperate and tropical
hardwoods, (3) wood consumption is helpful by substituting for more fossil fuels-
consuming construction products made of steel, concrete, aluminum, and brick, (4) and
sustainable forestry can provide environmental friendly products and employment in rural
areas, the situation in Costa Rica is totally different as much more wood is consumed
than is growing in productive forests and tree plantations (Arce-Benavides and Barrantes-
Rodriguez 2004; Calvo 2009). In fact, the most recent State of the Nation report 2009
mentioned that after 2010 the timber deficit in the country will reach 850,000 m3/year as
153
a result of shortages of timber coming from tree plantations (main source of timber in the
country), agroforestry systems, and sustainable forest management (Calvo 2009). For this
reason, the forestry sector proposal includes a reforestation program of 7,500 ha/year and
the management of 150,000 ha of forests, using PES and also private investments (Calvo
2009). The current market requires both softwood and hardwood (Arce-Benavides and
Barrantes-Rodriguez 2004; Calvo 2009).
In the PTBC, low quality wood can be obtained from tree plantations established
with fast-growing timber species such as melina, or the native species Vochysia
guatemalensis and Vochysia ferruginea (Leopold et al. 2001; Redondo-Brenes and
Montagnini 2006; Redondo-Brenes 2007b). In addition, secondary forests can also be a
good source of this type of timber (Redondo-Brenes et al. 2001; Valenciano and
Redondo-Brenes 2006). On the other hand, high-quality wood can be obtained from
primary forests and tree plantations of species such as teak and Terminalia amazonia
(Healey and Gara 2003; Kanninen et al. 2004; Carpenter et al. 2004; Streed et al. 2006;
Redondo-Brenes 2007b). Unemployment in the region was roughly 6% in 2004
(Newcomer 2007), so management of both tree plantations and natural forests would
provide not only timber to the forestry industry, but also jobs for local people.
Forest Health
Sustainable forestry can be promoted by keeping fires, insect and disease
outbreaks, and exotic weed invasions to non-catastrophic levels by appropriate
management (Oliver 2003). In Costa Rica, even though there have been more than 1,500
plant species introduced in the country (Obando 2002), there have been no reports of
154
serious insect and disease outbreaks or exotic weed invasions in the PTBC. However,
human-caused forest fires are becoming very common in the dry months, affecting more
than 200 ha in 2009.
Soil and Water quality
Sustainable forestry can increase economic development by alleviating soil
erosion, compaction, and nutrient depletion associated with subsistence farming and
grazing (Oliver 2003). Erosion problems are easily detected in the PTBC landscape. The
main causes of erosion are overgrazing and erosion from clear-cut areas to build new
residential areas. Sediments from both sources are polluting water sources (e.g., rivers
and springs), jeopardizing not only water quality for people and cattle consumption, but
also killing fish and other aquatic organisms in rivers. Without appropriate plans for its
sustainable management and usage, water is at risk of increased scarcity and decreased
quality (Welsh 2006). Costa Rica has recently been facing many watershed issues
because of a rising population and increased demand for clean drinking water, combined
with unsustainable land-use practices (Sanchez-Azofeifa et al., 2002b). In the PTBC,
Welsh (2006), studying water sustainability in two watersheds, discovered that the
decrease in water quality in two major rivers was related to a lack of forest cover in the
region, and that soil sediments coming from degraded pasture lands were the main source
of pollution.
The introduction of trees either in tree plantation or agroforestry systems has been
a common practice in Costa Rica to improve soil properties (Redondo-Brenes 2005). In
the country, several studies on the effects of trees on soil properties have been conducted
155
in the last two decades (Stanley and Montagnini, 1999; Montagnini, 2000). Restoration of
the degraded lands in the corridor through natural regeneration or the establishment of
tree plantations or agroforestry systems will not only ameliorate the negative effects of
erosion on water sources, but it would also provide all the economic and ecological
benefits described in the previous sections. To date, restoration of degraded lands around
main drinkable water sources in the PTBC is being achieved with the participation of
local people (Redondo-Brenes 2007a). Soil degradation and water quality are priorities
in the region, and both are also used as a means of preserving and increasing forest cover
because local people have become more conscious about the negative effects of forest
cover loss and degradation (Newcomer 2007; Basso and Newcomer 2009).
Carbon sequestration
Growing trees and storing the timber both within the forest and in forest products
reduce carbon dioxide (Winjum and Schroeder 1997; Oliver 2003). In Costa Rica, studies
of carbon sequestration estimates have been done in secondary forests (Vilchez et al.
2008), pure and mixed native tree plantations (Redondo-Brenes and Montagnini 2006;
Redondo-Brenes 2007b), and agroforestry systems (Montagnini et al. 2005). Overall,
carbon sequestration rates in agroforestry systems, native tree plantation, and secondary
forests range from 1.5 to 7 Mg C/ha/year (Montagnini and Nair 2004; Redondo-Brenes
and Montagnini 2006; Redondo-Brenes 2007b; Vilchez et al. 2007). Redondo-Brenes and
Montagnini (2006) and Redondo-Brenes (2007b) found that because of species growth
patterns in tree plantations, carbon sequestration estimations reflect two scenarios. First,
if the objective is to accumulate carbon in the short term, species such as V.
156
guatemalensis and V. ferruginea are two of the best options because of their fast growth.
On the other hand, if the objective is to create long term carbon sinks, species such as D.
panamensis and C. brasiliense are the best option. Higher density stands are also a good
alternative when carbon sequestration is the main objective (Redondo-Brenes 2007b).
Managing native tree plantations for timber or carbon sequestration may create
conflict among local farmers, government agencies, and potential carbon offsets buyers,
especially under the PES programs (Pfaff et al. 2000; Redondo-Brenes 2007b). Lamb et
al. (2005) mention that planting to generate ecological services (e.g., carbon
sequestration) as well as goods (e.g., timber) is difficult because trade-offs must be made
between the productivity of the desired goods and the provision of ecological services,
and the techniques to achieve these simultaneous goals are still being developed. For
instance, while local farmers are receiving government incentives (e.g., PES) for
establishing tree plantations in Costa Rica, most local farmers perceive plantations as a
timber production tool, and only a few landowners consider their environmental services
to be a unique goal (Delgado 2002; Piotto et al. 2003). Thus, if the government agencies
are evaluating the possibility of selling carbon offsets to a third party, they have to clarify
the objectives of the reforestation programs and the proper manner to manage the
plantations (Redondo-Brenes 2007b). However, even tree plantations designed for timber
production temporarily store carbon, until it is released into the atmosphere after the
burning of the wood (Oliver 2001; Aune et al. 2005).
157
Conclusions: Achieving Sustainability
The corridor could be sustainable if the natural resources are managed in a
gradation including protected areas, integrated management areas, and tree plantations
(Oliver 2003), also known as the Triad Approach (Seymour and Hunter 1999). These
three management units could provide the habitat required by the different wildlife
species in the corridor as well as create economic opportunities or incentives to protect
the PTBC habitats. Protected areas would be the core conservation areas for more
sensitive forest species. Integrated management areas such as agroforestry systems and
other agricultural areas would provide not only income for local people, but also habitat
for open-area species, and connectivity and buffer zones for forest fragments. Finally,
tree plantations would be used to restore degraded landscapes, to provide habitat for
wildlife species, buffer zones and connectivity to forested areas, and to provide goods
and services to local residents.
Using payment for environmental services, ecotourism, reforestation,
environmental education programs, and promoting national and foreign investment,
among others, in the region could provide the rural infrastructure that local people need
to practice integrated management effectively (Oliver 2003). However, it has to be
controlled such that ecotourism projects, agricultural practices, forest management, and
the new residential development projects do not exacerbate the natural resources through
fragmentation and/or degradation/destruction/pollution of forested ecosystems.
It is expected that after the world economic crisis ends forest cover will be
negatively affected in Costa Rica, especially in areas where tourism and agriculture are
the main source of income for local people (Calvo, 2009). In the case of the Path of the
158
Tapir Biological Corridor, this scenario is likely because tourism development has
delayed its expansion due to the crisis. However, because of lack of control from Costa
Rican authorities (Redondo-Brenes 2007a; Redondo-Brenes and Villalobos 2008) as well
as real estate and development sectors being focused on business rather than
sustainability (Basso and Newcomer, 2009), the future of forest cover and therefore
wildlife in the corridor is uncertain. Under the best scenerio, even if forest cover does not
reduce drastically, road and development would fragment the already fragmented
landscape further, affecting the species that require large areas of land connected to
forests to survive. Therefore, there is a need to adopt sustainable forestry practices to
reduce the impacts that development will cause at the landscape level. In addition, there
is a need to provide alternative management opportunities that ensure income for local
people as well as conservation of biodiversity, especially of the more sensitive species.
Conservation of forest cover in reserves and the establishment of tree plantations
and agroforestry systems, especially with native species, will provide many benefits to
society in general. Tropical trees integrating into the landscape as forest reserves, tree
plantations or as part of agroforestry systems (e.g., home gardens, living fences,
agrosilvopastoral systems, improved fallows) can serve diverse economic, social, and
ecological functions that may ultimately help reduce atmospheric CO2 accumulation,
restore degraded soils, protect water sources, protect and enhance biodiversity, contribute
to the wood supply, and improve human welfare (Oliver 2003; Lamb et al. 2005;
Redondo-Brenes and Montagnini 2006; Carnus et al. 2006; Redondo-Brenes 2007b).
159
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Table 5.1 Threatened and Endangered bird species found in the Path of the Tapir Biological Corridor
Common name Scientific name Family Guild Land-use in the PTBC
1 Baird's Trogon Trogon bairdii Trogonidae FR, FS BR, FE, FF, HG, WR, TP
2 Bat Falcon Falco rufigularis Falconidae CA, OA HG, ASP
3 Blue-headed Parrot Pionus menstruus Psittacidae FR, FG HG, VI
4 Brown-hooded Parrot Pionopsitta haematotis Psittacidae FR, FS BR, FE, ASP, WR
5 Collared Forest-Falcon Micrastur semitorquatus Falconidae CA, FS BR, FE, WR
6 Crested Guan Penelope purpurascens Cracidae FR, FS BR, WR
7 Crested Owl Lophostrix cristata Strigidae IS, FS FE
8 Crimson-fronted Parakeet Aratinga finschi Psittacidae FR, OA All
9 Golden-naped Woodpecker Melanerpes chrysauchen Picidae IS, FS All, except VI
10 Great Black-Hawk Buteogallus urubitinga Accipitridae CA, FS BR, WR
11 Great Curassow Crax rubra Cracidae FR, FS BR, FE, WR
12 Great Tinamou Tinamus major Tinamidae FR, FS BR, FE, FF, OP, WR, TP
13 King Vulture Sarcoramphus papa Cathartidae CAM FS FE, WR, TP
14 Long-tailed Woodcreeper Deconychura longicauda Furnariidae IS, FS BR, FE, WR
15 Mangrove Hummingbird Amazilia boucardi Trochilidae NE, FS FF, HG
16 Marbled Wood-Quail Odontophorus gujanensis Odontophoridae IS, FS BR, FE, FF, HG, WR, TP
17 Mealy Parrot Amazona farinosa Psittacidae FR, FS All
18 Orange-chinned Parakeet Brotegeris jugularis Psittacidae FR, OA All, escept WR
19 Orange-fronted Parakeet Aratinga canicularis Psittacidae FR, FG HG
20 Ornate Hawk-Eagle Spizaetus ornatus Accipitridae CA, FS BR
21 Red-crowned Ant-Tanager Habia rubica Thraupidae IS, FS BR
22 Red-fronted Parakeet Touit costaricensis Psittacidae FR, FS FF
23 Red-lored Parrot Amazona autumnalis Psittacidae FR, FS All
24 Scarlet Macaw Ara macao Psittacidae FR, FS HG, TP, VI
25 Speckled Mourner Laniocera rufescens Incertae Sedis IS, FS BR, WR
26 Three-wattled Bellbird Procnias tricarunculatus Cotingidae FR, FS FE, WR
27 Turquoise Cotinga Cotinga ridgwayi Cotingidae FR, FS HG
28 Wedge-tailed Grass-Finch Emberizoides herbicola Emberizidae GR, OA OP
29 White-crowned Parrot Pionus senilis Psittacidae FR, FG All
30 White-throated Shrike-Tanager Lanio leucothorax Thraupidae IS, FS WR, FE, BR
31 Yellow-billed Cotinga Carpodectes antoniae Cotingidae FR, FS ASP
32 Yellow-tailed Oriole Icterus mesomelas Icteridae IS, FG OP
Guilds: FR = forest specialist, FG = generalist, OA = open area species, FR = Frugivorous, IS = Insectivorous, NE = nectarivorous, CA = Carnivore
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Table 5.2 Endemic bird species found in the Path of the Tapir Biological Corridor
Common Name Species Family Guild Land-use Found Geographic Range

