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B.IODIVERSITY SERIES - IMPACT STUDIES

Public Disclosure Authorized

Biodiversity
Conservation in the
Public Disclosure Authorized

Context of Tropical
Forest Management

Francis E. Putz
Kent H. Redford
John G. Robinson
Public Disclosure Authorized

Robert Fimbel'
Geoffrey M. Blate

September 2000

TheWorld Bank
 O THE WORLDBANKENVIRONMENT
DEPARTMENT

Biodiversity
Conservation in the
Context of Tropical
Forest Management
Francis E. Putz
Kent H. Redford
John G. Robinson
Robert Fimbel
Geoffrey M. Blate

September 2000

Papers in this series are not formal publications of the World Bank. They are circulated to encourage thought and discussion. The use
and citation of this paper should take this into account. The views expressed are those of the authors and should not be attributed to
the World Bank. Copies are available from the Environment Department, The World Bank, Room MC-5-126.
The International Bank for Reconstruction
and Development/THE WORLD BANK
1818 H Street, N.W.
Washington, D.C. 20433, U.S.A.

Manufactured in the United States of America

First printing September 2000

About the authors:

Francis E. Putz (fep@botany.ufl.edu) is Professor of Botany at the University of Florida; Kent H. Redford
(KHRedford@aol.com) is Director of Biodiversity Analysis and Coordination at WCS; John G. Robinson
(jrobinson@wcs.org) is Senior Vice President and Director for International Conservation at WCS; Robert
Fimbel (robertf@parks.wa.gov) is Chief Scientist for the Washington State Parks and Recreation Commis-
sion; and Geoffrey M. Blate (gblate@botany.ufl.edu) is a Ph.D. candidate at the University of Florida.

Substantial background material for this paper was provided by D. Nepstad, M. Guariguata, R. Noss, and
C. Peters. We also acknowledge the contributions of E. Bennett, J. Ewel, P. Frumhoff, E. Losos, K.
MacKinnnon, A. Moad, G. Platais, A. Plumptre, and R. Plumptre.

This document does not represent an endorsement of any particular World Bank policy by the Wildlife
Conservation Society.

'WC s
The Wildlife Conservation Society (WCS) is dedicated to being the most effective conservation organiza-
tion, protecting and promoting a world rich in wildlife and wilderness.

WCS manages more than 300 field projects in 53 countries and develops award-winning environmental
education programs for schools in the U.S. and abroad.

At its celebrated wildlife parks in New York-the Bronx Zoo, the New York Aquarium, and Central Park,
Queens and Prospect Park Wildlife Centers-WCS inspires over 4.5 million annual visitors to care about
wildlife and wild lands and to participate in their conservation.

To leam more about WCS, visit www.wcs.org.

Biodiversity Series
Impact Studies

These "Impact Studies" are a subset of the Biodiversity Series of the World Bank's
Environment Department Papers. Within this subset the broader question of what the
positive and negative impacts of human activities on biodiversity is addressed.

The following studies have been published in this series:

1. BiodiversityConservationin the Context of TropicalForestManagement

2. Hunting of Wildlifein TropicalForests-Implicationsfor Biodiversityand ForestPeoples
Contents
PREFACE V

ExEcurlvESUMMARY VIl

Chapter 1
Introduction 1

Chapter 2
Disaggregating "Biodiversity" 3

Chapter 3
Disaggregating "Logging" 7
Logging intensities 7
Log yarding methods 9
Logging and other silvicultural treatments 11
Reducing the impacts of logging and other silvicultural treatments 12

Chapter 4
Impacts of Forest Management on Biodiversity 15
Background 15
Landscape impacts (see Appendix I) 16
Ecosystem impacts (see Appendix II) 19
Community impacts (see Appendix III) 21
Species impacts (see Appendices V and VI) 22
Genetic impacts (see Appendix VII) 25

Chapter 5
Overview of Biodiversity Conservation in Relation to Logging and other
Silvicultural Treatments 27
Synthesis 27
Troubling assumptions 29
Conclusions 30

Chapter 6
Recommendations 33

BiodiversitySeries - Impact Studies iii

BiodiversityConservation in the Context of TropicalForestManagement

APPENDICES
I Impacts of Logging (and other silviculture treatments where noted) on the Landscape Component
of Biodiversity in Tropical Forests 35
II Impacts of Logging (and other silviculture treatments where noted) on the Ecosystem Component
of Biodiversity in Tropical Forests 39
III Impacts of Logging (and other silviculture treatments where noted) on the Community Compo-
nent of Biodiversity in Tropical Forests 43
IV Bird responses (density based on inter-site comparisons) to Logging in Tropical Forests,
Organized by Feeding Guild 51
V Impacts of Logging (and other silviculture treatments where noted) on the Species Component of
Biodiversity in Tropical Forests 53
VI Primate responses (by feeding guild) to Logging in Tropical Forests 57
VII Impacts of Logging (and other silviculture treatments where noted) on the Genetic Component of
Biodiversity in Tropical Forests 61

GLOSSARY 63

BIBLIOGRAPHY 67

Boxes
1 Few tropical forests are logged or burned only once 9
2 Enrichment planting 12
3 Is reduced-impact logging (RIL) cheaper? 13
4 The critical role of production forests in maintaining biodiversity in the tropics 17
5 Hunting in logged tropical forests 23
6 Lesser-known species-Marketing challenges, silvicultural impediments, or desirable
diversity? 24

Figures
1 Logging intensities (m 3 /ha) for tropical forests 8
2 Generalized ranges of direct biodiversity impacts of timber yarding methods used for tropical
forest logging as a function of level of technological sophistication required 10
3 Some impacts of logging on biodiversity as a function of distribution of logging activities and
logging intensity 18
4 Expected effects of a range of forest uses on the components of biodiversity 27
5 Generalized biodiversity impacts plotted against expected short-term financial returns to forest
owners or concessionaires 28
6 Generalized biodiversity impacts resulting from different approaches to forest management for
timber 29

Tables
1 Components and attributes of tropical forest biodiversity that might be influenced by logging
and other silvicultural activities 4

iv Environment Departrnent Papers

Preface
This paper was prepared as part of the World
Bank's Overlay Program and as a contribution
to the Bank's ongoing Forest Policy review.
The Overlays Program, launched by the World
Bank in partnership with bilateral donors and
NGOs, seeks to internalize global externalities
into national environmental planning and the
Bank's sector work, operations, and its
dialogue with governments and partners. It is
an iterative process, combining conceptual
studies, reviews of state-of-the-art techniques
for measuring and mitigating local, national
and global externalities, and testing these
concepts and tools in country-level studies as a
means of identifying good practices for
country planners and Bank task managers. The
results will help guide national actions to
conserve biodiversity, reduce greenhouse gas
emissions, and protect international waters.
The Overlays add a new dimension to
traditional sector economic planning by
analyzing environmental impacts and
opportunities to internalize global
externalities. This analysis launches the
"Impact Studies" which asks the broader
question of what the positive and negative
impacts of human activities are on
biodiversity. An analytical framework is used
which allows for the systematization of these
impacts across sectors and areas of particular
interest.
BiodiversitySeries- ImpactStudies
One of the most important forest uses is logging
or timber harvesting. Logging is the most
lucrative forest use and can cause severe direct
and indirect environmental impacts. This
analysis evaluates the impacts of logging on
biodiversity in tropical forests based on a
thorough review of the literature and on the
best expert opinion.
The study was commissioned from the Wildlife
Conservation Society (WCS).A draft paper was
prepared by WCS and the University of Florida
and was distributed to an expert review panel.
Supported by the Forest Policy Implementation
Review and Strategy Process, these reviewers
convened in Washington with the authors and
Bank staff in late February, 1999for a
productive two day workshop after which the
paper was revised.
Forests will continue to be harvested and timber
harvesting will continue to be an important
component of sustainable forest management.
This is a reality we cannot escape. Therefore, as
this practice continues we need to assure
ourselves that it is ecologically and
environmentally sustainable. The World Bank
has a responsibility to its clients to inform them
of best practices and what the potential
consequences of natural resources management
decisions are. This paper helps fulfill this
responsibility.
v
Executive Summary
Existing parks and protected areas are
cornerstones of biodiversity conservation but on
their own are inadequate to assure the
continued existence of a vast proportion of
tropical forest biodiversity. As a result, priority
must be given to ensuring that the greatest
possible amount of biodiversity is conserved
outside protected areas by altering harvest
patterns in these landscapes of resource
extraction. Of all forest uses within tropical
forest regions, logging or timber harvesting, is
the most important to influence because not
only is it the most lucrative, it also causes the
most severe direct and indirect environmental
impacts.
Evaluating the impacts of logging on
biodiversity in tropical forests is not as simple a
task as many would believe. It depends on a
thorough review of the literature and on the
best expert opinion due to the complexities
hidden under the seemingly simple rubrics
"logging" and "biodiversity." As a first step in
effectively analyzing the relationship between
biodiversity and logging, it is essential to begin
by disaggregating the terms "logging" and
"biodiversity."
"biodiversity" into components (landscapes,
ecosystems, communities, species/populations,
and genes) and attributes (structure,
composition, and function). It then
disaggregates "logging" by detailing the vast
range of activities subsumed under this term
including variation of logging intensities,
Biodiversity Series- ImpactStudies
logging methods, collateral damage, and
silvicultural approaches. Using the richness
present in both these terms, a framework for
considering the impacts of logging and other
forest management activities on the various
components and attributes of biodiversity is
presented. This framework is, in turn, used to
evaluate the extensive literature covering
different studies of logging in tropical forests.
findings:
1. All consumptive uses affect some
component or attribute of biodiversity,
commonly affecting not only the target
resource but other elements as well.
2. As a result, only fully protected areas will
conserve all components and attributes of
biodiversity.
3. Recognizing that all significant
interventions in natural forests have
biodiversity impacts, all silvicultural
decisions necessarily represent
compromises. Management for some goods
or services necessarily involves
management against others.
4. Different intensities and spatial patterns of
timber harvesting, along with other
silvicultural treatments, result in different
effects on the different components of
biodiversity.
5. Some components and attributes of
biodiversity are more sensitive than others
to forest management activities.
6. There are some forested areas and some
areas within forests, which should never be
logged.
vii
BiodiversityConservationin the Contextof TropicalForestManagement
7. Biodiversity objectives can and should be
established within production forests.
8. In many cases the logging itself is not the
major cause of biodiversity loss, but rather
the indirect effects, often promoted by the
presence of roads, (for example, hunting,
forest fires, and the likelihood of
deforestation) represent the major
environmental threats.
Unfortunately, due to the complexities hidden
under the seemingly simple rubrics "logging"
and "biodiversity", the answer to the question
"Is logging compatible with biodiversity
protection?" can only be the very unsatisfying
"It depends." The paper does not conclude
with uncritical support for sustainable forest
management of timber as a conservation
strategy. Such an endorsement is unwarranted
given widespread illegal logging in the tropics,
widespread frontier logging and logging of
viii
areas of high priority for biodiversity
protection, the persistence of poor logging
practices despite substantial efforts in research
and training, and the generally slow rate at
which most loggers are transforming
themselves from timber exploiters into forest
managers.
Nevertheless, even harshly treated forests
maintain more biodiversity than oil palm
plantations, cattle pastures, or cornfields. From
a biodiversity maintenance perspective, natural
forest management is preferable to virtually all
land-use practices other than complete
protection. Forests that are carefully managed
for timber will not replace protected areas as
storehouses of biodiversity, but they can be an
integral component of a conservation strategy
that encompasses a larger portion of the
landscape than is likely to be set aside for strict
protection.
EnvironmentDepartmentPapers
1 Introduction
Existing parks and protected areas are essential
for biodiversity conservation but inadequate on
their own to assure the continued existence of
the majority of natural landscapes, ecosystems,
communities, species and genotypes in tropical
forests. Even if the.oft-quoted goal of 10-12
percent protection were attained and the
reserved areas were appropriately located and
properly managed, up to 50 percent of tropical
species would be expected to go extinct during
the next few decades (Soule and Sanjayan 1998).
Similar but often less well documented losses of
the other components of biodiversity are
expected. Another way of considering this dire
prediction is that if biodiversity outside of
protected areas is neglected, thousands of
species are likely to disappear. Faced with this
bleak future, the first priority should be to
increase the area of forests under strict
protection while at the same time improving
reserve management. Mechanisms should be
developed to halt road building and commercial
logging in forest wilderness areas as well as in
centers of diversity and endemism. However,
promoting more biodiversity-sensitive
management of forests outside protected areas
is of almost equal priority, given the
conservation potential of these still vast areas.
Biodiversity conservation within forests
managed primarily for timber production, for
example, should be fostered by a combination of
not logging ecologically important areas and by
employing biodiversity-sensitive management
methods within production areas.
Decisions about which large forest areas should
be designated for strict protection, which areas
BiodiversitySeries - ImpactStudies
within managed forests should be protected,
and which approaches to silviculture to follow
within forests used for timber production
should be made on the basis of a wide range of
considerations. Forests differ in their
biodiversity value, in their capacity to support
different intensities of silvicultural use, in
pressures for conversion to non-forest use, and
in the abilities of the relevant institutions to
regulate their management. General conceptual
approaches to zoning forests for different uses
have been presented recently by various authors
including Noble and Dirzo (1997), and
Frumhoff and Losos (1998). Methods for
integrating conservation functions at different
geographical scales were reviewed recently by
Poiani and others (2000). This paper focuses on
the forests zoned for timber production. The
goals are:
1. To contribute to the development of a
heuristic construct for considering the range
of impacts of forestry activities on tropical
forest biodiversity at.the levels of
landscapes, ecosystems, communities,
species, and genes.
2. To indicate the topics on which further
research will be particularly useful in
evaluating the impacts of different forestry
activities on the various components and
attributes of biodiversity.
3. To suggest ways to mitigate the deleterious
impacts of forestry activities on tropical
forestbiodiversity.
In the search for land-use practices compatible
with biodiversity maintenance, many
BiodiversityConservation in the Context of TropicalForest Management
environmentalists have focused upon logging
[see Grieser Johns (1997) and Haworth (1999)
for comprehensive reviews of the environmental
impacts of logging in tropical forests]. This
emphasis is not surprising given that of all
forest uses, logging is often the most financially
lucrative and has the most severe
environmental impacts. The indirect effects of
logging, particularly increased hunting (for
example, Robinson and others 1999) and the
increased likelihood of deforestation due to
improved access, reviewed by Kaimowitz and
Angelsen (1998), have also been highlighted
recently. What has emerged from the dust, mud,
chainsaw noise, and diesel fumes is the idea
that conservation of some components of
biodiversity could be facilitated by
collaboration between loggers and
environmentalists. Some of the latter oppose the
idea of promoting better forest management as
a means for achieving overall conservation
goals (or at least oppose financial investment in
such efforts, Rice and others 1997, Bawa and
Seidler 1998, Bowles and others 1998). This
opposition notwithstanding, consideration of
the inevitability of logging in much of the
tropics, the many constraints on expansion of
nature reserves in tropical countries, the
challenges of protecting and managing the
parks already demarcated on paper, sovereignty
issues, development needs, and the implicit
adoption of the "use it or lose it" assumption,
2 Environment Department Papers
motivate other conservationists to continue
holding sustainable forest management (SFM)
as a worthy conservation goal in forests outside
of protected areas (for example, Dickinson and
others 1996, Chazdon 1998, Poore and others
1999, and Whitmore 1999).
To help elucidate some of the factors upon
which the compatibility of tropical forest
logging and biodiversity protection depend, this
paper first attempts to disaggregate the terms
"logging" and "biodiversity." The wide range of
logging intensities, logging methods, collateral
damage, and silvicultural approaches
appropriate for tropical forests is discussed. A
framework for considering the impacts of
logging and other forest management activities
on the various components (=landscapes,
ecosystems, communities, species/populations, and
genes) and attributes (=structure, composition, and
function) of biodiversity is presented. In the
main section of the paper, the components and
attributes of biodiversity are used to review the
impacts of logging and other silvicultural
activities on tropical forests. To promote
readability of the text, much of the
bibliographical review is presented in
appendices organized by the components and
attributes of biodiversity. The paper concludes
by suggesting ways to enhance the
compatibility of forest management for timber
and biodiversity maintenance in tropical forests.
2 Disaggregating "Biodiversity"
Biodiversity refers to the natural variety and
variability among living organisms, the
ecological complexes in which they naturally
occur, and the ways in which they interact with
each other and with the physical environment.
This definition and the elucidation below is
based upon OTA (1987),Noss (1990) and
Redford and Richter (1999).Climate, geology
and physiography all exert considerable
influence on broad spatial patterns of biotic
variety; local ecosystems and their biological
components are further modified by
environmental variation (such as local climatic
and streamflow fluctuations) and ecological
interactions. This natural variety and variability
is distinguished from biotic patterns or
conditions formed under the influence of
human-mediated species introductions and
substantially human-altered environmental
processes and selection regimes (Bailey 1996,
Noss and Cooperrider 1994).
In discussions of diversity there is always the
lurking danger of assuming that "more is
better." Were this the case, plowing roads
through mature forests, clearcutting patches,
and introducing alien species would all be
reasonable. Although decisions about which
species or community-types are most valuable
are not straightforward, rarity is one obvious
dimension that needs to be considered.
Biologicaldiversity can be measured in terms of
different components (landscape, ecosystem,
community, population/species, and genetic),
each of which has structural, compositional, and
functional attributes (Table1). Structure refers to
BiodiversitySeries - ImpactSthdies
the physical organization or pattern of the
elements. Compositionrefers to the identity and
variety of elements in each of the biodiversity
components. Functionrefers to ecological and
evolutionary processes acting among the
elements.
Diversity of the landscapecomponentrefers to
regional mosaics of land uses, land forms, and
ecosystem types. Structure of this component
could be described on the basis of the areas of
different habitat patches, through the perimeter-
area relations of these habitat patches, or on the
basis of interpatch linkages. Landscape
compositionpertains to the identity, distribution,
and proportions of different habitat types.
Landscapefunction refers to issues such as patch
persistence and interpatch flows of energy,
species, and other resources.
Diversity of the ecosystemcomponentrefers to
interactions between members of a biological
community and their abiotic environment.
Structureof this component might be measured
through vegetative biomass and soil structural
properties. Ecosystem compositionrefers to
features such as biogeochemical standing
stocks. Ecosystemjiunction pertains to processes
including biogeochemical and hydrological
cycling.
Diversity of the community componentrefers to the
guilds, functional groups, and patch types
occurring in the same area and strongly
interacting through trophic and spatial biotic
relationships. Structure of this component
3
BiodiversityConservation in the Context of TropicalForest Management