1 Baird's Trogon Trogon bairdii Trogonidae FR,FS BR,FE,FF,HG,WR,TP Costa Rica - Western Panama
2 Black-bellied Wren Thryothorus fasciatoventris Troglodytidae IS,FS BR,FE,FF,RT Costa Rica - Western Colombia
3 Black-hooded Antshrike Thamnophilus bridgesi Thamnophilidae IS,FS All, except ASP Costa Rica - Western Panama
4 Charming Hummingbird Amazilia decora Trochilidae NE,FG FE, FF, HG, WR, RT Costa Rica - Western Panama
5 Cherries Tanager Ramphocelus costaricensis Thraupidae FR,FG All, except BR, WR Costa Rica - Western Panama
6 Crimson-fronted Parakeet Aratinga finschi Psittacidae FR,OA All Southern Nicaragua - Western Panama
7 Garden Emerald Chlorostilbon assimilis Trochilidae NE,FG FF,HG,OP,ASP,RT,VI Costa Rica and Panama
8 Golden-naped Woodpecker Melanerpes chrysauchen Picidae IS,FS All, except VI Costa Rica - Western Panama
9 Hoffmann's Woodpecker Melanerpes hoffmannii Picidae IS,FG FF,HG,OP,ASP,RT,TP Southern Honduras - Costa Rica
10 Mangrove Hummingbird Amazilia boucardi Trochilidae NE,FS FF, HG Costa Rica Pacific Mangrove Forests
11 Orange-collared Manakin Manacus aurantiacus Pipridae FR,FS BR,FE,FF,ASP,WR,TP Costa Rica - Western Panama
12 Riverside Wren Thryothorus semibadius Troglodytidae IS,FS All Costa Rica - Western Panama
13 Rufous-winged Woodpecker Piculus simplex Picidae IS,FS BR, WR Eastern Honduras - Western Panama
14 Scintillant Hummingbird Selasphorus scintilla Trochilidae NE,FG RT Costa Rica - Western Panama
15 Snowy-bellied Hummingbird Amazilia edward Trichilidae NE,FS FE Costa Rica and Panama
16 Spot-crowned Euphonia Euphonia imitans Fringillidae FR,FS All, except OP,ASP,VI Costa Rica - Western Panama
17 Three-wattled Bellbird Procnias tricarunculatus Cotingidae FR,FS FE, WR Eastern Honduras - Western Panama
18 Turquoise Cotinga Cotinga ridgwayi Cotingidae FR,FS HG Costa Rica - Western Panama
19 White-crested Coquette Lophornis adorabilis Trochilidae NE,FG Generalist Costa Rica - Western Panama
20 White-throated shrike-tanager Lanio leucothorax Thraupidae IS,FS BR, FE, WR Eastern Honduras - Western Panama
21 Yellow-billed Cotinga Carpodectes antoniae Cotingidae FR,FS ASP Costa Rica - Western Panama
22 Yellow-crowned Euphonia Euphonia luteicapilla Fringillidae IS,OA All, except BR, WR Nicaragua - Panama
Guilds: FR = forest specialist, FG = generalist, OA = open area species, FR = Frugivorous, IS = Insectivorous, NE = nectarivorous
BR = Biological Reserves, FE = Forest Edges, FF = Forest Fallows, HG = Homegardens, OP = Oil palm, ASP = Agrosilvopastoral systems, WR = Wildlife Refuges, RT =
Residential Tourism Projects, TP = Tree Plantations, VI = Villages
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Figure 5.1 Bird abundance by census (N = 12) of endangered and threatened species
found in the Path of the Tapir Biological Corridor. Different letters indicate
statistical differences at α = 0.05. Statistical analysis was performed to log
transformed of means.
171
Figure 5.2 Bird abundance by census (N = 12) of endemic species found in the Path
of the Tapir Biological Corridor. Different letters indicate statistical differences at
α = 0.05. Statistical analysis was performed to log transformed of means.
172
Frugivore Insectivore Nectarivore Carnivore Granivore
100%
80%
Relative abundance of species
60%
40%
20%
0%
Endemic species Threatened species
Birds
Figure 5.3. Relative abundance of endemic and threatened bird species (percentage
of species) by habitat guilds found in the Path of the Tapir Biological Corridor,
Costa Rica
173
Forest Specialist Forest Generalist Open Area
100%
Relative abundance of species
80%
60%
40%
20%
0%
Endemic Species Threatened Species
Birds
Figure 5.4 Relative abundance of endemic and threatened bird species (percentage
of species) by feeding guilds found in the Path of the Tapir Biological Corridor,
Costa Rica
174
sp28
sp32
OP
0.5
sp31
ASP
NMDS2
sp4 sp2
sp6
sp10
sp25 WR
sp20sp30
sp21sp11
sp5
sp14
sp26 BR sp8sp18
0.0
sp12 TP
sp13 sp9 sp29
sp17 RT
FE sp23
sp7 sp1 VI
sp16
sp24 sp3
FF
HG
-0.5
sp22 sp15 sp19

sp27
-1.5 -1.0 -0.5 0.0 0.5 1.0
NMDS1
Figure 5.5. Non-Metric Multidimentional scaling (NMDS) ordination of threatened
and endangered species found in ten land-use types in the Path of the Tapir
Biological Corridor.
BR = Biological Reserves, FE = Forest Edges, FF = Forest Fallows, HG = Homegardens, OP = Oil

palm, ASP = Agrosilvopastoral systems, WR = Wildlife Refuges, RT = Residential Tourism Projects,
TP = Tree Plantations, VI = Villages
175
sp21
0.5
TP
ASP
sp13
BR
sp20 sp1 sp11 sp9
sp8 sp6 sp22

sp3 OP
WR
0.0
sp16 sp12 RT sp14

sp5
VI
NMDS2
sp17
sp7
FE FF
sp2
sp19
sp15 sp4
-0.5
HG
sp10
-1.0
sp18
-1.5 -1.0 -0.5 0.0 0.5 1.0 1.5
NMDS1
Figure 5.6. Non-Metric Multidimentional scaling (NMDS) ordination for endemic
species found in ten land-use types in the Path of the Tapir Biological Corridor.
BR = Biological Reserves, FE = Forest Edges, FF = Forest Fallows, HG = Homegardens, OP = Oil

palm, ASP = Agrosilvopastoral systems, WR = Wildlife Refuges, RT = Residential Tourism Projects,
TP = Tree Plantations, VI = Villages
176
Summary and Conclusions
Bird species diversity in ten different land-use types in the Path of the Tapir
Biological Corridor, Costa Rica was studied. The ten land-use types were wildlife
refuges, biological reserves, tree plantations, forest fallows, forest edges, villages,
residential tourism projects, homegardens, agrosilvopastoral systems, and oil palm
plantations. The interest was determining how important the ten different habitat types
were for maintaining the bird diversity of the corridor and how these land uses can be
managed to enhance their conservation value. To address these questions 20 point counts
for each habitat type were used. Bird surveys were carried out over a two-year period
(2008-2009). Each point was surveyed three times over the dry and three times over the
rainy periods of each year, 12 visits per point. Total observation time was 400 hours.
Aside from bird identification over a period of 10 minutes in 50-meter radius plots, bird
activities (e.g. foraging, nesting, just moving, perching) and microhabitat were registered
(e.g. tree, shrub, bush, ground, utility wire, etc).
Overall, a total of 44,917 birds from 48 families, and 334 species were found.
Eighty one percent of birds were recorded utilizing forested habitats. However, 77% of
registered birds were found in the human-modified land-use types. At the landscape
level, while homegardens, agrosilvopastoral systems, and villages had the highest bird
abundance, tree plantations had the lowest. Forested habitats (i.e. biological reserves,
wildlife refuges, forest edges) also had low bird density values. Moreover, whereas forest
edges, forest fallows, homegardens, biological reserves, and wildlife refuges were the
sites with the highest species richness and diversity of species, tree plantations,
177
residential tourism, oil palm plantations, and villages had the lowest diversity values
within the biological corridor. The latter two were statistically the least diverse sites from
the ten land-use types assessed.
Species composition also varied across the ten habitat types in the corridor.
Biological reserves, wildlife refuges, and forest edges had the most similar assemblage of
species. This may be because these habitats were dominated by primary and old-
secondary forests, having mostly forest-specialist bird species. Also, tree plantations
seemed to share many species with these three habitat types. Agrosilvopastoral systems,
homegardens, residential tourism projects, and oil palm plantations also had similar
assemblage of species because they were dominated by generalist and open area bird
species. In contrast to this high similarity, more open area habitat species (i.e. villages, oil
palm plantations, and residential tourism projects) tended to have the lowest similarity
indexes in relation to forested biological reserves, wildlife refuges, and even forest edges.
When classifying bird species by habitat guild, 44.5% of species were forest
specialists, 38.9% forest generalists, and 16.6% open area specialists. Regarding feeding
guilds, 53.8% of species were classified as insectivores, 21.2% frugivores, 9%
nectarivores, and 8% each for carnivores and granivores. Also, a total of 32 threatened
species and 22 endemic species were identified. Eight of the 22 endemic species were
also under threat, and there were three endangered species registered: Ara macao,
Amazilia boucardi, and Icterus mesomelas. Overall, 64% and 75% of endemic and
threatened species, respectively, were forest-dependent species. Aside from the 46
species that were endangered, threatened, or endemic, there were other species that
should also be considered of conservation concern such as pollinators, seed dispersers,
178
insectivores, scavengers, among others, for the environmental services they provide.
From the insectivorous guild, forest understory dwellers are the most sensitive to land-
use change and were the species that were found the least in the human-modified
landscape. But, overall, all forest-dependent species, including 77 species registered in
these land-use types only, should be considered a priority for conservation planning in the
region.
Based on the results from this study, forested habitats seem to be the most
important land-use type for bird species conservation in the Path of the Tapir Biological
Corridor. No statistical difference was found among the four forested reserves regarding
bird species composition and abundance. Rancho La Merced and Hacienda Baru Wildlife
Refuges as well as Oro Verde Biological Reserve and La Cusinga Eco lodge were
selected because of the commitment of the owners to manage these properties in a
sustainable manner. All four reserves were a mixture of primary and secondary forests
surrounded by other land-use types such as agrosilvopastoral systems, homegardens, tree
plantations, and ecotourism development (e.g., trails and cabins), among others. Except
for Oro Verde, all reserves received tourists regularly, but birdwatching tours are offered
at all locations. Birds were sampled mostly in closed-canopy forests as well as along their
edges. There are around 30 other private initiatives and a national park in the PTBC
similar to these four forested private reserves.
Unfortunately, as mentioned throughout the dissertation, all of these reserves are
interspersed between private properties with multiple landowners. Personal experience in
the area illustrated that there are different interests in and goals for landscape
management within the PTBC. Generally, most local people care about the environment
179
and they are aware of the negative consequences associated with eliminating forest cover.
However, the minority, who do not really care about the environment also own important
forest fragments or have political power, and are promoting unsustainable development.
Furthermore, a lack of economic incentives, and in some cases negligence and lack of
control by the local and national authorities, are promoting illegal development and land-
use change in these forested habitats. At present, even though local groups and NGOs
such as ASANA have been working to mitigate the illegal development expansion in the
PTBC, it is difficult to control because of a lack of commitment by all stakeholders in the
region. Land-use change from forests to any other human activity, including residential
projects, is prohibited by law in Costa Rica, but the lack of law enforcement plus all
factors described above make it possible all over the country.
Tourism has became one of the most important sources of income in Costa Rica
and in many countries in the tropics, however, exponential and unplanned growth of
many of these projects jeopardize the biodiversity protected in forested landscapes. For
instance, in the Osa County, which encompasses around 40% of the PTBC area,
construction concessions grew 202% from January 2007 to September 2009. At the
current trend, and under the best scenerio, even if forest cover does not reduce drastically,
road and development would further fragment the already fragmented landscape,
affecting the species that require large areas of connected forests to survive. Therefore,
there is a need to adopt sustainable forestry practices to reduce the impacts that
development causes at the landscape level. In addition, there is a need to provide
alternative management opportunities to local people that ensure income as well as
biodiversity conservation, especially of the more sensitive species.
180
In Chapter (5), a few alternatives to manage the PTBC as a conservation unit were
proposed based on concepts of sustainable forestry, habitat-dependency of most bird
species, socio-political and economic situation in the corridor, and the existing country
laws. The main goal would be to manage the corridor in a gradation including protected
areas, integrated management areas, and tree plantations. The 30 private reserves are
expected to be the center of conservation in the region, favoring land-use types that
enhance connectivity for wildlife species while providing goods and services to the local
people and the environment, but that also provide habitat to open-area species. Payment
for environmental services, ecotourism, reforestation, environmental education programs,
and private investment, among others, could provide the incentives and infrastructure that
local people need to practice integrated management effectively in the human-dominated
PTBC.
The human-modified land-use types contribute to the high bird diversity of the
PTBC. For instance, more than 77% of bird species in the corridor utilized at least one of
these six land-uses in comparison to 81% found in forested habitats. Forest cover
conservation in reserves and the establishment of tree plantations and agroforestry
systems, especially with native species, could provide many benefits to society in
general. Tropical trees integrating into the landscape as forest reserves, tree plantations or
as part of agroforestry systems (e.g., home gardens, living fences, agrosilvopastoral
systems, improved fallows) can serve diverse economic, social, and ecological functions
that may ultimately help reduce atmospheric CO2 accumulation, restore degraded soils,
protect water sources, protect and enhance biodiversity, contribute to the wood supply,
and improve human welfare.
181
I anticipate that the results from this dissertation will inform local stakeholders
about the importance of the corridor as well as the need to adopt sustainable forestry
practices in the region. However, in order to be effective, the decision making process
should incorporate all participants in the corridor, including government institutions (e.g.
Minister of Environment, Minister of Agriculture and Livestock, Minister of Education,
Water Authorities, the National Institute of Electricity), local people and local groups
(e.g. environmental groups such as ASANA, cooperatives, water management groups
(ASADAS), rural development groups), local universities (e.g University of Costa Rica,
Instituto Tecnológico de Costa Rica, CATIE), international organizations working in the
area (e.g. Pitzer College, TNC), tourism sector (e.g hotel’ owners and managers, travel
agencies), and the real estate and development sectors (e.g. Coldwell Bankers, Punta
Dominical, Costa Rica real state).
Finally, this dissertation is only a two-year bird assessment. To successfully
accomplish and assess the PTBC’s conservation goals or lack thereof, long-term
monitoring programs that include not only birds but also other wildlife taxa and plants is
necessary. Also, more commitment from all stakeholders mentioned above is important
to ensure a truly sustainable corridor in the long term.
182
APPENDIX 1. Abundance data for 334 bird species found at ten land-use types in the Path of the Tapir Biological Corridor,
Costa Rica
Common name Scientific name BR FE FF HG OP ASP WR RT TP VI