Table I Components and attributes of tropical forest biodiversity that might be influenced by logging and
other silvicultural activities

Organizedbythe componentsandattributesof biodiversity

Components Struaure Composition Function
Landscape Sizeand spatialdistribution of Identity, distribution, and Habitat patch persistenceand
habitat patches(e.g.,seral proportion of habitattypes turnover rates; energyflow rates;
stagediversity and area); and multi-habitat landscape disturbanceprocesses(e.g., extent,
physiognomy;perimeter-area types; collectivepatterns of frequency,andintensity of fires);
relations; patchjuxtaposition speciesdistributions humanlanduse trends; erosion
and connectivity;fragmentation rates;geomorphicand hydrologic
processes

Ecosystem Soil(substrate)characteristics; Biogeochemicalstocks; Biogeochemicaland hydrological

vegetationbiomass,basalarea lifeform proportions cycling;energyflux; productivity;
andvertical complexity,density flows of speciesbetween patches;
anddistribution of snagsand local climate impacts
fallen logs

Community Foliagedensityand layering; Relativeabundanceof Patchdynamicsand other

canopyopennessandgap
proportions; trophic andfood
web structures
speciesand guilds;richness
and diversity indices;
proportions of endemic,
exotic, threatened,and
endangeredspecies;
proportions of specialistsvs.
generalists
successional processes;
colonizationandextinction rates;
pollination,herbivory, parasitism,
seeddispersalandpredation rates;
phenology

Speciesl Sexand age/sizeratios; range Speciesabundance Demographicprocesses(e.g.,

Population anddispersion;infraspecific distributions, biomass,or survivorship,fertility, recruitment,
morphologicalvariation density;frequency; anddispersal);growth rates;
importance or cover value phenology

6enetic Effectivepopulationsize; Allelic diversity; presenceof Geneflow; inbreedingdepression;

heterozygosity;polymorphisms; rare alleles;frequencyof rates of outbreeding,genetic drift
generationoverlap;heritability deleteriousalletes and mutation;selection intensity;
dysgenicselection
Note:Modified from Noss (1990)and Redford and Richter (1999).

includes consideration of vegetative be measured through population age structure

physiognomy and trophic structure.
Community composition refers to relative
abundances of species and guilds. Community
functions include flows between patch types,
disturbance regimes (such as fires and floods),
successional processes, and species interactions.
Diversity of the species/population component refers
to the variety of living species and their
component populations at the local, regional, or
global scale. Structure of this component might
or species abundance distributions. Species/
population composition pertains to which
particular species are present. Function of this
component refers to demographic processes
such as recruitment and death.
Diversity of the genetic component refers to the
variability within a species, as measured by the
variation in genes within a particular species,
subspecies, or population. Structure of this
component can be expressed on the basis of

4 Environment Department Papers

 Disaggregating"Biodiversity"

heterozygosity or genetic distances between and in what proportions. Genetic functions can
populations in different patches (that is, be expressed on the basis of gene flow, genetic
metapopulation genetic structure). Genetic drift, or loss of allelic diversity in small, isolated
compositionrefers to which alleles are present populations.

Biodiversity Series - Impact Studies 5

3 Disaggregating "Logging"
When properly planned and conducted, logging
(=timber harvesting) is an integral component
of forest management systems designed to
promote sustained timber yields (STY), or the
more all-encompassing goal of sustainable
forest management (SFM). Unfortunately,
logging in tropical forests all-too-often
represents a timber "mining" activity carried
out without regard for the renewability of this
natural resource (for example, Putz and others
2000). Other silvicultural treatments designed to
promote sustainability are often prescribed but
are rarely applied. Due mostly to the desire to
obtain voluntary third-party certification of
good management from the Forest Stewardship
Council (FSC), the transition from forest mining
to forest management has finally started to
occur in some areas in the tropics (Nittler and
Nash 1999). Despite this progress, even within
certified forests there are questions about how
to most effectively and efficiently minimize the
deleterious environmental impacts of logging
and other silvicultural activities. This paper
addresses many of these questions by reviewing
the state of knowledge about biodiversity
maintenance in exploited and managed tropical
forests.
It is critical to recognize that forest interventions
of all types, from harvesting of fruits for home
consumption to clearcutting for timber, all have
impacts on forests that need to be understood
and often deserve mitigation (Peters 1996).
Nevertheless, logging is often the most
damaging and generally the most financially
lucrative of such forest interventions (Pearce
and others 1999). Unfortunately, discussions
Biodiversity Series - Impact Studies
about the compatibility of logging with
biodiversity conservation are complicated
because logging is carried out over a huge range
of intensities, using a variety of techniques
which may be applied carefully or in ways that
result in a great deal of avoidable damage. The
following sections focus on the issues of
harvesting intensities, yarding methods (=how
timber is extracted from the stump to haul
roads), and ways of reducing logging damage.
A brief overview of other silvicultural
treatments used in timber stand management is
also provided.
Logging intensities
Trying to conclude anything about the
Try ity oflogging aboutethe
compatibility of logging and biodiversity
maintenance is made complicated by the wide
range of logging intensities, yarding methods,
and accompanying forest management practices
to which tropical forests are subjected. The
simple observation that logging intensities span
more than two orders of magnitude (<1 m 3 /ha
to >100 m 3 /ha) illustrates the challenge of
generalizing (Figure 1).
Even within the same forest management unit,
logging intensities vary greatly at different
spatial scales. At a small scale (1-10 ha), the
typical aggregated distributions of tropical trees
(Hubbell 1979) leads to locally severe logging
impacts unless harvesting controls are
implemented (that is, "tree-marking rules" are
followed, Uhl and Vieira 1989, Crome and
others 1992, Cannon and others 1994). The
localized but severe direct impacts of roads, log
7
BiodiversityConservation in the Context of TropicalForest Management

landings, and skid trails hardly need to be
emphasized (Guariguata and Dupuy 1997). At a
slightly larger scale (10-100 ha), stands vary in
stocking of commercial species and in their
accessibility due to terrain or edaphic factors.
Where logging is carried out in areas designated
for each year of management (=annual coupes),
logging impacts tend to be aggregated at larger
scales.
Logging intensities also vary over time, tending
to increase with local timber shortages,
improved access, and greater willingness of
markets to accept lesser-known species
(Plumptre 1996). For example, in a remote part
of the Bolivian Amazon loggers may harvest an
average of only 0.3 m 3 /ha of 1-3 species
(Gullison and Hardner 1993) whereas in
similarly stocked stands in the more accessible
eastern portions of Amazonian Brazil, it is likely
that 30-50 m3 of 20-30 species would be
harvested Uohns and others 1996).
The substantial number of studies conducted on
the impacts of logging in tropical forests all
concluded that soil impacts and damage to the
residual forest all increase with increasing
logging intensity (Ewel and Conde 1976, Sist
and others 1998a). Proportions of both soil and
residual trees damaged by logging range from 5
to 50 percent, depending on harvesting
intensity, yarding method, and the care with
which the operations are carried out.
Interpretation of data pertaining to the
relationship between logging intensity and
residual stand damage is complicated by
concomitant change in residual stand density; at
the extreme, there is no residual stand in

Figure I Loggingintensities (m3/ha) for tropical forests. In most of these studies, as in most logging areas in
the tropics, felling was with chainsaws and yarding with bulldozers or articulated skidders with rubber tires.

< I rn/ha 50 m 3/ha 00 m 3 /ha 150 m 3/ha

Bolivia Venezuela Indonesia Malaysia

(Gullisonand Hardner 1993) (Kammesheidt1998) (Bertaultand Sist 1997) (Pinardand Putz 1996)

Suriname Costa Rica Philippines

(Hendrison 1990) (Webb 1998) (Nicholson 1979)

Venezuela
(Mason 1996)

Australia
(Cromeet al. 1992)

Brazil Malaysia
Oohns et al. 1996) (Pinard and Putz 1996)

Australia
(Crome et al. 1997)

Uganda
(Chapman and Chapman 1997)

8 Environment Departnent Papers

 Disaggregating "Logging"

clearcuts. Further complicating assumptions at the landscape, ecosystem, community,

about logging damage is the fact that few
tropical forests are logged (or burned) only once
(Box 1).
Log yarding methods
In this paper, particular attention is paid to the
yarding phase of the timber harvesting process
because much of the direct damage to tropical
forest caused by logging occurs while logs are
being extracted from the stump to roads or
riversides from which they are then hauled or
towed out of the forest. Yarding methods
utilized in tropical forests range in technological
sophistication, the collateral damages with
which they are associated, and in yarding costs
(Conway 1982). Accurate cost figures for
yarding are difficult to obtain and problematic
to calculate (for example, should impacts on
future timber yields be included, and, if so, at
what discount rate?). For the same volume of
timber yarded, the range in biodiversity impacts
population/species, and genetic levels varies
greatly along the continuum of technological
sophistication of yarding methods that stretches
from manual extraction to the use of helicopters
(Figure 2). Note that Figure 2 hides a great deal
of relevant variation. In particular, attempts to
include another dimension expressing yarding
costs (per cubic meter yarded to the roadside)
have so far failed due to the multitude of factors
that need to be considered, including labor
costs, technical capacity, and discount rates,
along with the values assigned to future harvest
yields, non-timber forest products, biodiversity
protection, and ecosystem services.
Yarding can be carried out in an environmen-
tally/silviculturally sensitive manner, or can be
extremely destructive. The rankings of
environmental impacts on Figure 2 are just
suggestions of the typical amounts of damage
associated with different yarding techniques.

Box 1
Few tropical forests are logged or burned only once
Logging and wildfires in tropical forests are causally linked (Holdsworth and Uhl 1997) and also share the
characteristic of being self-promoting processes. Although the environmental damage resulting from unplanned
logging of primary forest by untrained crews is justifiably receiving attention from researchers and policy-
makers, few logged stands are allowed to recover for the full cutting cycle or rotation stipulated in their man-
agement plans. Instead, forests are repeatedly entered by loggers who sequentially 'high-grade" the best re-
maining timber. Although the costs of road building are substantial, the first cut is generally the most lucrative.
As more timber species enter commercial markets, and mills begin to accept smaller and lower quality logs,
incentives for multiple entry logging increase. The same logging firms are sometimes involved in a series of
entries into the same forest, but more often firms with lower operating costs and smaller equipment take
advantage of the increased access provided during the first harvest. A familiar adage in Amazonia is "ma-
hogany builds the roads." Reliable data on frequencies of multiple entry logging, either legal due to waivers
issued by governmental agencies or illegal, are apparently rare, but the process is unquestionably common-
place.
As with multiple entry logging, wildfires seldom occur only once (Peres 1999, Cochrane and Schulze 1999). In
forests that are not naturally maintained by fire, the first fire opens the canopy and thereby subjects the under-
story to rapid drying. Fuel mass in the understory is augmented by trees killed by the first fire, and by grasses
and other plants that regenerate under conditions of reduced competition. The resulting fire frequencies far
exceed the historical rates. In the eastern Amazon, for example, fire frequencies in Paragominas have recently
doubled and now occur far too frequently to allow forest recovery. Human made savannas and grasslands
along with severely degraded forests due to frequent fires are becoming more the rule than the exception
throughout much of the wet and moist tropics. On the basis of either the area affected or the magnitude of
impacts, the direct damages attributable to tropical forestry are minor compared to those of wildfires.