Accipitridae
Bicolored Hawk Accipiter bicolor 0 1 0 0 0 0 0 0 0 0
Black-Hawk Eagle Spizaetus tyrannus 0 0 0 0 0 0 1 0 0 0
Broad-winged Hawk Buteo platypterus 3 5 0 0 0 0 5 2 0 0
Double-toothed Kite Harpagus bidentatus 6 6 0 0 0 0 13 0 0 0
Gray Hawk Buteo nitidus 0 2 0 0 0 3 0 1 0 0
Gray-headed Kite Leptodon cayanensis 0 1 0 0 0 0 1 0 1 0
Great Black-Hawk Buteogallus urubitinga 2 0 0 0 0 0 3 0 0 0
Mangrove Hawk Buteogallus subtiles 1 1 0 0 0 0 1 0 0 0
Ornate Hawk-Eagle Spizaetus ornatus 1 0 0 0 0 0 0 0 0 0
Osprey Pandion haliaetus 0 0 0 0 0 0 0 0 0 3
Pearl Kite Gampsonyx swainsonii 0 0 0 0 0 3 0 0 0 0
Plumbeous Kite Ictinia plumbea 0 0 0 0 0 0 0 1 0 0
Roadside Hawk Buteo magnirostris 1 6 49 32 63 73 1 43 73 24
Swallow-tailed Kite Elanoides forficatus 4 2 0 1 0 0 0 0 0 0
White Hawk Leucopternis albicollis 4 2 1 0 0 0 3 0 0 0
Alcedinidea
Amazon Kingfisher Chloroceryle amazona 0 1 0 0 0 0 0 0 0 1
Belted Kingfisher Ceryle alcyon 0 0 1 0 0 0 0 0 0 0
Green Kingfisher Chloroceryle americana 0 4 0 0 1 1 0 0 0 0
Ringed Kingfisher Ceryle torquatus 0 0 1 0 0 0 0 0 0 1
Anatidae
Black-bellied Whistling-Duck Dendrocygna autumnalis 0 0 0 2 61 6 0 0 0 4
Apodidae
Costa Rican Swift Chaetura fumosa 0 0 0 0 0 0 0 0 0 1
Lesser Swallow-tailed Swift Panyptila cayennensis 0 0 0 0 0 0 0 0 0 6
Ardeidae
Bare-throated Tiger-Heron Tigrisoma mexicanum 0 0 0 0 0 4 1 0 0 2
Cattle Egret Bubulcus ibis 0 0 0 13 62 316 0 29 5 12
183
Fasciated Tiger-Heron Trigrisoma fasciatum 0 0 0 0 0 1 1 0 0 0
Great Egret Ardea alba 0 0 0 1 3 20 0 0 0 0
Green Heron Butorides virescens 0 0 0 0 4 10 0 0 0 2
Little Blue Heron Egretta caerulea 0 0 0 0 2 5 0 0 0 0
Snowy Egret Egretta thula 0 0 0 0 3 17 0 0 0 1
Bucconidae
White-necked Puffbird Notharchus macrorhynchos 3 13 3 3 0 3 6 1 2 0
White-whiskered Puffbird Malacoptica panamensis 2 3 0 0 0 0 3 0 0 0
Caprimulgidae
Common Pauraque Nyctidromus albicollis 1 0 3 0 0 1 2 0 4 0
Cardinalidae
Blue-black Grosbeak Cyanocompsa cyanoides 20 36 50 9 4 15 17 18 18 5
Buff-throated Saltator Saltator maximus 10 31 62 46 0 37 1 30 14 7
Dickcissel Spiza americana 0 0 0 0 0 1 0 0 0 0
Indigo Bunting Passerina cyanea 0 0 1 1 0 0 0 0 0 0
Painted Bunting Passerina ciris 0 0 1 1 0 0 0 0 0 0
Rose-breasted Grosbeak Pheucticus ludovicianus 0 1 3 0 0 2 0 0 0 0
Streaked Saltator Saltator striatipectus 0 0 1 0 0 1 0 0 0 0
Cathartidae
Black Vulture Coragyps atratus 0 2 0 15 9 42 2 9 5 54
King Vulture Sarcoramphus papa 0 7 0 0 0 0 1 0 1 0
Turkey Vulture Cathartes aura 2 7 1 1 11 5 6 1 5 40
Charadriidae
Southern Lapwing Vanellus chilensis 0 0 0 1 0 0 0 0 0 0
Columbidae
Band-tailed Pigeon Patagioenas fasciata 0 0 0 0 0 7 20 0 0 0
Blue Ground-Dove Claravis pretiosa 2 3 4 1 0 1 3 2 0 0
Gray-chested Dove Leptotila cassini 21 26 22 17 6 6 10 6 14 18
Gray-headed Dove Leptotila plumbeiceps 4 5 2 0 0 0 5 0 0 0
Inca Dove Columbina inca 0 0 0 19 9 49 0 12 1 41
Pale-vented Pigeon Patagioenas cayennensis 0 4 8 116 12 34 0 29 1 125
Plain-breasted Ground-Dove Columbina minuta 0 0 0 9 0 0 0 0 0 9
184
Red-billed Pigeon Patagioenas flavirostris 0 0 0 3 0 1 1 0 0 0
Rock Pigeon Columba livia 0 0 0 0 0 2 0 0 0 30
Ruddy Quail-Dove Geotrygon montana 9 5 2 0 0 0 13 0 0 0
Ruddy-ground Dove Columbina talpacoti 0 0 35 238 232 185 0 94 2 154
Scaled Pigeon Patagioenas speciosa 1 0 5 12 0 4 0 2 0 0
Short-billed Pigeon Patagioenas nigrirostris 142 125 72 35 21 44 138 58 34 2
White-tipped Dove Leptotila verreauxi 4 35 192 105 83 101 10 111 64 29
Corvidae
Brown Jay Cyanolyca cucullata 2 21 24 26 45 122 2 30 47 12
White-throated Magpie-Jay Calocitta formosa 0 0 0 9 11 0 0 0 6 0
Cotingidae
Three-wattled Bellbird Procnias tricarunculatus 0 2 0 0 0 0 4 0 0 0
Turquoise Cotinga Cotinga ridgwayi 0 0 0 1 0 0 0 0 0 0
Yellow-billed Cotinga Carpodectes antoniae 0 0 0 0 0 1 0 0 0 0
Cracidae
Crested Guan Penelope purpurascens 12 0 0 0 0 0 20 0 0 0
Gray-headed Chachalaca Ortalis cinereiceps 2 5 74 55 0 24 0 50 0 7
Great Curassow Crax rubra 30 5 0 0 0 0 42 0 0 0
Cuculidae
Groove-billed Ani Crotophaga sulcirostris 0 0 0 13 145 20 0 14 3 6
Mangrove Cuckoo Coccyzus minor 0 1 0 2 2 3 1 1 1 0
Pheasant Cuckoo Dromococcyx phasianellus 0 1 0 0 0 0 0 0 0 0
Smooth-billed Ani Crotophaga ani 0 0 3 37 49 86 0 44 2 57
Squirrel Cuckoo Piaya cayana 2 10 2 2 0 5 1 0 0 0
Striped Cuckoo Tapera naevia 0 2 2 7 13 9 0 0 0 0
Emberizidae
Black-striped Sparrow Arremonops rufivirgatus 3 7 136 52 140 153 0 56 17 16
Blue-black Grassquit Volantinia jacarina 0 4 96 108 333 431 0 152 11 104
Orange-billed Sparrow Arremon aurantiirostris 27 16 6 0 0 0 17 0 6 0
Ruddy-breasted Seedeater Sporophila minuta 0 0 3 0 9 5 0 0 0 2
Slate-colored Seedeater Sporophila schistacea 0 0 0 0 1 1 0 0 0 0
Stripe-headed Brush-Finch Buarremon torquatus 2 0 0 0 0 0 0 0 0 0
185
Stripe-headed Sparrow Aimophila ruficauda 0 0 0 0 4 0 0 0 0 0
Thick-billed Seed-Finch Oryzoborus funereus 0 0 9 7 32 48 0 6 0 2
Variable Seedeater Sporophila americana 0 0 124 248 350 362 0 160 17 194
Wedge-tailed Grass-Finch Emberizoides herbicola 0 0 0 0 3 0 0 0 0 0
White-collared Seedeater Sporophila torqueola 0 0 2 14 45 17 0 3 0 5
Yellow-bellied Seedeater Sporophila nigricollis 0 0 3 3 4 3 0 0 0 1
Yellow-faced Grassquit Tiaris olivaceus 0 0 5 3 3 37 0 15 0 0
Falconidae
Barred Forest-Falcon Micrastur ruficollis 0 0 0 0 0 0 2 0 0 0
Bat Falcon Falco rufigularis 0 0 0 1 0 3 0 0 0 0
Collared Forest-Falcon Micrastur semitorquatus 2 1 0 0 0 0 6 0 0 0
Crested Caracara Caracara cheriway 0 0 0 1 19 14 0 4 4 3
Laughing Falcon Herpetotheres cachinnans 2 8 7 4 7 0 11 3 5 1
Yellow-headed Caracara Milvago chimachima 0 0 4 13 13 24 0 3 6 13
Formicariidae
Black-faced Antthrush Formicarius analis 7 3 0 0 0 0 8 0 0 0
Streak-chested Antpitta Hylopezus perspicillatus 5 1 0 0 0 0 13 0 0 0
Fringillidae
Spot-crowned Euphonia Euphonia imitans 26 20 9 7 0 0 36 4 6 0
Thick-billed Euphonia Euphonia laniirostris 0 14 9 28 13 5 0 10 4 16
White-vented Euphonia Euphonia minuta 5 6 0 0 0 0 2 0 0 0
Yellow-crowned Euphonia Euphonia luteicapilla 0 19 41 94 94 163 6 80 93 159
Yellow-throated Euphonia Euphonia hirundinacea 0 8 23 36 35 18 2 23 16 12
Furnariidae
Black-striped Woodcreeper Xiphorhynchus lachrymosus 19 12 0 0 0 0 25 0 4 0
Buff-throated Foliage-gleaner Automolus ochrolaemus 3 0 0 0 0 0 7 0 0 0
Cocoa Woodcreeper Xiphorhynchus susurrans 59 74 55 35 24 19 77 16 47 0
Long-tailed Woodcreeper Deconychura longicauda 2 1 0 0 0 0 3 0 0 0
Northern Barred-Woodcreeper Dendrocolaptes sanctithomae 14 10 5 0 0 0 22 0 1 0
Olivaceous Woodcreeper Sittasomus grisicapillus 0 0 0 0 0 0 0 0 1 0
Pale-breasted Spinetail Synallaxis albescens 0 0 0 0 2 53 0 1 0 0
Plain Xenops Xenops minutus 18 9 12 10 0 6 26 8 18 0
186
Ruddy Woodcreeper Dendrocincla homochroa 3 1 0 0 0 0 1 0 0 0
Scaly-throated Leaftosser Sclerurus guatemalensis 5 0 0 0 0 0 2 0 0 0
Slaty Spinetail Synallaxis brachyura 0 1 11 0 0 5 0 0 0 0
Spotted Woodcreeper Xiphorhynchus erythropygius 1 1 1 0 0 0 0 0 0 0
Streak-headed Woodcreeper Lepidocolaptes souleyetii 1 4 12 28 12 29 0 14 29 9
Striped Woodhaunter Hyloctistes subulatus 1 0 0 0 0 0 0 0 0 0
Tawny-winged Woodcreeper Dendrocincla anabatina 7 7 2 1 0 0 10 0 9 0
Wedge-billed Woodcreeper Glyphorhynchus spirurus 66 39 5 0 0 0 50 0 19 0
Galbulidae
Rufous-tailed Jacamar Galbula ruficauda 0 1 0 0 0 0 0 0 0 0
Hirundinidae
Bank Swallow Riparia riparia 0 0 0 0 3 0 0 1 0 7
Barn Swallow Hirundo rustica 0 0 0 1 0 0 0 0 0 0
Blue-and-white Swallow Pygochelidon cyanoleuca 0 0 0 2 0 13 0 0 0 0
Brown-chested Martin Progne tapera 0 0 0 0 0 0 0 0 0 7
Gray-breasted Martin Progne chalybea 0 0 0 3 15 5 0 1 0 23
Northern Rough-winged
Swallow Stelgidopteryx serripennis 0 0 0 8 21 13 0 2 0 22
Southern Rough-winged
Swallow Stelgidopteryx ruficolis 0 0 0 18 41 38 0 7 0 23
Icteridae
Baltimore Oriole Icterus galbula 0 3 20 31 20 30 0 16 18 26
Black-cowled Oriole Icterus prosthemelas 0 1 0 1 5 1 0 0 0 0
Bronzed Cowbird Molothrus aeneus 0 0 0 9 39 41 0 0 0 3
Eastern Meadowlark Sturnella magna 0 0 0 0 0 2 0 0 0 0
Giant Cowbird Molothrus oryzivorus 0 0 0 0 2 1 0 0 0 0
Great-tailed Grackle Quiscalus mexicanus 0 0 1 350 160 183 0 52 13 985
Melodious Blackbird Dives dives 0 0 1 88 9 45 0 7 1 61
Orchard Oriole Icterus spurius 0 3 0 7 1 0 0 1 0 1
Red-breasted Blackbird Sturnella militaris 0 0 0 0 0 2 0 0 0 0
Scarlet-rumped Cacique Cacicus uropygialis 84 83 38 25 0 2 78 3 53 0
Yellow-billed Cacique Amblycercus holosericeus 0 0 20 0 0 0 0 0 0 0
187
Yellow-tailed Oriole Icterus mesomelas 0 0 0 0 1 0 0 0 0 0
Incertae Sedis
Bananaquit Coereba flaveola 0 4 46 70 0 44 0 41 2 30
Black-and-white Becard Pachyramphus albogriseus 2 0 0 0 0 0 0 0 0 0
Black-crowned Tityra Tityra inquisitor 2 12 21 33 13 19 4 10 9 17
Cinnamon Becard Pachyramphus cinnamomeus 0 0 1 0 0 0 0 0 0 0
Masked Tityra Tityra semifasciata 2 22 11 28 3 15 5 5 10 12
Rose-throated Becard Pachyramphus aglaiae 1 8 4 2 0 1 3 0 7 0
Rufous Piha Lipaugus unirufus 32 23 0 0 0 0 49 0 1 0
Speckled Mourner Laniocera rufescens 1 0 0 0 0 0 1 0 0 0
Thrushlike Schiffornis Schiffornis turdina 12 2 0 0 0 0 6 0 0 0
White-winged Becard Pachyramphus polychopterus 0 3 1 2 0 5 1 0 7 0
Jacanidae
Northern Jacana Jacana spinosa 0 0 0 2 0 33 0 0 0 7
Momotidae
Blue-crowned Motmot Momotus momota 50 58 37 25 0 21 25 17 18 0
Nyctibiidae
Common Potoo Nyctibius griseus 0 0 0 0 0 0 1 0 0 0
Odontophoridae
Marbled Wood-Quail Odontophorus gujanensis 11 11 3 3 0 0 2 0 3 0
Parulidae
American Redstart Setophaga ruticilla 2 1 0 1 0 1 1 0 1 0
Bay-breasted Warbler Dendroica castanea 0 0 1 0 0 0 0 0 0 0
Black-and-white Warbler Mniotilta varia 2 5 2 2 0 1 2 0 2 0
Blue-winged Warbler Vermivora pinus 0 0 0 1 0 0 0 0 0 0
Buff-rumped Warbler Phaeothlypis fulvicauda 18 14 1 1 0 0 3 0 0 1
Cerulean Warbler Dendroica cerulea 0 1 0 0 0 0 0 0 0 0
Chesnut-sided Warbler Dendroica pensylvanica 19 41 65 27 5 40 23 17 71 6
Common Yellowthroat Geothlypis trichas 0 0 0 1 0 2 0 0 0 0
Golden-winged Warbler Vermivora chrysoptera 1 1 1 0 0 0 1 0 0 0
Gray-crowned Yellowthroat Geothlypis poliocephala 0 0 2 0 0 6 0 0 0 0
Hooded Warbler Wilsonia citrina 0 0 0 2 0 0 0 0 0 1
188
Kentucky Warbler Oporornis formosus 1 0 4 0 0 0 0 0 1 0
MacGillivray's Warbler Oporornis tolmiei 0 0 0 0 0 1 0 0 0 0
Magnolia Warbler Dendroica magnolia 0 0 0 1 0 0 0 0 0 0
Mourning Warbler Oporornis philadelphia 0 0 5 0 1 4 0 0 1 0
Northern Waterthrush Seiurus noveboracensis 0 3 0 0 2 0 0 0 0 0
Ovenbird Seiurus aurocapilla 0 0 0 0 0 0 2 0 0 0
Prothonotary Warbler Protonotaria citrea 0 0 0 0 0 2 0 0 0 0
Tenessee Warbler Vermivora peregrina 9 20 82 34 25 50 7 27 41 20
Tropical Parula Parula pitiayumi 0 0 0 2 0 0 0 0 0 0
Wilson's Warbler Wilsonia pusilla 0 1 3 1 4 3 1 0 1 2
Worm-eating Warbler Helmitheros vermivorum 1 0 0 0 0 0 0 0 0 0
Yellow Warbler Dendroica petechia 0 1 10 50 68 47 0 19 30 55
Yellow-breasted Chat Icteria virens 0 0 1 0 0 0 0 0 0 0