Biodiversity Series - Impact Studies 9

BiodiversityConservation in the Context of TropicalForest Management

Figure 2 Generalized ranges of direct biodiversity impacts of timber yarding methods used for tropical
forest logging as a function of level of technological sophistication required (assumes equal volumes of timber
yarded). For mechanized ground-based yarding operations, conventional (non-RIL)and reduced-impact
logging(RIL)impacts are contrasted.

Severe

Modest

Low (;reat
Technological
Sophistication

For ground-based yarding, using the smallest
yarding equipment possible contributes to
reducing the deleterious impacts of logging,
W.Ahich yarding equipment is used is important,
but so is the care with which yarding operations
are carried out, as indicated by the contrasts
between conventional and reduced-impact
logging (RIL) on Figure 2. Unfortunately,
although the silvicultural, environmental, and
economic benefits of planning of log extraction
routes have been recognized for many decades
(see Putz and others 2000 for a review), well
planned logging operations are still the
exception in tropical forests.
Most destructive yarding in tropical forests is
carried out with bulldozers (=crawler tractors).
Bulldozers are excellent devices for constructing
roads but are unfortunately versatile enough to
yard timber as well. The excessive damage to
soils and residual trees during conventional
bulldozer yarding, especially at high intensities
on steep slopes logged during wet weather by
untrained crews is well known, but bulldozers
still yard much of the timber in commercial
logging areas in the tropics.
With the loss of forest in accessible areas in
much of the tropics, logging is increasingly
being relegated to flooded, steep, rocky, or
otherwise adverse terrain. Because ground
based yarding from such sites can be
prohibitively expensive, they have often been
left as unlogged refugia within harvested areas.
Their status as refuges is jeopardized by
helicopter yarding, because helicopters can yard
timber from even the most adverse sites.
Obtaining timber from these sites is becoming
more cost effective under some conditions.
Although areas from which timber is harvested
by helicopters are not dissected by skid trails,
haul roads are still needed, and with them, all

10 Environment Department Papers


the problems associated with increased access.
An equally important concern about helicopter
yarding is that areas traditionally avoided by
loggers are rendered accessible and are likely to
be harvested. Because helicopters leave no
obvious trails, it is going to be very challenging
to monitor their harvesting impacts.
Logging and other silvicultural treatments
Logging can be either a cause of a great deal of
avoidable damage or one of a series of
silvicultural treatments designed to promote the
regeneration and growth of commercial timber
species while protecting ecosystem services and
biodiversity. In logging areas where sustained
yield of timber is a priority, various silvicultural
treatments can be used in combination with the
appropriate logging regime to promote the
regeneration or growth of commercial species.
Carrying out silvicultural treatments in
conjunction with logging reduces their cost
while simultaneously reinforcing the idea that
logging itself can be silviculturally useful.
Silviculturally appropriate harvesting
prescriptions (from a timber stand management
perspective) run the full gamut from single tree
selection to clearcutting (Smith and others
1997). For forests with ample stocks of trees of
all sizes of the commercial species (that is,
negative exponential size-class frequency
distributions), "polycyclic" (=uneven aged)
methods such as single tree and group selection
methods are generally suitable. In such a forest,
the next crop is derived from the mixture of
trees of intermediate size (for example, 20-40
cm dbh) at the time of first cutting. In contrast,
at the time of the initial harvest the trees in the
next crop in "monocyclic" (=even-aged) systems
are seeds, seedlings, or saplings. Clearcutting is
the most obvious example of a monocyclic
system, but in "shelterwood management" a
single-aged stand develops after two stages of
harvesting. The first shelterwood harvest opens
the canopy and otherwise stimulates
Biodiversity Series -Impact Stutdies
Disaggregating"Logging"
regeneration, the second cut releases the
established seedlings and saplings. There are
innumerable variations on the themes of
monocyclic and polycyclic harvesting, and each
is appropriate under different conditions and in
forests for which there are different silvicultural
goals (Lamprecht 1989). Even within the same
forest, the silviculturally most appropriate
harvesting regime sometimes can change over
distances of less than 100 m (Pinard and others
1999).
The most common method attempted for
controlling timber harvesting in tropical forests
is to simply set a minimum stem diameter for
felling. Although theoretically easy to
implement and monitor, minimum diameter
rules are often inimical to achieving silvicultural
goals. The problem with this system is obvious
where minimum diameter limits are set below
the sizes at which trees start to reproduce
(Appanah and Manof 1991, Plumptre 1995).
Minimum diameter rules also do not prevent
harvesting of clusters of trees and thereby
creating silviculturally unsatisfactory conditions
such as where commercial species are favored
by small canopy gaps or vines proliferate in
large ones.
Logging is often the most severe of silvicultural
interventions, but there are other prescriptions
designed to increase the stocking of commercial
tree species, or to increase the growth of trees
already present. Retaining seed trees in
harvested stands is one possible way to increase
stocking, but for species that require mineral
soil seed beds or minimal competition for
germination, establishment, and subsequent
growth, seed tree retention needs to be
combined with various other treatments. Seed
beds can be modified and competition can be
reduced by controlled burning, mechanical
scarification, or herbiciding plants competing
with seedlings of the crop species. Where
natural regeneration fails, or where particularly
11
BiodiversityConservation in the Context of TropicalForest Management

high stocking levels are desired, many foresters
have tried planting seedlings of commercial
species in gaps or along lines cleared through
the forest (see Box 2). For stands in which
regeneration of the commercial species is
already established, various thinning and weed
control treatments are often prescribed to
accelerate tree growth and otherwise for "stand
improvement." Thinning around potential crop
trees, often referred to by tropical foresters as
"liberation thinning," and vine cutting are two
commonly prescribed but less commonly
applied silvicultural treatments. In
experimental areas where liberation treatments
have been applied to enhance volume
increments of commercial species, treatments
are often prescribed at 10-year or more frequent
intervals (de Graaf and others 1999). As in the
case of logging, all of these other silvicultural
treatments have impacts on biodiversity, but
they have been much less well studied.
Reducing the impacts of logging and other
silvicultural treatments
Substantial attention has been given recently to
Suced-ial loggin
has Box 3ie Most o
"reduced-impact logging" (RIL, Box 3). Most of
the practices in the various RIL guidelines that
have been promulgated of late have long been
recognized as being environmentally sound and
silviculturally appropriate (Bryant 1914). Full
application of RIL techniques would represent a
major step towards sustainable forest
management (SFM), but RIL alone does not
guarantee sustainability. Especially where tree
species being harvested only regenerate in large
clearings, the required silvicultural
interventions to assure sustained yield of the
same species may often be substantial (for
example, mimicking the impacts of slash-and-
burn agriculture or hurricanes followed by fires,
Snook (1996), Fredericksen (1998), Dickinson
and Whigham (1999), Pinard and others (1999),

Box 2
Enrichment planting
Despite scarcity of successes and millions of dollars wasted, enrichment planting in gaps or along cleared lines
continues to be invoked as a way of restoring the commercial timber production potential of severely degraded
forests. The principal problem with enrichment planting is failure to tend the out-planted seedlings (Dawkins
1961,Dawkins & Philip 1998).Admittedly there are cases where few silvicultural options remain. Unfortu-
nately, enrichment planting is often utilized in forests that should not need such an expensive intervention.
Furthermore, it is often not recognized that enrichment planting as often applied results in conversion of
natural forests into plantations. Mason and Thiollay (in press) compared the impacts of enrichment planting
with conventional selective logging in Venezuela and found the former to have greater negative impacts on
birds. With these caveats issued, a description of a large-scale enrichment planting project in Sabah, Malaysia
may be useful because the project implementers are avoiding some of the pitfalls of this technique and are
trying to mitigate some of its deleterious environmental impacts.
Funds from the FACE (Forests Absorbing Carbon Dioxide Emissions) Foundation of The Netherlands, Rakyat
Berjaya (a subsidiary of the Innoprise Foundation) have been used to plant nursery grown seedlings of com-
mercial species in several thousand hectares of forests degraded by extremely intensive and poorly controlled
logging in Sabah, Malaysia. The project has developed rapidly since its inception in 1992. Contract planters
are now trained in dendrology so that they can recognize natural regeneration, and are paid for its retention at
the same rate as for planted seedlings. Planters are also responsible for tending operations and are not paid for
seedlings that do not survive 3 years. Site matching of species has also improved, as have nursery techniques
for tending wildlings (=seedlings extracted from the forest) and raising large numbers of seedlings at rela-
tively low cost. Finally, to promote retention of wildlife in the planting area, seedlings of fleshy- fruited trees
are planted along with the mostly wind dispersed timber species in the Dipterocarpaceae. When the planted
stands are ready to harvest in 50 or so years, the concessionaire hopes to harvest 50-100 trees per hectare, an
intensity tantamount to clearcutting but perhaps preferable to the forest conversion to pulpwood plantation
option that has been adopted in much of the region.

12 Environment Department Papers

 Disaggregating "Logging"

Box3
Is reduced-impact logging (RIL) cheaper?
Many people believe that RIL has been proven cheaper than conventional logging (CL) and are therefore per-
plexed at the lack of its adoption by loggers. Adoption of RIL techniques is of particular importance in the
tropics where few countries have regulations that are both explicit enough and well enough enforced to pre-
vent exploitative harvesting.

Numerous studies over the past decades have shown that by planning skid trails and directional felling to
facilitate yarding, logging damage can be substantially reduced (Bryant 1914, Nicholson 1958, Redhead 1960,
Fox 1968, Nicholson 1979, Ewel and Conde 1980, Hendrison 1990, Johns and others 1996, Pinard and Putz 1996,
Blate 1997, Elias 1997, Winkler 1997, Uhl and others 1997, Haworth 1999). Several experimental plot-based
studies have demonstrated that due primarily to reduced yarding costs, such straightforward improvements
in logging methods also translate into direct financial benefits to the loggers (Marn and Jonkers 1981, de Graaf
1986, Malvas 1987, Jonkers 1987, Hendrison 1990, Gerwing and others 1996, Bertault and Sist 1997, Barreto and
others 1998, Holmes and others 1999, Boltz 1999). If RIL is cheaper than CL per cubic meter yarded to the
roadside, then why haven't loggers spontaneously adopted RIL techniques out of enlightened self interest? A
partial answer to this question may be that under normal operating conditions, at industrial scales, and espe-
cially on adverse terrain in wet forests, RIL may not always be cheaper (Putz and others 2000, Hammond and
others 2000). Data on the comparative costs of RIL and CL from the "Reduced-Impact Logging as a Carbon
Offset Project" in Sabah, Malaysia may serve to elucidate this issue. Tay (1999) and Healey and others (in press)
reported that the financial profits from logging were substantially lower in a 450 hectare area harvested ac-
cording to RIL guidelines than in a comparable area subjected to CL. The principal reason for lower profitabil-
ity of RIL was the reduction in yield caused by reductions in the proportion of the area logged due to restric-
tions on bulldozer access to steep slopes. Despite regulations against timber yarding from steep slopes in
Sabah, the practice is commonplace and loggers consider the timber foregone when RIL guidelines are fol-
lowed as lost profits and thus do not spontaneously adopt RIL out of enlightened self interest. These findings
indicate that the issue of cost effectiveness of RIL under adverse conditions deserves scrutiny and that loggers
not using RIL techniques may not be acting in a financially irrational manner.

In contrast to the apparent situation regarding RIL adoption in Southeast Asia, large vertically integrated
forestry companies in Brazil have begun to invest heavily in RIL apparently out of enlightened self-interest.
Although there is an additional incentive for adoption of more environmentally friendly logging in the form of
better enforcement of regulations governing forest management operations, it appears that some companies
have been convinced by the work of Baretto and others (1998), Holmes and others (1999), and others that their I
profits will increase if they adopt more efficient (and resource friendly) practices. Loggers in the region are also
applying RIL techniques in response to an increase in demand for timber certified by the Forest Stewardship
Council.

Given the differences in the results of comparative studies on the financial profitability of different logging
techniques, adoption of RIL techniques under some conditions may require either subsidies (such as carbon
offset funds) or stricter regulations with better enforcement. The conditions under which incentives for RIL
adoption are needed, and the best form and magnitude of these incentives are yet to be determined.