Yellow-throated Warbler Dendroica dominica 0 0 2 2 0 0 0 0 0 0
Passeridae
House Sparrow Passer domesticus 0 0 0 0 0 0 0 0 0 65
Picidae
Golden-naped Woodpecker Melanerpes chrysauchen 58 68 26 22 3 11 53 16 17 0
Hoffmann's Woodpecker Melanerpes hoffmannii 0 0 1 1 3 1 0 2 2 0
Lineated Woodpecker Dryocopus lineatus 2 12 8 12 13 18 1 5 19 2
Olivaceous Piculet Picumnus olivaceus 0 7 22 13 0 9 5 10 6 0
Pale-billed Woodpecker Campephilus guatemalensis 19 23 1 1 0 0 48 1 1 0
Red-crowned Woodpecker Melanerpes rubricapillus 3 12 92 188 153 225 1 91 116 160
Red-rumped Woodpecker Veniliornis kirkii 1 0 0 0 0 0 2 0 0 0
Rufous-winged Woodpecker Piculus simplex 1 0 0 0 0 0 2 0 0 0
Pipridae
Blue-crowned Manakin Pipra coronata 151 140 23 0 0 0 144 0 27 0
Orange-collared Manakin Manacus aurantiacus 19 14 20 0 0 1 5 0 6 0
Red-capped Manakin Pipra mentalis 56 33 4 2 0 0 45 0 2 0
White-ruffed Manakin Corapipo leucorrhoa 1 0 2 0 0 0 1 0 0 0
Psittacidae
Blue-headed Parrot Pionus menstruus 0 0 0 4 0 0 0 0 0 15
189
Brown-hooded Parrot Pionopsitta haematotis 12 15 0 0 0 8 33 0 0 0
Crimson-fronted Parakeet Aratinga finschi 8 11 8 77 10 106 11 16 37 55
Mealy Parrot Amazona farinosa 47 74 4 52 2 8 70 19 7 19
Orange-chinned Parakeet Brotegeris jugularis 4 29 71 194 121 188 0 105 197 287
Orange-fronted Parakeet Aratinga canicularis 0 0 0 4 0 0 0 0 0 0
Red-fronted Parakeet Touit costaricensis 0 0 3 0 0 0 0 0 0 0
Red-lored Parrot Amazona autumnalis 49 37 4 129 2 10 42 53 15 66
Scarlet Macaw Ara macao 0 0 0 50 0 0 0 0 2 64
White-crowned Parrot Pionus senilis 67 103 128 188 34 101 57 59 73 71
Rallidae
Gray-necked Wood-Rail Aramides cajanea 2 5 13 16 36 8 8 2 2 7
Purple Gallinule Porphyrio martinica 0 0 0 0 3 8 0 0 0 10
White-throated Crake Laterallus albigularis 0 0 0 0 0 2 0 0 0 0
Ramphastidae
Chesnut-mandibled Toucan Ramphastos swainsonii 130 206 110 77 12 64 177 114 54 14
Fiery-billed Aracari Pteroglossus frantzii 71 99 57 63 0 43 63 64 39 9
Strigidae
Crested Owl Lophostrix cristata 0 5 0 0 0 0 0 0 0 0
Ferruginous Pygmy-Owl Glaucidium brasilianum 0 0 0 5 0 0 0 0 0 3
Mottled Owl Ciccaba virgata 1 1 0 0 0 0 0 0 0 0
Spectacled Owl Pulsatrix perspicillata 2 0 0 6 0 0 2 0 0 0
Striped Owl Pseudoscops clamator 0 0 0 0 1 0 0 0 0 0
Vermiculated Screech-Owl Megascops guatemalae 0 0 0 0 0 0 1 0 0 0
Sylviidae
Long-billed Gnatwren Ramphocaenus melanurus 46 33 12 0 0 0 50 1 10 0
Tropical Gnatcatcher Polioptila plumbea 8 28 35 14 9 0 10 4 36 0
Thamnophilidae
Bare-crowned Antbird Gymnocichla nudiceps 4 3 0 0 0 0 6 0 0 0
Barred Antshrike Thamnophilus doliatus 2 12 54 19 4 29 0 29 15 1
Bicolored Antbird Gymnopithys leucaspis 3 0 0 0 0 0 6 0 0 0
Black-hooded Antshrike Thamnophilus bridgesi 139 154 120 7 2 0 183 18 54 1
Chesnut-backed Antbird Myrmeciza exsul 202 183 108 13 7 1 292 8 66 0
190
Dot-winged Antwren Microrhopias quixensis 25 9 2 0 0 0 23 0 0 0
Dusky Antbird Cercomacra tyrannina 14 8 2 0 0 0 12 0 2 0
Great Antshrike Taraba major 0 15 16 1 0 2 0 0 2 0
Plain Antvireo Dysithamnus mentalis 24 5 2 0 0 0 21 0 0 0
Russet Antshrike Thamnistes anabatinus 4 5 0 0 0 0 2 0 0 0
Slaty Antwren Myrmotherula schisticolor 3 1 2 0 0 0 9 0 0 0
Thraupidae
Bay-headed Tanager Tangara gyrola 4 1 12 6 0 5 1 0 0 0
Blue Dacnis Dacnis cayana 0 14 8 14 0 16 2 9 2 7
Blue-gray Tanager Thraupis episcopus 1 3 129 343 80 223 0 202 105 349
Cherries Tanager Ramphocelus costaricensis 0 44 460 445 141 291 0 342 85 332
Golden-hooded Tanager Tangara larvata 25 66 129 98 11 123 9 100 40 57
Gray-headed Tanager Eucometis penicillata 87 81 41 14 0 4 66 0 31 0
Green Honeycreeper Chlorophanes spiza 7 3 7 16 0 2 2 2 1 0
Palm Tanager Thraupis palmarum 0 2 117 288 98 113 0 191 23 269
Red-crowned Ant-Tanager Habia rubica 4 0 0 0 0 0 0 0 0 0
Red-legged Honeycreeper Cyanerpes cyaneus 4 8 19 37 0 23 7 25 6 6
Scarlet Tanager Piranga olivacea 0 2 4 3 2 5 1 0 0 0
Scarlet-thighed Dacnis Dacnis venusta 0 3 3 1 0 0 0 0 0 0
Shining Honeycreeper Cyanerpes lucidus 0 10 11 29 0 3 0 4 1 1
Summer Tanager Piranga rubra 8 16 22 30 6 12 6 6 21 3
Western Tanager Piranga ludoviciana 0 3 0 0 0 0 0 0 0 0
White-lineated Tanager Tachyphonus rufus 0 0 0 0 4 0 0 0 0 0
White-shouldered Tanager Tachyphonus luctuosus 24 1 0 0 0 0 25 0 4 0
White-throated Shrike-Tanager Lanio leucothorax 35 6 0 0 0 0 55 0 0 0
Threskiornithidae
White Ibis Eudocimmus albus 0 0 0 0 0 26 2 0 0 2
Tinamidae
Great Tinamou Tinamus major 33 17 9 0 1 0 46 0 1 0
Little Tinamou Crypturellus soui 14 13 8 1 0 0 25 1 4 0
Trochilidae
191
Snowy-bellied Hummingbird Amazilia edward 0 1 0 0 0 0 0 0 0 0
Band-tailed Barbthroat Threnetes ruckeri 7 20 2 1 0 0 9 0 7 0
Blue-tailed Hummingbird Amazilia cyanura 0 0 2 0 0 0 0 0 0 0
Blue-throated Goldentail Hylocharis eliciae 27 38 48 27 14 8 25 16 64 5
Bronzy Hermit Glaucis aeneus 0 19 1 0 0 0 1 0 0 0
Charming Hummingbird Amazilia decora 0 5 4 7 0 0 2 2 0 0
Garden Emerald Chlorostilbon assimilis 0 0 5 13 1 4 0 4 0 5
Green-breasted Mango Anthracothorax prevostii 0 2 0 14 0 1 0 2 0 1
Green-crowned Brilliant Heliodoxa jacuna 1 1 0 0 0 0 0 0 0 0
Long-billed Hermit Phaethornis logirostris 56 37 6 14 1 0 45 1 9 0
Long-billed Starthroat Heliomaster longirostris 0 5 5 5 1 2 0 0 5 0
Mangrove Hummingbird Amazilia boucardi 0 0 1 1 0 0 0 0 0 0
Purple-crowned Fairy Heliothryx barroti 11 20 18 25 1 5 9 20 9 2
Ruby-throated Hummingbird Archilochus colubris 0 0 5 2 0 1 0 0 0 0
Rufous-tailed Hummingbird Amazilia tzacatl 1 8 36 65 15 37 0 49 11 35