Fredericksen and Mostacedo (2000). This
silvicultural challenge notwithstanding, careful
planning and implementation of harvesting
guidelines would represent a big step towards
sustainable forest management.
For all yarding methods, logging damage can be
substantially reduced, and logging costs
reduced as well, by proper design, construction,
and maintenance of road networks. Much of the
cost of harvesting timber and a large proportion
of the hydrological damage (for example,
stream sedimentation) due to logging is
associated with roads (Bruijnzeel 1992). For
ground-based yarding, efficient skid trail
layouts are also essential for reducing logging

BiodiversitySeries -Impact Studies 13

BiodiversityConservation in the Context of TropicalForest Management
damage and increasing yarding efficiency, but
improper road siting often restricts skid trails to
inappropriate places. Although the required
density of roads is lower where aerial extraction
techniques are used (such as skyline and
helicopter yarding), roads are still needed and
their locations can greatly influence logging
costs and environmental damage. Guidelines
for road construction are abundantly available
and are well outlined in the FAO Model Code of
Forest Harvesting Practices (Dykstra and
Heinrich 1996). Unfortunately, forest
engineering standards in most tropical logging
areas are extremely low and there are too few
experienced forest engineers involved in most
tropical logging operations. It is important to
note that because soil damage due to poor skid
trail design or improper use, for example,
reduces productivity, increases surface erosion,
and has various other deleterious
14
environmental impacts, adhering to RIL
guidelines has advantages regardless of
whether the forest is allowed to regenerate or is
replaced by an oil palm plantation or a maize
field (Congdon and Herbohn 1993, Nussbaum
and others 1995, Pinard and others 1996).
The impacts of other silvicultural treatments on
biodiversity (such as thinning and vine-cutting)
depend on the intensity with which they are
applied and on the proper designation of areas
deemed inappropriate for stand "improve-
ment." For example, vinecutting can enhance
tree growth but undoubtedly has negative
impacts on a wide variety of animals (Putz and
others in press). Both biodiversity impacts and
labor costs of vine cutting depend on whether
only selected future crop trees are liberated or
vine cutting is carried out as a blanket
prescription.
Environment Department Papers
 Imnpactsof Forest Management
on Biodiversity
Background
Human activities are highly variable in their
influence on the components and attributes of
binfleer onytheompone ant thattribults of
biodiversity. Any human activity that results intmxliaino
substantial resource extraction or modification
will always entail significant, often unknown,
and almost always unappreciated consequences
for one or more biodiversity components,
primarily by redirecting matter and energy
flows. The cumulative redirection is enormous
at the planetary scale as three examples
illustrate: 1) Vitousek and others (1997)
calculated that 40 percent of the Earth's
terrestrial primnary productivity is being
appropriated by humans, 2) 25 to 35 percent of
the primary productivity of continental shelf
marine ecosystems is consumed by humans
(Roberts 1997), and 3) Postel and others (1996)
report that humans now appropriate 26 percent
of total evapotranspiration and use 54 percent
of all runoff in rivers, lakes, and other accessible
sources of water.
Despite these statistics on current human
impacts and the observations of the long-term
impacts of previous generations of our species
(Denevan 1992), many in the conservation and
sustainable development community still
maintain it is possible to both use and preserve
biodiversity (Huston 1993) with no costs to
either side. This claim is made regardless of a
history of human over-exploitation of resources
that began in prehistory (Goudie 1990) and is
manifested most recently in the negative effects
of tropical logging (Bawa and Seidler 1998,
Frumhoff 1995) and non-timber forest product
BiodiversitySeries - ImpactStudies
exploitation (Coomes 1995, Homma 1992). This
ahistorical and wishful thinking is extremely
angerocau a llowsits adhrentlt
dneosbcuei losisahrnst
believe that there exist easy, cost-free solutions
h lnt
t e
As societal concerns about the fates of tropical
forests increase and human demands on forests
change, so should the ways in which they are
silviculturally treated. For example, a few
decades back when the primary component of
sustainability of interest to most forest decision-
makers was sustained timber yields (STY), it
was often recommended that logging be
followed by silvicultural stand "improvement"
treatments such as poison girdling of non-
commercial trees, for a historical review see
Dawkins and Philip (1998). Unfortunately, few
of the tens of thousands of hectares of tropical
forests that were treated according to the
various silvicultural systems used in the 1950s
(for example, the Malayan Uniform System and
the Tropical Shelterwood System) remain, most
having already been logged again, converted to
oil palm plantations, or otherwise lost.
Fortunately, there is now evidence from a
lowland dipterocarp forest in Malaysia that at
least the Malayan Uniform System (MUS) could
achieve its silvicultural goals (Lee and others
1998) with mixed impacts on biodiversity,
depending on the taxon in question. The forest
treated by MUS and then studied 45 years later
was, as intended, enriched in commercial
timber trees, but also maintained much of the
fauna and flora of primary forest. More studies
of this sort are needed, and they need to be
15
BiodiversityConservation in the Context of TropicalForest Management
better circulated so that more people will realize
that at least "qualified" successes in tropical
silviculture are attainable.
In the following sections (4.2-4.6) the impacts of
logging and other silvicultural treatments on the
components and attributes of biodiversity using
the framework elaborated in section 2.0 (see
Table 1) are outlined; details are provided in the
appendices. Summarizing the impacts of
forestry activities on biodiversity in tropical
forests is an effort fraught with problems in
large part due to their immense diversity. Even
at the species level, the diversity is difficult to
imagine and each of the literally millions of
species in tropical forests responds to different
logging impacts in distinct ways. While some
general response patterns are obvious, and
others have emerged from field research, the
idiosyncrasies and apparent inconsistencies of
species responses need to be recognized. For
example, chimpanzee (Pan troglodytes)
populations have been reported to increase
(Howard 1991, Hashimoto 1995), decrease
(White 1992), and not respond to logging
(Plumptre and Reynolds 1994, see Appendix
VI). In contrast, studies of terrestrial and bark-
gleaning insectivorous birds have consistently
reported negative impacts of logging (see
Appendix IV). Many more examples of this sort
are discussed briefly below, and both cosset
aredisconsistent pattefly
below,erge. consistent
and inconsistent patterns emerge.
Landscape impacts (see Appendix I)
Logging affects the landscape component ofSaak,Mlyi)Whtelogdsnsar
biodiersit
cangig lan
by form andSarawak,
y chaging lnd fors andinterspersed
biodivrsity
ecosystem types across large geographic areas
(Box 4). Logging shifts the regional mosaic of
land uses. Although the landscape component
of biodiversity is the least sensitive to logging, influences long-term species maintenance.
changes in the size, spatial distribution, and
connectivity of habitat patches across the
landscape occur especially as the intensity of
management interventions increases. These
changes in the habitat mosaic alter species
distribution patterns, forest turnover rates, and
16
hydrologic processes. The most severe impacts
described in this section, however, result from
indirect consequences of logging such as
increased access to remote areas, fragmentation,
and altered fire regimes.
Structure: The size and spatial distribution of
tropical forest patches and the juxtaposition and
connectivity of different forest patches across
the necape are most patche across
the landscape are most affected byite indirect
impacts of logging. One of these impacts-
increased access to humans-is often
deleterious. Partcularly when logging roads
penetrate far into the forest frontier in countries
where there are numerous potential forest
colonists, logging is almost invariably
accompanied by increased hunting pressure
(Robinson and others 1999, Robinson and
Bennett 2000) and is often followed by
deforestation as lands are cleared for agriculture
(Kaimowitz and Angelsen 1998).
Another indirect impact of logging on this
attribute is fragmentation of previously
contiguous or otherwise connected forest
patches. The resulting forest fragments,
however, are typicaly not completely isolated
for all species for all time. For example, wide
lor roads may time. Forossable
logging roads may represent uncrossable
barriers for some forest interior species but
roadsides with secondary vegetation attract
many large ungulates where they are more
easily hunted (Robinson and Bennett (2000) in
Malaysia). Whether loggred stands are
within s ecies-rich forest or in a
p p
low diversity landscape dominated, for
example, by pulpwood plantations, also
Furthermore, even small unlogged patches
within harvest areas can serve as source
populations for some species after logging. The
degree of fragmentation depends on whether
logging is dispersed over large areas, or is
concentrated in small areas (Figure 3). Where
Environment Department Papers
 Impacts of Forest Management on Biodiversity

Box4
The critical role of production forests in maintaining biodiversity in the tropics
Tropical rainforests harbor approximately 50 percent of all terrestrial biodiversity and much of that biodiversity
occurs in the lowland forests which are most accessible for, and thereby most threatened by, logging and
agricultural conversion. In Peninsular Malaysia, for example more than 50 percent of all mammal species occur
below 350 m and a startling 80 percent occur below 650 m (Stevens 1968, MacKinnon and others 1996). World-
wide tropical forest loss is estimated as at least 17 million hectares/yr, an area the size of Cambodia (FAO
1993). Dramatic as it is, this global total is probably a gross underestimate given that recent figures show that
Indonesia alone has lost 18 million hectares of tropical forests between 1985 and 1997 in just three of the major
Outer Islands-Kalimantan, Sumatra and Sulawesi. Most of this forest loss has occurred in lowland forest and
logging has played a major role in that destruction. In many areas forests have been cleared and converted for
agriculture, plantations and transmigration. However, it is not logging activities per se that have caused forest
and biodiversity loss but poor concession management and new access along logging roads. These factors
have allowed shifting farmers and hunters to colonize and clear logged areas. Moreover, areas that have been
logged (whether legally or illegally) are more vulnerable to wild fires during El Nifio years; there is no doubt
that poor logging practices contributed to the great Indonesian fires in 1998 when 5 million hectares of forest in
Kalimantan burned, an ecological, economic and social disaster.

It is not just the reduction in total area of natural habitat that concerns conservationists, it is also the fact that
remaining habitat is being broken into ever smaller fragments within which species' populations are no longer
viable. As habitats become more fragmented by clearance along new roads, protected areas will increasingly
become "islands" and more and more species populations will be fated for local extinction. Fragmentation is
particularly serious because few protected areas still cover a full range of altitudinal habitats as large areas of
lowland forests, the most species-rich habitats and prime habitat for many large and wide-ranging mammals,
have been excised from designated protected areas for logging (such as Kerinci, and Gunung Leuser National
Parks in Sumatra). For Gunong Leuser National Park, for example, the main elephant, orangutan and tiger
populations occur in the lower-lying production forests outside present park boundaries. Even more alarming
is the recent increase in logging and other illegal activities within the boundaries of several National parks in
Indonesia.

The loss of bird species on Java is an indicator of what can be expected in Borneo and other large islands as
forests are cut and fragmented. Studies of forest bird distribution shows that whereas in Borneo, Sumatra, and
New Guinea, maximum avian species richness is still found in lowland forests and species numbers decrease
with altitude, Java is atypical in having a depauperate avian fauna in the hill zone between 300 and 15OOrn.
This pattern is interpreted as a result of long term deforestation since some predominantly lowland species
have failed to survive in hill forests because they have been cut off from lower altitude populations which were
a source of colonizers. Families of large birds such as malkohas have lost proportionally more species than
families of small birds such as flowerpeckers. Moreover, Java has lost more species which are exclusive to
lowland rainforests than species which can occupy secondary habitats, forest edge or open habitats. Small
forest patches have lost more species than larger blocks, with extinction rates as high as 80 percent in 10-40
hectare plots compared to rates of 25 percent for areas over 10,000 hectares. These results have important
implications for reserve design, forest management and biodiversity conservation. They emphasize thatforests
must be maintained asforests and that reserves must be large, cover a wide altitudinal range and be connected by
corridors of natural habitats. This necessitates a landscape approach to biodiversity conservation by manag-
ing production forests adjacent to core protected areas to maintain both permanent forest cover and biodiversity
so that production forests become buffer zones that effectively extend and supplement the conservation estate
(MacKinnon and others 1996, MacKinnon and Phillipps 1993).

preservation of forest interior biodiversity is the conservation biologists (Noble and Dirzo 1997).
priority, concentrating logging in small areas is Fortunately, due to associated cost savings,
generally the pattern recommended by concentration of logging activities is one of the

Biodiversity Series - Impact Studies 17

BiodiversityConservation in the Context of TropicalForest Management
Figure 3 Some impactsof loggingon biodiversityas a functionof distributionof loggingactivities(percent of
area logged)and loggingintensity(m3/ha of timber harvested)
Concentrated
Damage
.t .to Moderate
Yield
LargeAreaImpact-Free
Do
C Low Impact
J ? MinimumYield
LargeAreaImpact-Free
Low
Area Logged (percentage)
steps towards improved forest management
that loggers may find acceptable.
The impacts of logging and other timber stand
management activities on landscape structure
are minimized when plantation products are
used in place of those from natural forest.
Opting for this so-called "New Zealand
Solution" (Hunter 1998) is not currently an
option likely to be exercised in many tropical
countries. Nevertheless, as forests dwindle and
economies grow (such as in Thailand, Malaysia,
Chile, and South Africa), this solution will
become more viable.
Composition: Logging activities may directly
and indirectly affect the identity, distribution,
and proportion of habitat types in tropical
forests. Forestry may directly affect composition
of the landscape component of biodiversity by
intentional creation of new types of habitats
(such as forests converted into plantations).
Furthermore, if silvicultural objectives are
uniform across the landscape, inter-stand
diversity is sacrificed by widespread
application of the same stand "improvement"
treatments. Perhaps most importantly,
18
WorstCasefor Biodiversity
Maximum
Yield
SmallAreaImpact-Free
Dispersed
Damage
ModerateYield
SmallAreaImpact-Free
High
improved access provided by logging roads
indirectly fosters post-logging habitat changes
by human forest colonizers, weeds, and
wildfires.
Setting aside reserves within logging areas may
mitigate some of the deleterious impacts of
logging and other silvicultural treatments. The
specific location of reserves substantially
influences their value in biodiversity
conservation. Optimally, the full range of
landscape features and habitats should be
represented within protected areas.
Function: Logging may markedly alter several
landscape level ecological processes subsumed
under the functional attribute of the landscape
component of biodiversity. For example,
logging roads, and activities associated with
their construction, can greatly influence the
permanence of the forest fragments they create
by altering landscape level disturbance regimes.
In large part, disturbance is altered because
roads and skid trails provide ready access to the
forest for both colonists and fire (see below).
High intensity and widespread logging,
especially if not carefully controlled, also
Environment Department Papers