Scaly-breasted Hummingbird Phaechroa cuvierii 5 5 10 6 0 2 12 4 5 1
Scintillant Hummingbird Selasphorus scintilla 0 0 0 0 0 0 0 1 0 0
Steely-vented Hummingbird Amazilia saucerrottei 0 1 2 0 0 0 0 0 0 0
Stripe-throated Hermit Phaethornis striigularis 13 30 8 7 0 0 10 3 7 2
Violet Sabrewing Campylopterus hemileucurus 1 2 0 0 0 0 1 0 0 0
Violet-crowned Woodnymph Thalurania colombica 33 25 10 4 0 2 27 4 10 0
Violet-headed Hummingbird Klais guimeti 3 4 0 4 1 1 4 0 2 1
White-crested Coquette Lophornis adorabilis 1 1 2 5 0 0 2 0 0 0
White-necked Jacobin Florisuga mellivora 6 15 21 29 3 11 2 15 10 3
White-tipped Sicklebill Eutoxeres aquila 3 0 0 0 0 0 0 0 0 0
Troglodytidae
Black-bellied Wren Thryothorus fasciatoventris 2 30 13 0 0 0 0 2 0 0
House Wren Troglodytes aedon 0 0 45 308 348 157 0 180 77 254
Plain Wren Thryothorus modestus 0 4 60 73 133 74 0 81 19 39
Riverside Wren Thryothorus semibadius 97 184 95 25 2 5 83 27 25 3
Rufous-breasted Wren Thryothorus rutilus 0 4 0 1 0 0 0 0 0 0
192
Rufous-naped Wren Campylorhynchus rufinucha 0 0 0 5 0 2 0 0 0 22
Scaly-breasted Wren Microcerculus marginatus 18 0 0 0 0 0 20 0 0 0
White-breasted Wood-Wren Henicorhina leucosticta 1 0 0 0 0 0 1 0 0 0
Trogonidae
Baird's Trogon Trogon bairdii 12 6 3 1 0 0 5 0 5 0
Black-throated Trogon Trogon rufus 19 18 0 1 0 0 27 0 3 0
Slaty-tailed Trogon Trogon massena 36 30 22 18 0 6 26 6 23 0
Violaceous Trogon Trogon violaceus 7 16 18 10 0 2 5 3 6 0
Turdidae
Clay-colored Robin Turdus grayi 0 11 47 268 21 56 0 94 2 212
Swainson's Thrush Catharus ustulatus 21 43 26 11 1 10 10 11 8 4
White-throated Robin Turdus assimilis 1 0 0 0 0 0 0 0 0 0
Tyrannidae
Acadian Flycatcher Empidonax virescens 0 2 1 0 0 0 1 0 1 0
Alder Flycatcher Empidonax alnorum 0 0 3 0 0 3 0 0 0 0
Black-tailed Flycatcher Myiobius atricaudus 11 11 0 0 0 0 6 0 0 0
Boat-billed Flycatcher Megarhynchus pitangua 3 13 16 34 29 43 4 20 34 26
Bright-rumped Attila Attila spadiceus 22 14 19 17 15 12 14 16 18 6
Common Tody-Flycatcher Todirostrum cinereum 0 0 9 27 3 26 0 11 4 25
Dusky-capped Flycatcher Myiarchus tuberculifer 1 7 3 2 5 4 2 1 14 0
Eastern Wood-Pewee Contopus virens 0 0 0 1 0 12 0 0 1 0
Eye-ringed Flatbill Rhynchocyclus brevirostris 4 0 0 0 0 0 3 0 0 0
Golden-crowned Spadebill Platyrinchus coronatus 18 5 0 0 0 0 38 0 0 0
Gray Kingbird Tyrannus dominicensis 0 0 0 0 1 0 0 0 0 0
Gray-capped Flycatcher Myiozetetes granadensis 1 1 10 37 0 22 0 21 2 39
Great Crested Flycatcher Myiarchus crinitus 8 20 30 21 14 12 11 26 28 3
Great Kiskadee Pitangus sulphuratus 4 9 92 250 214 236 0 168 50 329
Greenish Elaenia Myiopagis viridicata 0 3 3 1 0 0 0 0 0 0
Least Flycatcher Empidonax minimus 0 2 1 0 0 0 0 0 1 0
Lesser Elaenia Elaenia chiriquensis 0 0 0 1 0 4 0 0 0 0
Northern Bentbill Oncostoma cinereigulare 2 2 4 0 0 0 1 1 1 0
Northern-scrub Flycatcher Sublegatus arenarum 0 0 0 0 0 0 0 0 1 0
193
Ochre-bellied Flycatcher Mionectes oleagineus 29 24 3 0 0 0 58 0 12 0
Olive-sided Flycatcher Contopus cooperi 1 1 2 1 0 4 0 0 1 0
Paltry Tyrannulet Zimmerius vilissimus 1 5 7 5 2 8 0 9 8 2
Panama Flycatcher Myiarchus panamensis 0 0 1 0 0 0 1 0 0 0
Piratic Flycatcher Legatus leucophaius 56 57 74 144 36 133 35 100 56 124
Royal Flycatcher Onychorhynchus coronatus 0 6 0 0 0 0 0 0 0 0
Ruddy-tailed Flycatcher Terenotriccus erythrurus 9 5 0 0 0 0 18 0 0 0
Rufous Mourner Rhytipterna holerythra 16 8 0 0 0 0 22 0 0 0
Rusty-marginated Flycatcher Myiozetetes cayanensis 0 0 0 0 0 0 0 0 0 2
Scissor-tailed Flycatcher Tyrannus forficatus 0 0 1 2 0 3 0 4 0 0
Slate-headed Tody-Flycatcher Poecilotriccus sylvia 2 0 4 4 0 0 2 0 0 0
Social Flycatcher Myiozetetes similis 0 0 22 95 6 54 0 59 5 111
Southern Beardless-Tyrannulet Camptostoma obsoletum 0 3 8 14 0 5 2 10 4 1
Streaked Flycatcher Myiodynastes maculatus 3 22 20 46 72 58 0 34 99 55
Sulphur-bellied Flycatcher Myiodynastes luterventris 1 2 3 6 0 3 1 0 3 2
Sulphur-rumped Flycatcher Myiobius sulphureipygius 12 1 0 0 0 0 10 0 0 0
Tropical Kingbird Tyrannus melancholicus 0 2 81 237 134 180 0 202 23 261
Tropical Pewee Contopus cinereus 0 1 3 1 0 13 0 0 2 2
Western Kingbird Tyrannus verticalis 0 0 3 9 31 17 0 10 0 8
Western Wood-Pewee Contopus sordidulus 0 1 2 0 0 6 0 0 0 0
White-throated Spadebill Platyrinchus mystaceus 2 0 0 0 0 0 0 0 0 0
Yellow Tyrannulet Capsiempis flaveola 0 1 10 13 16 5 0 3 2 3
Yellow-bellied Elaenia Elaenia flavogaster 0 0 11 71 8 83 0 59 2 28
Yellow-bellied Flycatcher Empidonax flaviventris 2 1 3 2 1 6 2 2 3 1
Yellow-bellied Tyrannulet Ornithion semiflavum 0 4 3 2 0 1 0 1 0 0
Yellow-crowned Tyrannulet Tyrannulus elatus 0 11 1 2 0 4 0 1 1 0
Yellow-olive Flycatcher Tolmomyias sulphurescens 0 10 7 7 3 0 2 2 7 0
Vireonidae
Green Shrike-Vireo Vireolanius pulchellus 5 0 0 0 0 0 6 0 0 0
Lesser Greenlet Hylophilus decurtatus 91 70 45 12 1 5 82 14 40 1
Philadelphia Vireo Vireo philadelphicus 1 8 20 16 4 18 5 13 9 2
Red-eyed Vireo Vireo olivaceus 0 0 0 1 0 3 1 0 0 0
194
Scrub Greenlet Hylophilus flavipes 0 0 0 6 4 1 0 1 0 0
Tawny-crowned Greenlet Hylophilus ochraceiceps 65 13 1 0 0 0 65 0 0 0
Yellow-green Vireo Vireo flavoviridis 0 2 3 2 0 1 0 1 0 0
Yellow-throated Vireo Vireo flavifrons 1 3 5 4 1 5 1 0 3 2
BR = Biological Reserves, FE = Forest Edges, FF = Forest Fallows, HG = Homegardens, OP = Oil palm, ASP = Agrosilvopastoral systems, WR =
Wildlife Refuges, RT = Residential Tourism Projects, TP = Tree Plantations, VI = Villages
195

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Alvaro Redondo Dissertation April 2010

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Effects of Land-Use Change on the Conservation of Bird Species in the

Path of the Tapir Biological Corridor, Costa Rica

Presented to the Faculty of the Graduate School

Alvaro Redondo Brenes

Dissertation Director: Dr. Florencia Montagnini and Dr. Chadwick Oliver

edges, villages, residential tourism projects, homegardens, agrosilvopastoral systems, and

insectivores, 21.2% frugivores, 9% nectarivores, and carnivores and granivores 8% each.

threatened species, respectively, were forest-dependent species. It is suggested managing

Payment for environmental services, ecotourism, reforestation, environmental education

Presented to the Faculty of the Graduate School

Dissertation Director: Dr. Florencia Montagnini and Dr. Chadwick Oliver

Table and Figures……………………………………………………………………. vii

Chapter 1: A Review of the Effects of Land-Use Change and Habitat

Chapter 3: Avian diversity of ten land-use types in a human-modified biological

Chapter 4: Response of bird guilds to ten land-use types in a human-modified

Chapter 5: Conservation of threatened and endemic bird species in a human-

Summary and Conclusions………………………………………………………….. 177

Chapter 2: Physical, Political, and Socio-Economic Aspects of the Path of the

Chapter 3: Avian diversity of ten land-use types in a human-modified biological

Chapter 4: Response of bird guilds to ten land-use types in a human-modified

Chapter 5: Conservation of threatened and endemic bird species in a human-

deeply grateful to my other committee members: Chad Oliver, Jonathan Reuning-Scherer

dissertation thesis and time at Yale more meaningful and enjoyable.

In Costa Rica, especially in the Path of the Tapir Biological Corridor, I am

Aside from ASANA and Hacienda Baru, several families/organizations permitted

provided invaluable bird information for my dissertation.

Several governmental and non-governmental organizations also contributed to my

Municipalities, local organizations and of course local people.

surrounding areas. These areas and their people are unique.

A Review of the Effects of Land-Use Change and Habitat Fragmentation on Wildlife

The establishment of protected areas is one means of counteracting some of the

threats to biodiversity such as habitat loss and fragmentation (Sanchez-Azofeifa et al

been an exponential growth in the number of protected areas worldwide, particularly in

tropical countries where biodiversity is greatest (Zimmerer et al. 2004; Naughton-Treves

classifications such as the IUCN Conservation categories (Rodrigues et al. 2004).

areas are jeopardizing biological conservation (Powell et al. 2000; Sanchez-Azofeifa et

human-modified landscapes in assessments of biodiversity and ecosystem services,

planning of corridors, establishment of buffer zones, and restoration of degraded lands

(Chazdon et al. 2009).

of Central America (TNC-ASANA 2000), but whose biodiversity is at risk because of

approximately 10,000 people in 55 rural communities.

biological reserves, wildlife refuges, homegardens, agrosilvopastoral systems, forest

Chapter (1) presents a theoretical background to understand the effects of

fragmentation, land-use change, and human-dominated landscapes on the conservation of

to species of conservation concern (e.g endemic and threatened species). Furthermore,

Fragmentation and Land Use Change effects on wildlife species

Destruction or alteration of natural habitats by people because of land-use change,

conserving biodiversity in fragmented landscapes now dominated by human land use

isolated patches (Saunders et al. 1991). As a consequence of these changes at the

(1) the elimination or dangerous reduction of populations of large, wide-ranging species,

including many top carnivores;

species or destabilize communities;

effects such as altered microclimates or invasive species; and

agents – such as pollination, seed dispersal, predator-prey interactions, and nutrient

The response of wildlife species to habitat destruction and fragmentation is

patterns, dispersal capabilities, nesting and shelter requirements, as well as tolerance to

habitat disturbance and sensitivity to altered microclimates (Bennett 2003). Based on

these animal characteristics, the species vulnerable to forest fragmentation can be

organized into six groups (Groom et al. 2005):

heterogeneous landscapes such as amphibians and turtles are vulnerable to

Examples are carrion beetles in Amazonian forests being fragmented by pasture

that are unwilling to enter the matrix (Laurance et al. 2002).

(Lycaeides argyrognomon lotis), an endemic species of California, USA that requires