influences hydrological processes at all levels
perhaps including the regional climate. Where
logging roads are wide and logging intensities
are high, landscape level movements of animals
can be disrupted (Goosem 1997), as can gene
flow in plants when pollinators are restricted to
isolated fragments by inhospitable
surroundings. It should be noted, however, that
the impacts of roads depends on where and
how they are constructed, and change with time
as the road and its margins mature (Lugo and
Gucinski 2000).
In this section the emphasis is on the impact of
fires on the functional attributes of landscapes
in full recognition of the importance of fires to
all components and attributes of tropical forest
biodiversity. Although fires have long played
substantial roles in the ecology of tropical
forests throughout the world (Goldammer 1990,
Saldarriaga and West 1986), wildfires as
exemplified by the recent Indonesian fires have
recently increased in frequency, extent, and
intensity in part due to widespread logging
(Cochrane and Schulze 1999, Cochrane and
others 1999). Canopy opening due to timber
harvesting operations increases the rate of forest
drying and thereby increases inflammability
even in characteristically wet forests and
swamps (Holdsworth and Uhl 1997). Perhaps
equally importantly, vegetation proliferation
along logging roads and skid trails greatly
enhances fire penetration into forest interiors
even where logging damage is slight or non-
existent. Whether or not the extensive forest
fires that have recently become common can be
indirectly attributed to logging, it is clear that
fire damage often exceeds logging damage in
many tropical countries. In Bolivia in 1999, for
example, fires burned an estimated 20,000 km2
whereas logging was carried out in perhaps
only one-tenth of that area (W. Cordero, pers.
comm.).
Although difficult to detect on satellite images,
but see Souza and Barreto (in press) for recent
Biodiversity Series- ImpactStudies
Impacts of ForestManagement on Biodiversity
improvements in relevant remote-sensing
methods, the impacts of the understory burns
typical for tropical forest fires are substantial.
Especially in forests that have not burned for
many decades to centuries, the effects of even a
single understory burn are great, even if they
are not fully realized for many years. Typically
small trees are killed whereas large trees often
receive only minor basal damage. Unfortu-
nately, basal damage exposes trees to wood
rotting organisms; a high incidence of heart rot
is typical of forests that burn at low intensities
at infrequent intervals (F. E. Putz, pers. obs.).
Elevated rates of tree mortality rates for many
years post-fire are also characteristic in seldom-
burned forests. Such long-term data are difficult
to obtain because accumulation of fuel in the
form of fallen branches as well as post-fire
proliferation of palms, grasses, and other fire-
carrying plants render forests very fire prone.
Surprisingly, the effects of tropical forest fires on
biodiversity have not been extensively studied
(Leighton and Wirawan, 1986).
Ecosystem impacts (see Appendix II)
The deleterious ecosystem-level impacts of
logging on tropical forests are widespread,
substantial, enduring, and well studied
(especially compared to the landscape and
genetic components). The ecosystem component
of biodiversity is somewhat more sensitive to
logging impacts than the landscape component
in part because management activities are
usually implemented at this scale. In contrast to
the landscape component, most ecosystem-level
impacts are a direct consequence of logging
activities. Logging purposely removes biomass
from ecosystems, but it also alters their vertical
complexity and soil characteristics. Depending
on the silvicultural objectives, changes in
structural heterogeneity may be intended.
Whether intended or not, the structural impacts
of logging alter the relative proportions of
lifeforms and biogeochemical stocks, as well as
nutrient and hydrologic cycling, productivity
and energy flows.
19
BiodiversityConservation in the Context of TropicalForestManagement
Structure: Logging may affect the structural
attribute of the ecosystem component of
biodiversity by changing the biophysical
properties of soils, spatial heterogeneity of
forest stands, and biomass. The extent and types
of ecosystem damage to these structural
attributes depend on logging intensity, yarding
system, and the care with which the operations
are conducted. Soil compaction, for example, is
a major problem during ground-based yarding
operations especially where skid trails are
unplanned and yarding continues during wet
weather. Where compaction is severe, soil
permeability and bulk density often require
many decades to recover. Exposure of mineral
soil after litter layers and root mats are bladed
off by bulldozers is also a concern. Mineral soil
disruption during bridge building, road
construction and maintenance, and skidding
operations also represent forest management
activities that affect ecosystem structure and
thereby have biodiversity impacts.
Logging induced soil compaction negatively
affects hydrology, which in turn alters the
characteristics of watercourses. For example,
water infiltration rates into soil, surface flow
rates and volumes, and stream channel
characteristics are all adversely affected by
uncontrolled logging. Deposition of large
volumes of unconsolidated sediments into
streams during road construction and due to
increased erosion from exposed mineral soils on
log extraction routes can also greatly modify
stream characteristics such as pool-riffle-run
ratios. The impacts of such changes on aquatic
organisms have been little studied in the tropics.
Another impact of logging on ecosystem-level
structure is reduction in biomass and alteration
of necromass. Losses of biomass due to forest
management activities include the amounts in
the removed timber, damage to trees in the
residual stand that result in mortality, and any
silvicultural treatments that result in tree death.
Biomass losses range from 5-10 Mg/ha at the
20
lowest logging intensities, to substantially
greater amounts where heavy logging is
followed by poison girdling of non-commercial
trees in the residual stand. Necromass,
including coarse woody debris, increases
immediately after logging but may then
decrease to levels below pre-logging conditions
due to increased temperatures near the ground
and associated increases in decomposition rates.
Wildfires promoted by logging road
construction and canopy opening, as well as
controlled bums carried out for silvicultural
purposes, can have obvious effects on biomass
and necromass stocks in tropical forests.
Maintenance of healthy communities, species
populations, and gene pools is predicated upon
protection of hydrological functions, nutrient
cycles, and other ecosystem properties.
Fortunately, methods for mitigating the
ecosystem-level impacts of logging on tropical
forests are well known. Switching from ground-
based to skyline yarding techniques, for
example, greatly reduces damage to residual
stands, soils, and streams but allows harvest on
steep slopes. The opposite trend in tech-
nological change also can have environmental
benefits; log yarding with draft animals or by
manual means generally results in substantially
less logging damage than yarding with
bulldozers (Cordero 1995). To enjoy these
potential benefits, the expertise of experienced
forest engineers should be more often called
upon where logging does have to occur.
Composition: One important way that logging
affects the ecosystem-level compositional
attribute of biodiversity is by changing
biogeochemical stocks. For example, soil
compaction reduces water holding capacity,
which in turn leads to increased surface runoff.
Limited storage capacity in natural streams is
further reduced by sedimentation, which means
that flow regimes can be greatly modified by
logging, especially during the first years after
logging is completed. Various RIL techniques,
Environment Department Papers

such as installation of cross drains on skid trails,
can greatly diminish these impacts.
Where large volumes of timber are harvested
and post-logging forest recovery is retarded by
soil damage, fire, or weed infestations, standing
stocks of nutrients in biomass are greatly
reduced. Storage of nutrients released from
biomass in necromass and soil organic matter is
generally brief in lowland tropical forests due to
accelerated rates of decomposition and low
cation-exchange capacities in the soil. The
results can be substantial leaching and, if the
forest is burned, volatilization of nitrogen and
sulfur.
Function: Logging affects the functional
attribute of ecosystem-level biodiversity by
adversely affecting hydrological and
biogeochemical fluxes as well as productivity.
Reduced plant productivity results in part from
impeded root growth, a consequence of logging-
induced soil compaction. Because most of the
available nutrients are usually found near the
top of the soil profile, blading of the soil surface
also diminishes nutrient availability in local
areas and otherwise interferes with nutrient
cycling. In more extensive portions of logging
areas where RIL guidelines are not followed,
nutrient cycling and hydrological functions are
greatly modified by reduced canopy
interception of rain and mist, decreased uptake
of water and nutrients by the diminished
biomass, and increased occurrence of surface
erosion and landslides especially associated
with improperly located and poorly constructed
roads and skid trails.
Changes in carbon storage and flux associated
directly and indirectly with logging and other
silvicultural activities influence whether forests
are net sources or sinks of "greenhouse" gases.
For example, substantial logging-induced
transfers of living trees to coarse woody debris
can have substantial effects on understory
structure and dynamics leading to more carbon
Biodiversity Series - Impact Studies
Impacts of Forest Management on Biodiversity
release. The deleterious impacts of logging on
forest carbon balance can be greatly diminished
by application of RIL techniques (Pinard and
Putz 1996). Substantial biodiversity benefits are
also likely to result from RIL, but this and other
co-benefits have not been well studied. Other
silvicultural treatments such as fire
management, weed control, thinning, and
enrichment planting have various impacts on
both greenhouse gas emissions and biodiversity
that should be investigated.
Logging, especially if followed by silvicultural
treatments such as liberation of future crop trees
from competition, can substantially change the
physiognomy, composition, and trophic
structure of forest stands. To a large extent,
these modifications represent the goal of forest
"refinement" treatments applied to increase
volume increments and relative densities of
commercial timber species. This "stand
domestication" by nature reduces species
richness; rare, threatened and endangered
species may become locally extinct especially if
they have no perceived commercial value.
These changes in composition and structure
affect numerous community-level ecological
processes including colonization, predation and
mortality rates, pollination, seed dispersal, and
timing and abundance of flower and fruit
production.
Structure: The most obvious logging-induced
impact on the structural attribute of
community-level biodiversity is the change in
proportions of successional stages in forest
stands. Depending on harvesting intensities,
planning of roads and skid trails, and training
and supervision of workers, logging can result
in large changes in the proportion of forest in
mature, recovering, and early successional
stages. In some severely disturbed areas,
succession might be "arrested" by post-logging
proliferation of vines, bamboo, and other non-
arboreal growth forms. Silvicultural treatments
such as thinning and vine-cutting can increase
21
BiodiversityConservation in the Context of TropicalForest Management
the rate of succession and increase the
proportion of stand growth concentrated in
commercial species, but not without affecting
biodiversity in more than the intended ways.
Composition: For the community component of
biodiversity, logging affects composition by
changing (often purposefully) the relative
abundance of species and guilds inhabiting
forest stands. Relative abundances of tree
species with light demanding vs. shade tolerant
regeneration, wind vs. animal dispersed seeds,
vertebrate vs. invertebrate pollinated flowers,
and thick vs. thin bark, for example, are all
subject to change in logged and otherwise
silviculturally treated forests. Likewise,
sieviculturatiyntreatedfforest. uikewofanise, s
rersntto odifrnguilsonml
(such as understory insectivores and arboreal
folivores) is influenced by forestry activities,.
Depending on a great number of factors related
to the intensities, spatial scales, and modes of
forest intervention as well as characteristics of
the focal taxa, effects of forestry activities can be
negative, positive, or neutral. For example, in
eight studies that considered the impacts of
logging on frugivorous birds, two reported
positive impacts at the guild level, three
reported negative impacts, and three reported
no change at all (Appendix IV).
Function: The functional attribute of
community-level biodiversity includes
numerous key ecological processes (such as
pollination, herbivory, seed dispersal and
predation) all of which are affected by logging
especially under the most intensive
management interventions. Many of the effects
on these processes are a direct consequence of
altered resource abundance (for example fruit
for frugivores or young leaves for folivores),
which in turn result from the logging-induced
changes in community structure and
composition. In addition to being influenced by
resource-base changes, these ecological
processes are also affected by changes in forest
22
microclimates that result from exploitation,
silvicultural treatment, and by hunting (see Box 5).
Species impacts (see Appendices V and VI)
The species component of biodiversity has
received the most attention from researchers
concerned about the impacts of logging and
other silvicultural treatments in tropical forests.
The most obvious species-level impact of
logging is on the abundance and age/size
distribution of harvested and damaged trees.
Depending on the intensity of logging and the
care with which it is carried out, the
reproduction, growth, and survival of a great
number of species can be adversely affected. In
reviewing this literature, it is important to note
that the taxa studied were not selected at
random. Instead, in many cases, the species
chosenswer d, to becsensitie to,cand
thus goo i da of fensitr impacs
Structure: The most immediate and direct
impacts of logging on the structural attribute of
the species component of biodiversity are
suffered by the harvested tree species. Their
populations are often left greatly depleted,
especially in the larger size classes of
reproductive individuals when management is
based solely on minimum diameter felling
rules. Because of the spatial clustering
characteristic of many commercial timber trees,
the richest patches of forest are generally the
most severely disturbed, unless logging
guidelines specify minimum spacing between
harvested trees.
Changes in forest structure are suffered most by
specialist species of the forest interior. After
logging, many formerly shaded microenviron-
ments in the forest interior become drier,
brighter, warmer, and more easily exploited by
some predators. For example, severe canopy
opening adversely affects litter invertebrates
and their predators. For species that are
generalists in their diets and wide-ranging in
their habitat use, such as many frugivorous
canopy birds, the direct impacts of logging vary
Environment Department Papers
 Impacts of ForestManagement on Biodiversity

Box 5
Hunting in logged tropical forests
The escalating scale of wildlife harvest in logged areas is an insidious problem that may undermine the
biodiversity conservation potential of forests managed for wood production (Robinson and others 1999,Fimbel
and others in press). Logging operations multiply the harvest of wildlife from tropical forests primarily by
increasing access to previously remote areas. These same access roads serve as conduits for commercially
traded meat and other wildlife products. Although the magnitude of the impacts of subsistence hunting of
tropical forest animals (for food, feathers, and skins for example) by indigenous people are non-trivial (Robinson
and Redford 1991, Redford 1992,Robinson and Bodmer 1999,Peres 2000),the impacts of commercial hunting
are several orders of magnitude greater. Of particular concern is the tendency for hunters to target almost any
species larger than 1 kg, including many species that are vulnerable to extinction due to their long life spans
and low rates of population recovery (Bodmer and others 1997).
Hunting pressures increases in logging areas partially because of the number of people involved in logging
operations as well as by the increased access provided by logging roads. In formerly isolated forest areas,
loggers themselves hunt (Ruiz and others in press) or provide ready markets for meat and other wildlife
products (Wilkie and others 1992, Bennett and Gumal in press). Local communities quickly shift toward com-
mercial hunting to supply markets to which logging roads provide access. This shift initiates a positive feed-
back cycle wherein money gained from wildlife products buys better weapons with which to hunt, which in
tum increases the harvest. Such escalations are unsustainable, however, and some species become locally ex-
tinct.
Loss of wildlife threatens the sustainability of tropical to the extent that the animals targeted by hunters play
key roles in ecological processes including seed dispersal, seed predation, and herbivory (Redford 1992,Jansen
and Zuidema in press). Even when animal species are not extirpated, their numbers may be reduced suffi-
ciently such that they are rendered ecologically extinct. The loss of wildlife may have cascading effects on the
structure and composition of the tropical tree community with deleterious consequences for recruitment of
commercial timber species. Such dire predictions are most likely to be borne out in areas such as the Guyanas
where a large proportion of canopy trees are dispersed by large animals (Hammond and others 1996).Finally,
as human populations expand and forest landscapes become more fragmented, the potential for wildlife to re-
colonize defaunated areas will likely diminish.
While the deleterious consequences for wildlife of logging and other silvicultural treatments (such as vine
cutting) deserve further investigation and mitigation, it should be recognized that the indirect impacts of log-
ging resulting from increased access often far outweigh the initial damage done by even the worst predatory
logging.

from being somewhat negative, to neutral, to
positive. For the understory species that are
adversely affected by logging, the effects may
persist for decades (Wong 1985, but see Lee and
others 1998). It is worth reiterating that the
imnpacts of hunting on populations of large and
slow-reproducing animals generally overwhelm
any potential benefits that they might have
enjoyed after the canopy was opened by logging
(Bennet and Dahaban 1995, see Box 5).
Population sizes and structures of most species
are also drastically modified by the fires that so
often accompany uncontrolled logging.
Species impacts of logging and stand refinement
treatments are of particular concern in small
forest management units. Private landowners
with less than 100 hectares to manage, for
example, may be unwilling to set aside 10
percent of their forest for species preservation. If
their forests are surrounded by similarly
managed or deforested areas, then blanket
apation o refiement treatmet
application of stand refinement treatments or
heavy logging can take a heavy toll on
commercial and non-commercial species alike.
Composition: Logging affects the composition
attribute of species-level biodiversity by

Biodiversity Series - Impact Studies 23

BiodiversityConservation in the Context of TropicalForest Management

changing the abundance and distribution of
species. Unless logging is accompanied by other
silvicultural treatments designed to foster their
reproduction and growth, the abundance and
population structure of the harvested tree
species are greatly modified by logging (see Box
6). Logging impacts on tree populations
continue for many years after logging is
completed because damaged trees suffer high
mortality rates, proliferation of weeds (such as
vines) interferes with tree reproduction and
survival, and population size reduction and
fragmentation can decrease pollination levels and
change patterns and intensities of seed dispersal
and predation. Species composition of animals also
changes in response to the direct impacts of logging
(such as canopy opening) and the associated
indirect impacts as well (for example increased fire
frequency and intensity, hunting, and forest
conversion).
Changes in species composition in response to
forestry operations are by no means consistent
across or even within taxa. Our review of the
literature on primates, for example, revealed
few cases of consistent responses of species to
logging (Appendix VI). This variation can be
attributed to differences in logging intensities as
well as to differences in the duration of post-
logging population monitoring. Silvicultural
treatments other than logging, especially vine
cutting and crown liberation of future crop
trees, might have more consistent deleterious
impacts on canopy animals, but such impacts
have been little studied.
Small fragments of untouched forest that
remain within even heavily logged forests serve
as important refugia for plants and animals.
Wildlife densities in these unlogged fragments
can be very high during and shortly after
harvesting, but then diminish as animals
recolonize the surrounding matrix. Many
1munplanned" reserves are on steep or otherwise
adverse sites, which certainly influences their
function as refugia. Implementation of

Box 6
Lesser-known species-Marketing challenges, silvicultural
impediments, or desirable diversity?
Sustaining volumetric yields of timber is made challenging in tropical forests where many trees have little
market value (Plumptre 1996). Particularly where just a few species of light-demanding trees have ready
markets, species that are commercially lesser known or that have less desirable timber properties often render
regeneration-enhancing treatments prohibitively expensive (Fredericksen 1998, Pinard and others 1999). Where
markets for formerly lesser-known species have developed and they can therefore be harvested profitably,
forest managers can financially justify creating the large canopy openings required for regeneration of the
most valuable light-demanding timber species.

The same lesser known species that interfere with timber stand management treatments and present chal-
lenges for wood technologists and marketing firms represent an important component of forest diversity. One
indication of their importance is that whereas tree species with wind dispersed seeds (such as Dipterocarpaceae
and Meliaceae) dominate the international timber trade (ITTO 1996), a high proportion of species producing
less well known timbers produce fleshy fruit and animal dispersed seeds (Jansen and Zuidema in press).

Silvicultural treatments applied to promote regeneration and growth of commercially desirable species might
be made more feasible by increased market demand for what are currently lesser known timbers. While pro-
moting sustained yield of the species being harvested, these silvicultural treatments intentionally result in at
least locally substantial modification of pre-intervention stand structure and composition. Compromises be-
tween sustained yield and the more all-encompassing goal of sustainable forest management need to be in-
formed by research on the direct and indirect impacts of increased timber harvesting (Fredericksen and others
1999).

24 Environment Department Papers


helicopter yarding, and to a lesser extent other
aerial yarding techniques, may place many of
these unlogged patches in jeopardy.
Function: Demographic processes (such as
survivorship, fertility, and recruitment) and
growth rates are two key functional attributes of
the species component of biodiversity that
logging affects. Populations of many organisms
are susceptible to large fluctuations after
logging due to both the direct impacts on forest
conditions (for example microclimate and
fragmentation)
fragmentingfioandand the indirect
forest impacts
condirectsimTe s of
hunting, fire, and forest conversion. The
proliferation of disturbance-adapted taxa in
logged-over forests, some species of which are
not native or were not previously common in
the area, can have large but as yet little studied
imnpactson the resident flora and fauna.
Genetic impacts (see Appendix VII)
The genetic component of biodiversity is likely
to be the most sensitive of all components to
logging because of reductions in effective
population size and interruptions in gene flow.
At present, however, little is known about the
genetic structure of any tropical organisms,
even commercially valuable timber trees (Ledig
1992). Furthermore, the techniques required for
assessing the genetic structure of populations
are sophisticated and expensive. Except in a few
cases, concerns about dysgenic selection, genetic
drift, and other genetic problems are based on
controversial theory that is rapidly developing
as evidence accumulates.
Structure: Logging affects the structural
attribute of the genetic component of
biodiversity by reducing effective population
sizes and heterozygosity. Effective population
sizes of both commercial and non-commercial
species are reduced by harvesting, other
silvicultural treatments, forest fragmentation,
weed proliferation, and wildfires. There are also
good reasons to be concerned about the effects
of logging and stand improvement treatments
on dioecious species and in small forest
Biodiversity Series - Impact Studies
Impactsof ForestManagementon Biodiversity
management units in which population sizes of
all species are correspondingly small. Allelic
frequencies of commercial species change after
removal of a large proportion of healthy
reproductive adults. For species with high
densities of advanced regeneration, genetic
structure of their populations are unlikely to
change dramatically after selective harvesting,
unless collateral damage is severe. Timber stand
improvement treatments also may affect the
genetic structure of species targeted for removal
(such as woody vines) as wetr as their
(uha
associates, od ie)aimpacts
but these elahavehiapparently
not been studied. Given the high proportion of
vines and other plants that resprout after
cutting (=coppice), large impacts on genetic
cure are lkely.
structure are unlikely
Composition: The fact that most species are rare
in tropical forests implies that allelic diversity
will decrease with increasingly intensive
management interventions. Unregulated
harvesting of all merchantable individuals of
commercial species, for example, has immediate
impacts on allelic frequencies that continue to
change due to decreased effective population
sizes. Deleterious recessive genes may become
more apparent due to dysgenic selection and
heterozygosity may decline due to the
"bottyeneck" effect in the small, isolated
potions that inut smarvested
populations that result from harvesting, forest
fragmentation, and other direct and indirect
impacts of forestry activities (Styles and Khosla
1976,Murawski and others 1994aand b, but see
Newton and others 1996).
Function:Logging may affect the functional
attribute of the genetic component of
biodiversity by interrupting gene flow, which in
turn influences outbreeding rates. Decreased
effective population sizes coupled with losses of
pollinators and seed dispersal agents can result
in reduced gene flow and inbreeding depression
in populations of both commercial and non-
commercial species. Especially vulnerable are
populations represented by scattered mature
25
BiodiversityConservation in the Context of TropicalForest Management

individuals and very few juveniles (for example obligate outcrossers, only very severe
many "long-lived pioneers" such as the reductions in effective population size are likely
mahoganies). Given the high proportion of to have much effect on gene flow (Ghazoul and
tropical tree species that are dioecious or others 1998).

26 Environment Department Papers

 Overview of Biodiversity Con-
servation in Relation to Logging
5 and other Silvicultural Treatments

Synthesis The impact of these silvicultural approaches on

A way of graphically displaying the impacts of
logging on tropical for-ests (Figure 4) based
upon the various components and attributes of
biodiversity as described above was developed.
Along the vertical axis of our framework the
five components of biodiversity were arrayed in
the order of increasing susceptibility to logging
impacts (landscape, ecosystem, community,
population/species, and genetic). The
horizontal axis arrays a variety of approaches to
silviculture in order of increasing intensity.
each biodiversity component using three
categories were assessed. First, each
biodiversity component was scored as 'mostly
conserved" for cases in which their attributes
are expected to usually stay within their natural
range of variation. Second, biodiversity
components were scored as "affected" for cases
in which their attributes are expected to
frequently fall outside their natural range of
variation. Finally, biodiversity components were
scored as "mostly lost" for cases in which their

Figure 4 Expectedeffects of a rangeof forest useson the componentsof biodiversity

___- - .Legend:
NTFP= Non-timber forest prod-
. ~~~~~~~~~~~~~~~ucts
Genetic _
fA
c
> MQSTLYRIL
~~~~~MOSTILY
= Reduced-impact
logging
3} .1% Reserves = Protected areas
5 Species -oa LOST within logged units
0
Refinement = Silvicultural
E treatments such as liberation of
6 Community AFFECTED
future crop trees from competi-
tion, which can substantially
change the physiognomy, com-
._% .position, and trophic structure
L \ . of forest stands which are ap-
> \csse plied to increase volume incre-
.2 Ecosystem \ ' and relative densities of
ments
0
n0
MOSTLY \ commercialtimberspecies
m Landscape
CONSERVED Enrichment planting = Increas-
ing the stocking of commercial
speciesby planting seedlings (or
seeds) in logging gaps or along
A ~ x x~ Yx clearedlines
t~ -900 CL = Conventional logging

BiodiversitySeries - ImpactStudies 27
BiodiversityConservation in the Context of TropicalForest Management

attributes are expected to almost always fall
outside their natural range of variation.
The scoring process used for Figure 4 was
admittedly subjective; the scores were based on
a reading of the literature and the authors'
experience. It is fully recognized that particular
situations might warrant different scores, and it
is emphasized that the purpose is to illustrate
what is believed to be a useful analytical
process rather than to obtain perfectly accurate
scores. It is also emphasized that this
framework serves to suggest hypotheses that
seem to be important research topics for
conservation biologists from a number of
different disciplines.
The responses summarized on Figure 4 actually
represent a multitude of impacts from a
diversity of silvicultural approaches applied
with varying degrees of concern for biodiversity
over a large range of scales. Additionally, Figure
4 addresses neither the issue of profitability of
different forest uses nor issues related to land-
use capability, biodiversity value, or the
capabilities and desires of local stakeholders. At
least two other dimensions of this multi-
dimensional topic are captured in Figure 5, on
which it is indicated that every forest activity
can be carried out over a range of intensities.
Correspondingly, each forest-use activity
generates timber volumes and financial profits
that also range widely (Figure 6). Recognizing
that forest use practices vary over time with, for
example, market fluctuations and political
change, and that biodiversity impacts are a
function of a multitude of interacting factors
operating at different temporal and spatial
scales, it is hoped that Figures 4-6 present an
accurate "snapshot" of the relationship between
biodiversity conservation and forest
management.
The impacts summarized in Figure 4 and the
ranks and ranges of impacts and profits on
Figure 5 are based on a combination of literature
reviews and the authors' subjective estimations.
For example, under the range of stocking levels,

Figure5 Generalized biodiversity

impactsplottedagainstexpectedshort-termfinancialreturnsto forest
ownersor concessionaires (NTFPs= non-timberforest products)

Severe

(A et nv/e tona
v Y 1g oggin (CL)/
'hi~~~~~~~~~~~~~~~i
E
.t Reced
u /

.0

Modest P areas
Small Great
Financial Profitability

28 EnvironmentDepartmentPapers
 Overview of BiodiversityConservation in Relation to Loggingand other SilviculturalTreatments

Figure 6 Generalized biodiversityimpacts resultingfrom different approaches to forest management for

timber. (CL= conventionallogging,RIL= reduced-impact logging,
refinement= anyof a varietyof silvicultural
treatmentsappliedto increaseratesof timbervolumeincrement; and,reserves= protectedareaswithinloggedunits)

Severe

E mn

4-.

sia 1
Modest
0 1 2 3 4 5 6
CommercialTimber Production(m3 /ha/yr)

terrain, and accessibility that logging is carried
out, reduced-impact and conventional logging
overlap in both impacts and profitability.
Nevertheless, particularly where logging is
carried out on steep slopes or under otherwise
adverse conditions, application of most RIL
guidelines results in immediate profits that are
lower than those enjoyed by unconstrained
conventional loggers. This observation helps
explain why where regulations are non-existent
or unenforced, conventional logging will likely
be used to log such areas. Correspondingly,
some of the impacts of high intensity RIL
overlap those of low intensity conventional
logging, but the former generally has fewer
adverse environmental impacts than the latter.
1) Tropical forests are often assumed to have
developed their current structure,
composition, and functional properties
under a disturbance regime characterized
by small, localized, and natural
perturbations. The importance of
unrecorded widespread cataclysmic
anthropogenic and natural disturbances,
especially those of past centuries, is seldom
recognized.
2) It is often assumed that regenerating trees
harvested for timber is simply a matter of
protecting advanced regeneration (for
example, seedlings, saplings, and poles) and
providing small canopy openings in which

Troubling assumptions they can mature.

Three widely held and interlocking assumptions 3) Many people believe that devolution of
about tropical forests complicate the issue of the authority over forests to indigenous groups
compatibility of forest management for timber and other rural communities will enhance
and biodiversity conservation: efforts towards sustainable management

Biodiversity Series - Impact Stuidies 29

BiodiversityConservation in the Context of TropicalForest Management
and biodiversity conservation (Luckert
1999, Putz 2000).
While these assumptions hold true in some
tropical forests, many of the trees currently
being harvested actually regenerated under
conditions very different than those of today or
those that will be created by harvesting alone.
The most familiar example is provided by the
neotropical and African mahoganies (Meliaceae
spp); these trees and many others like them are
the heritage of slash-and-burn agriculture,
hurricanes, river meanders and/or catastrophic
fires of centuries past. The problem confronting
silviculturalists managing for such light-
demanding species is that promoting
regeneration requires intensive stand
manipulations which are expensive and have
great impacts on the biodiversity present
immediately prior to intervention (Lugo 1999).
Despite accumulating evidence of substantial
and long-term pre-historical and more recent
impacts of humans on what were formerly
considered "natural" ecosystems, it is important
to recognize that the current rates, spatial scales,
and intensities of human-induced perturbations
often far exceed the resilience of tropical forests.
Forest recovery after disturbances such as
mechanical clearing for pastures, for example, is
known to be extremely slow, especially if the
pastures are large or intensively used. Natural
regeneration in such areas can be accelerated
through application of silvicultural treatments,
but full recovery of diversity, from the genetic to
the landscape level, will require centuries, if it
occurs at all. For example, although leaf
biomass may recover quickly, in abandoned
agricultural clearings coarse woody debris
accumulates at rates measured in centuries.
Research progress on methods for afforestation
and reforestation should not beguile us into
believing that, once destroyed, tropical forests
can be recreated.
The cause of social equity may be well served
by devolution of management responsibility to
local communities, but the biodiversity benefits
30
are less secure. Some communities, such as the
municipalities in Bolivia that have recently
received control over substantial forest areas,
may be indifferent about conservation
(Kaimowitz and others 1998). In other cases,
such as some communities in Nepal, devolution
of forest control has had substantial
conservation benefits. Obviously the process of
devolution deserves scrutiny lest some turn out
to be worse forest stewards than the logging
companies and national governmental agencies
they replace. Empirical evidence suggests that
community behavior towards forests depends
on the forest richness or forest poorness of their
particular situation, but many other factors are
likely involved and deserve investigation.
Conclusions
All consumptive use affects some component or
attribute of biodiversity, commonly affecting not
only the target resource but other factors as well
(Redford and Richter 1999). The population/
species component is most commonly
understood to be affected by silvicultural
activities although effects, some of which are
subtle or cumulative, are undoubtedly often
missed even in this comparatively well studied
component. Of increasing importance is an
understanding of how the community and
ecosystem components have been (Runnels
1995) and are being affected by logging and
other silvicultural activities (Noss and
Cooperrider 1994, Vitousek and others 1997,
Fimbel and others, in press).
Recognizing that all significant interventions in
natural forests have biodiversity impacts, all
silvicultural decisions necessarily represent
compromises. Management for some goods or
services necessarily involves management
against some others. What are "weeds" to
timber stand managers are food sources, rare
species, carbon stores, or inter-crown pathways
for other human and non-human stakeholders.
The biodiversity compromises involved in
deciding whether to cut vines, retain seed trees,
or enhance seedling establishment by carrying
out controlled burns should be informed by
Environment Department Papers
 Overview of BiodiversityConservation in Relation to Logging and other Silvicultural Treatments
research. Unfortunately, very little is known
about how tropical forests can be managed in
the most biodiversity friendly manner.
Researchers have instead focussed on
enumerating the deleterious environmental
impacts of uncontrolled logging by untrained
and unsupervised crews. To inform decisions
about tropical forest management and to assure
that biodiversity is protected to the maximum
extent, more research is needed on how to
maintain diversity in forests selected for
logging. The large and rapidly growing body of
literature on ecosystem management in both
north and south temperate forests (such as
Kohm and Franklin 1997, Lindenmayer 1999)
should provide inspiration and starting points
for tropical researchers intent on solving
biodiversity related problems associated with
forest management.
The primary conclusions to derive from these
analyses are that:
D
Different intensities and spatial patterns of
timber harvesting, along with other
silvicultural treatments, result in different
effects on the different components of
biodiversity.
a Some components and attributes of
biodiversity are more sensitive than others
to forest management activities.
* Only extremely limited use will protect all
components (that is, large protected areas
are essential for biodiversity conservation).
The capacity to mitigate the deleterious
environmental impacts of logging and other
silvicultural treatments should not be construed
as constituting unilateral support for
sustainable forest management as a
conservation strategy. Such an endorsement is
unwarranted given widespread illegal logging
in the tropics, widespread frontier logging and
logging of areas of high priority for biodiversity
protection, the persistence of poor logging
practices despite substantial efforts in research
Biodiversity Series - Impact Studies
and training, and the general slow rate at which
most loggers are transforming themselves from
timber exploiters into forest managers.
Nevertheless, even the most harshly treated
forests maintain more biodiversity than tree
farms for pulpwood, oil palm plantations,
maize fields, or cattle pastures. Furthermore,
logging is often the environmentally least
damaging of land uses that are also financially
viable (Pearce and others 1999). Given these
conclusions, effective mechanisms for financing
forest protection and environmentally sound
forest management are needed. It might also
prove useful to evaluate critically the limits to
the assumption that well managed forests are
less likely to be converted to other land uses
than forests that are logged without apparent
concern for sustainability.
By focusing on the deleterious environmental
impacts of tropical forest management
activities, we often lose sight of the fact that
from a biodiversity maintenance perspective,
natural forest management (that is, maintaining
forests as forests) is preferable to virtually all
land-use practices other than complete
protection. The number of taxa with reportedly
inconsistent, neutral, or positive responses to
logging is an indication that timber harvesting
is not necessarily incompatible with protection
of many components and attributes of
biodiversity. This conclusion derives even
greater support from the fact that nearly all of
the studies conducted to date on the
biodiversity impacts of logging were carried out
after unplanned logging by crews with no
training in reduced-impact logging techniques
and no incentives to reduce their impacts. As
forest management practices improve under
market pressure or pressure from land-owners,
the deleterious environmental impacts of
logging and other silvicultural activities are
likely to be substantially reduced. Forests that
are carefully managed for timber will not
replace protected areas as storehouses of
biodiversity, but they can be an integral
31
BiodiversityConservation in the Context of TropicalForest Management

component of a conservation strategy that
encompasses a larger portion of the landscape
than is likely to be set aside for strict protection.
In other words, forests managed primarily for
timber, if managed properly, will supplement
and effectively extend the conservation estate.
Finally, it should be recognized that landscape
management is consistent with the ecosystem
approach emphasized by the Convention on
Biological Diversity (CBD).

32 Environment Department Papers

6 Recommendations
1. Given that all substantial forest
interventions affect some component or
attribute of biodiversity, the best way to
assure biodiversity conservation is through
the establishment and protection of large,
properly located, and well managed
reserves and protected areas.
2. In some parts of the world and some types
of forest, there should be no logging at all.
3. Existing legislation and regulations should
be enforced. Where they do not exist, or are
weak they should be created or updated to
reflect current conservation and sustainable
resource management concerns.
4. Within forests used for logging and other
silvicultural activities, biodiversity
conservation is enhanced by setting aside a
portion of the area for complete protection.
Optimally, these reserves should be large,
shaped so as to minimize edge effects, cover
representative areas of all the ecosystem
types present, and include features of
special concern for biodiversity
maintenance such as water courses, rock
outcrops, and salt licks.
5. Where logging is to be carried out, reduced-
impact logging guidelines (Dykstra and
Heinrich 1996) should be fully implemented
by well trained crews.
6. Researchers need to determine the financial
and environmental costs and benefits of the
BiodiversitySeries - ImpactStudies
components of reduced-impact logging and
various other pre- and post-logging
silvicultural activities. Particular attention
should be paid to plant and animal species
with attributes that render them susceptible
to logging-induced extirpation (Martini and
others 1994, Pinard and others 1999).
Regional guidelines for certification need to
be developed and regularly updated to
reflect increasing knowledge about
biodiversity so as to assure that certification
efforts realize their conservation potential.
7. Measures to protect biodiversity should be
included in all forest management plans.
Particular attention should be paid to the
retention of nest and den trees as well as
species of great importance to biodiversity
conservation.
8. Wildfires need to be controlled. Successful
fir e contro lled. Sude
fire control programs will need to include
public awareness building as well as
implementation of existing technologies and
methods. Research is needed for developing
cost-effective fire control measures with
minimal biodiversity impacts.
9. While continuing to promote and develop
community-based wildlife management
prograrns, hunting by loggers and market
hunting of slow reproducing species should
be prohibited. Enforcing existing laws will
in many cases confer substantial protection
to species threatened by over-hunting.
33
BiodiversityConservation in the Context of TropicalForest Management
10. Training is needed for researchers who will
develop biodiversity-sensitive silvicultural
methods as well as cost-effective methods
for monitoring the environmental impacts
of silviculture. Training is also needed for
the field crews that are responsible for
implementing the recommended practices.
Perhaps the most critical shortage is of
forestry/conservation "research
practitioners" who understand both the
broad scientific and the more specific
technical aspects of tropical forest
management.
34
11. The biodiversity impacts of devolution,
plantation conversion, certification,
12. forest-based carbon offsets, and global
trends in timber markets need to be
monitored.
13. Linkages to the policy arena must be
pursued. Ecosystems provide important
services which typically are not internalized
into the decision making process.
Legislation and their regulations are an
integral part of this process and should be
reviewed, updated and enforced.
Environment Department Papers
Appendix I -
Impacts of Logging (and other
silviculture treatments where noted)
on the LandscapeComponent
of Biodiversity in Tropical Forests

Biodiversity Series - Impact Studies 35

 Impacts of logging (and other silviculture treatments where noted) on the landscape component of biodiversity in tropical forests

LandscaPe attributes
Category Structure Composition Function
Vegetation,
Soil& - Fragmentation
/ roads+ greateraccess
to > 50%forestarealogged+ specieslost > Increased humancolonizationandfire &
Water humans4 increased
deforestation
(Kaimowitz
& (Eastern
Amazon:Nepstadet al. 1992) decreasedmigrationof animalsandgeneflow
Argelsen1998) in plants(if pollinatorsare restrictedto
fragments)
> Tree populations,biomass,
andcommunity > Ecologicaldegradation
of 1°forestfragments > Morethan 50yrs requiredfor recoveryof pre-
compositionaffectedmoreby edgeeffects from increasededgeeffects& newhabitat loggingstructure(Uganda: Plumptre1996)
than by patchsize(Amazonia:
Gascon& matrix(Amazonia:Gascon& Lovejoy1998) 200yrs necessary for recoveryof spp
Lovejoy1998) composi-tion followingloggingdamage
(Malaysia, forestgapmodel:Kurpicket al.
1997)
Increasedspatialheterogeneityof habitatsafter > Increaseinmixedforest(w/ valuabletimber o Impoundments dueto undersizedor failed
140yrs of agroforestry,
logging,
& charcoal spp)& decrease of Cynometraforestpatches culverts& bridgeabutmentskill treesand may
productionfollowedby 50yrs regeneration afterarboricidetreatmentfrom 1951-90 influencefauna(F.E.Putz,pers.obs.)
(PuertoRico:Garcia-Montiel & Scatena1994) (Uganda: Plumptre1996) )> Increased likelihoodof largescaleaccidental
> Maintenance
& recoveryof species firesdue to canopyopeningand roads
composition
in loggedpatchesdependson (Amazonia:Cochrane& Schulze1999)or fuel
landscape
context(Liu& Ashton1999) accumulation fromfire protection(Miombo
woodlands,Africa:Chidumayo1988)
;O Complexmosaicof foresttypes& disturbance > Harvestgapswarmerthan naturaltreefallgaps
w/ 85% swampforest& only 15%lowland (Amazonia:
Vitt et al. 1998)
forestescaping
loggingdisturbance
(W.
Kalimantan,
8 yrs post-logging:
Cannonet al.
1994)
> Patches(I - 100ha)of unlogged
forestsin
heavilyloggedarea(500-1000ha,80-120
m3 /ha) on steepslopes(>350) and in rocky
areas(Sabah,
Malaysia:
Pinard& Putz1996)
Primates
Non-rodent
mrommals
Rodents
& > Increased
smallmammalrichness in forest
Marsupials fragmentsb/cof increased
edgeandnew
habitat matrix Gascon
& Lovejoy1998)

Birds
Herps > Neotropical frogs- see rodents above(Gascon
& Lovejoy 1998)
Insects > Lossof species& genesof butterfly
communitiesb/c of conversion of 30% of
landscapeto agricultureandforestry
(Neotropics: Brown 1997)
Soilorganisms
Appendix II
impacts of Logging (and other
silviculture treatments where noted)
on the Ecosystem Component
of Biodiversity in Tropical Forests

Biodiversity Series - Impact Studies 39

 Impacts of logging (and other silviculture treatments where noted) on the ecosystem component of biodiversity in tropical forests

Ecosystem Attributes
Category Structure Composition Function
Vegetation,
Soil& P Reduced biomass,basalarea,andstemdensity > Thin barkedspeciesmostsusceptible to P Increased
vulnerabilityto fire: ignitionafter2
Water (Uhl et al. 1991,Silvaet al. 1995,Pinard& Putz logging(andfire) (Amazonia:Uhl & Kaufman weeksw/out rainin gapsvs.monthsin
1996,Webb1998,Finegan & Camacho1999) 1990;Pinard& Huffman1997) understory(Amazonia: Cochrane& Schulze
> Reductionin largetreeswith hollows& holes 1999,Nepstadet al. 1999)
importantfor wildlife(Australia:
Gibbons&
Lindenmayer 1996)
> 2/3 basalarea,3/4 numberundamaged trees, > Higherrichness (trees> 20cm dbh)in logged P Decreased evapotranspiration & infiltration;
I/5 no of emergents(>30 m), & 1/3 vs.unlogged,butwhenequalnumbersof trees increasedsurfaceflow 4 nutrientlosses
commercial volumeof unloggedforest sampledrichnesswasequal(W. Kalimantan, 8 (Brtower1996, Poels1987,Bruijnzeel 1992), but
(Venezuela, 19yrs post-harvest
of 10trees/ha: yrs post-logging:
Cannonet al. 1998) nutrientbssesdon'trestrictproductivity(Proctor
Kammesheidt 1998) > T related to 1992)
> Forestsurfacefireskill40-95%of alltreesw/ geography thanwhethera patchwaslogged; > Soilbulkdensity6 yrspost-logging60%greater
dbh > 10cm(Amazon:Nepstadpersobs) richnesshigherin westof reserveandin logged thaninundisturbed forest(Malaysia:
Malmer&
patches(L.tanda,60yrspost'sustainable
logging': Grip1990); skidtrailsestimated
to require> 50
P 36% biomass lossfrom IncreasedtreePkp-e16)yst co'rhdaiccduty(aly:
motlt (Aaoi,
mortality(Amazonia,IAA fro frgmn
I00 m from fragment Plumptre1996) ~&Knrzna yrsto recoverhydrauliccorductvity
96(MaVsia:
edge,10-17yrspost-fragmentation: Laurance K& K 1996)
etal. 1997)
> Increaseinorganicfuels(56to 180tons/ha)on > Reduceddensitiesof timberspp& shiftin > 100-170M tonsof carbon/yremittedfrom
forestfloor (easternAmazon,post-logging:
Uhl dominance& agestructureof canopyspp 1981-1996 (Asia:Houghton& Hackler1999)
&Kaufman1990,Uhl& Vieira 1989) (PuertoRico,post 140yrs agroforestry, > 0.5%of totalglobalcarbonemissions
(Brazilian
> Soildisturbance mainfactorlimitingseedling logging,
charcoalproduction& St yrs Amazon,1996-97:Nepstadet al. 1999)
regeneration& establishment (Kalimantan: regeneration:
Garcia-Montiel& Scatena1994)
Gardingen et al. 1998)
> 5-40%of forestfloor disturbedby roads,skid > Epiphytic& stranglerFicussppsufferhighrates > PostCL, streamsuspended solidsandturbidity
trails& tractortracks(Cannonet al. 1994 of damage (Malaysia,
post-logging:
Lambert were 12x& 9x > controland levelspersisted
Johnset al. 1996,Pinard& Putz 1996,Sistet al. 1991) at least5 yrs; RILlevelswere 2x > controlbut
1998a,Holmeset al. 1999) recoveredafter2 yrs (Malaysia: Zulkifli&
Anhar 1994)
Vegetation,
Soil& > Soilproperties(i.e.BD, total porosity,sathydr > Woodyvinesincreased w/in 100m of edges > Watershedsedimentloadswere 18x higher5
Water conductivity,
& resistance to penetration) butdidn't compensate for lost biomassfrom mospost& 3.6xhigherI yr post-logging
Vegetation required12-52yrs (skidtrailslongest)to increased tree mortalityAmazonia forest (Malaysia:Douglaset al. 1992)
recover(Malaysia: Kamaruzaman & fragments yrspcstfragmentation:
10-17 Laurance> 200 yrs necessaryfor recoveryof sppcomp
Kamaruzaman 1996) et al. 1997) followingloggingdamages (Malaysia,
forestgap
> Limitingharvestgapsto < 650 m2 (< 2 model:Kurpicket al. 1997)
trees/gap)enhanceddipterocarpregen& > Runoffas% precipitationrangedfrom58% -
inhibitedpioneerregen(Kalimantan:Gardingen 94% in plantationsconvertedfrom forest
et al. 1998) depending on method(Malaysia: Malmer1992)
> Harvestgapsmuchlargerthannaturaltreefall > Road& skidtrail impoundments mayincrease
gaps& thushavelesslitter andshade& more methaneemissions (F.E.Putz,pers.obs.)
intenselight(Amazonia:Vitt et al. 1998)
> Sedimentloadshavenegativeeffectson
> Tree basalareaandcommercial volumes dwsra qai ie(utai:Cmbl
recoveredwithin40yrs postloggingw/ MUS downstreamaquaticlife (Australia:Campbell&
(Malaysia:
Manokaran 1998) Doeg1989)
Primates > Longdistanceseeddispersal andspecies
movements to/from differenthabitatpatches
likelyimpeded(nospecificdata)
Non-rodent > Species movements to/from differenthabitat
mammals patcheslikelyimpeded(no specificdata)
Rodents
& > Speciesmovements to/from differenthabitat
Marsupials patcheslikelyimpeded;seedpredationrates
likelyaltered(nospecificdata)
Bats > Speciesuseof differenthabitatpatches&
pollination& seeddispersalrateslikelyaltered
(nospecificdata)
Birds > Birdsppdeclinew/ increasing
disturbance > Speciesuseof differenthabitatpatcheslikely
(Lawtonet al. 1998) altered;dependingon spp.,pollination,
predation,seeddispersal rateslikelyaltered;
(nospecificdata)
Herps
 CommunityAttributes
Category Structure Composition Function

Insects > Arthropod diversity decreaseswhen structural > Various ecosystemprocesseswill be relatively
complexity decreases(Holloway et al. 1992; unaffected(Vitousek 1990)
Gardner et al. 1995)and/or b/c of changesin
understory,litter, & soil microclimate (Blau
1980,Kremen 1992,Spitzer et al. 1997,
Camilo & Zou in press)
Soilorganisms ) 25%-75% reductionin soil mycorrhizae > Termite richnesslower in clear-cuts, but
(Malaysia,post-logging:Alexanderet al. 1992) higherin regeneratingplots than I' forest
(Cameroon),but overall richnesswas similar in
1°, 3 yr and 17yr secondarysites (Malaysia:
Eggletonet al. 1995);changesin termite comp
maybe delayedafter disturbance(Eggletonet
al. 1997)
> Nematodesdeclined alongdisturbance
gradient & were 40% lower in clearcut vs. 1'
forest (Bloemerset al. 1997)
> Soilfungi increasedfrom 17 spp in 7-yr
regrowth to 27 in 16-yr regrowth; soil
microbial popin'spositivelycorrelated w/ tree
density & basalarea(India:Arunachalamet al.
1997)
> Ectomycorrhizaldiversity greater under open
& regeneratingcanopiesthan in undisturbed
forest (Kalimantan,9 months post clear-cut or
partial logging,trees > 50 cm dbh: Inglebyet
al. 1998)
> Termite populationsincreaseafter
conventionalloggingmore than after RIL
(Sabah,Malaysia,80-120 m3/ha harvested:F.E.
Putz, pers.obs.)
(MUS = MalayanUniform System,a monocyclic
approachto timberstandmanagement)
Appendix III
Impacts of Logging (and other
silviculture treatments where noted)
on the Community Component of
Biodiversity in Tropical Forests

Biodiversity Series - Impact Studies 43

 Impactsof logging(and other silviculturetreatments where noted) on the community componentof biodiversityin tropical forests
Community Attributes
Category Structure Composition Function
Vegetation > Increasedproportion
of open > Increased
speciesrichness(esp.light-demanding pioneertrees,shrubs i Decreasedseeddispersalof 6 timber
canopy(Uhl&Vieira1989, andlianas)> 10yrs post-logging (Brazil:Verissimoet al. 1995, speciesin logged+ huntedforestvs.
Thiollay1992,
Cannon et al. Magnusson et al. 1999;Salicket al. 1995,Silvaet al. 1995,Delgadoet al. logged+ protectedforest(CostaRica:
1994,Mason1996); upto 1997,CostaRica:Webb 1998) Guariguata et al. in press)
50%ingapsvs.5-10%
D Aggressive
shade-intolerant sppdominateandcompetitivelyexclude > Fruit productionof treeslower IOyrs
(Hartshor 1978,Denslow otherplants(Uganda: Struhsaker1997) post-logging (Amazonia: Johns1992)

> Greaterdensityof > Multipletree fellinggapsmostdiverse,butdominatedby pioneertree > Highdegreeof outcrossing and low density
understoryvegetation5-6 spp(CostaRica:Webb 1998) of matureindividuals(neotropics:
yrs postharvest(Venezuela: Murawski1995,Stacyet al. 1996)
Mason1996)
Remaining
largetrees o Highdominanceof few sppin heavilydisturbedhabitats(e.g.,roads) > Seedproductionlower inseveraltimber
inadequate
to sustainfuture (CostaRica:Guariguata
& Dupuy1997) species(Uganda:Plumptre1995)
harvests(Philippines,
I 5 yrs Large-scale,long-termshift insppcomposition
afterlogging,augmented - Increasedself-pollination
attributedto low
post-SLS:Tombec& spp(Uganda:
by arboricidetreatmentof non-commercial Plumptre densityof floweringindividuals(Murawski
Mendoza1998) 1996) & Hamrick 1991,1992)4 lessseed

> No changein overallrichness & compositioninloggedvs.logged+

thinned(Nicaragua;Salicket al. 1995)
Reduced tree density& > Increasedirradiance4 moreearlysuccessional spp(Swaine1996, > Wind-dispersedseedsfallin higher
canopyclosure,denser Mulkeyet al. 1996,Whitmore1998)especially vines& pioneers(Lowe numbersingapsthan in understory
cover < 2 m aboveground, & Walker 1977,Cannonet al. 1994) (Augspurger& Franson1988,Loiselleet al.
& equalgroundcover Post-logging is differentthano flli (Peters 1996)4 highercolonizationlevelsfrom
(Indonesia,5 yrs post- ostg spp.
seedfor wind-dispersed
selective:Hill et al. 1995)
> Increased dominanceof fastgrowingtimberproducingtree spp.after
silviculturalrefinement(Malaysia,
MUS:Chua1998)
Reducedleafbiomass but > Vinespp.richnessanddensitiesrecoverto pre-MUSloggingand - Reduceddispersal
of seedscarriedby
increased
leafquality; treatmentby 40 yrs (Malaysia:
Gardette1998) large-bodied,
gap-shyfrugivores(Gorchov
reduced log distribution, Lich d ir d mo d d 40 d et al. 1993,Forget & Sabatier 1997)or that
regenerating
stems,and siMcultural
treatirent(Maoas div
MUS:Wolseley
et al01998) arescatterhoarded
by caviomorphrodents
overall trsieeutualteamnt(Mlysa,MSsWize e l.I(Forget & Milleron 199I; Forget 1990,
(Madagascar,RIL,<10% > Palmspp.richnesslowerinloggedandMUStreatedforestthanin
1°forest 1993,1994;Asquithetal. 1999)esp.if
forest:Ganzhorn1995, (Malaysia
NurSupardi
et al. 1998) huntingpresent
Ganzhornet al. 1990)
 Vegetation > Fewerlargediametervines40 > Mortality rates 4x higher for spp w/ dbh >
yrspostMUSthanin II forest 40 cm (S. India, 10-I5 yrs post- selective
Gardette1998)
(Malaysia: logging:Pelissieret al. 1998)

> for
Denningtreeslessavailable > Tree mortality increasedafter
arborealrodents& marsupols liberation/refinement treatments (Costa
afterselectiveloggng Rica:Finegan& Camacho 1999)
Laurance
(Auistralia: & > Vine cutting promotes tree growth &
(autra .A
Laurance1996;Invchoa &
Sorianoin ) reduces loggingdamagebut removes
important intercrown pathwaysfor wildlife

> Repeated selectivelogging (reviewed in Putz 1991)

reduces abundance of large
treesandshiftscomposition
to earliersuccessional species
(Australia:Home& Hickey
1991
Primates > Increasedinfantmortality > Omnivore densities(1-25 trees harvested/ha,I-SO yrs post-logging, > Herbivory & frugivory - enhancedby low
for variousspp. Africa, Asia,Amazon)- 8 spp decreased,13spp increased,5 spp intensity (< 10% affected) logging
immediatelyafter logging, showed no change;severalspp showed opposite results in different disturbance (Madagascar:Ganzhorn 1995);
but some spp recovered 6- studiesand/or different sites (reviewed in Plumptre & GrieserJohnsin data lackingfor higher intensities, but
12yrs post logging press) presumed asvariable as changesin spp
(Malaysia:Johns& Johns > Folivore/frugivoredensities (asabove)- 6 spp decreased,5 spp densities

1995) increased,3 spp showed no change;as above,severalspp showed > Seeddispersal& predation - no specific
opposite responsesin different studies/sites(Plumptre & GrieserJohns data; probably varieswith animaldensities
in press);huntingaffects folivores more than loggingitself (Africa: Oates > Vine cutting and liberation thinning
eta]. 1996) increasedlocomotory costs and

> Frugivore/folivoredensities (asabove)- 3 spp decreased,2 increased,4 susceptibilityto predators for non-volant
spp showed no change;again,severalspp responded differently in arboreal animals(reviewed in Putz et al.
different sites/studies(Uganda:Plumptre & Grieser Johnsin press) 2000)

> Lemursin dry forest (< 10% affectedby logging)- increasedsightingsof > Predation of smallprimates by raptors may
all spp attributed to increasedfruit & higherfoliage quality on remaining increasein responseto vine removal
trees (Madagascar:Ganzhorn 1995) (Suriname:S. Boinski, pers comm.)

> Monkeysin Zaire - 4 spp preferred secondaryforest: 3 others preferred

primary forest (Zaire: Thomas 1991)
> Folivores- hunting hasgreater negativeimpact than logging(Africa:
Oates et al. 1996)

vp~ > Primatedensitiesdecreasedin responseto logging& MUS (Malaysia:

CA1 Laidlaw 1998)
 CommunityAttributes
Category Structure Composition Function
Non-rodent > Browsers/grazers (e.g.,Asianelephant,Africanbuffalo,gaur,banteng,& > If no hunting,increasedherbivory;if
mcmnmds sambardeer)increase(if no hunting)b/c of increasedfood quality huntingdecreased herbivory
(Oateset al. 1990,Waterman& Kool 1994)& quantityJohns1992);the
onlysensitivesppin thisguildwasthegiantforesthog(Hylochoerus
meinertzhageni)(reviewedin Davieset al. inpress)
> Carnivores- mixedresponses & few data;sppwith restricteddiets
mostsensitive(reviewedin Davieset al. inpress)
> Speciesfavoredby hunters(e.g.,duikers,pigs,peccaries),for special
trade(e.g.,elephants)
androadsidefeeders(e.g.,Synceroscaffer,Bos
spp)decline(Davieset al. inpress)
> Large,slow(moving& reproducing) speciesdecline(huntinga
confoundingfactor)(Lahm1994,Grieserjohns 1997)
> Speciesdependenton fruits/seeds of timberspecies(e.g.Susbarbatus,
Caldecott1991)onclosedcanopy(e.g.,Muntiocus atherodes, Heydon
1994;Giaoet al. 1998),or with limiteddietaryflexibilitydueto
energeticrequirements (e.g.,Tragulus spp.Heydon& Bulloh1997)
decline.
Rodents
and > Arborealmarsupials
- I spdeclined & 4 sppunaffected
after8-10trees(50-55> Increasedseedpredationrates(espif
Marsupials m3) harvested/ha
(Australia:
Laurance & Laurance1996) hunting)(e.g.,Terborgh& Wright 1994)
> Smallmammalpopulations mostsensitive- according to models:(a) > Decreased predationon largeseedsif large
Sciuridae,
Echimyidae & othercanopydependentfrugivores/granivores; seedpredatorsextirpated(Putzet al.
and(b) sppw/ low RAandrestrictedgeogrange(Ochoaet al. 1993, 1990)
1995;Utrera 1996;Ochoa1997;reviewinOchoa& Sorianoin press) ' Increasein Muridaemayincrease
colonizationof smallseededtree spp.
Smallmammal populationsleastsensitive
- according
to models:Didephidae & (Hammond& Thomasin press)
Muridae;
semi-arborealomnivores/predators increase
(e.g.,Didelphis
spp.&
Philander
opossum; seeaboverefs)
> Rodentspprichnessincreased soonafterlogging(Uganda, 9 trees/ha,
62% canopyopen:lsabirye-Basuta & Kasenene 1987,Muganga1989),
butdecreased by 16yrs post(Lwanga1994)
> Rodentspp.richnessincreased, 2 dominantspp.decreased inabundance
butincreasedin bodymassafterlogging(China:Wu et al. 1996)
Bats > Pioneerplantdispersers (e.g.,Carollia.Artibeus& Sturnira)increase > Increased dispersalof pioneerplantspp;
(Charles-Dominique 1986,Fleming1988,Fentonet al. 1992,Kikkawa& decreasedpredation
Dwyer 1992,Brossetet al. 1996,Utrera1996,reviewin Soriano&
Ochoain press)
> Phyllostominae (insectivores & vert predators)- decline(e.g.,Artibeus > Reductions could
in insectivorepopulations
obscurus, Vampyressa bidens, Phyllostomus elongatus,Tonatia resultin increased herbivoryrateswith
sauropila)or disappear(e.g.,Vampyrumspectrum,Tonatiasylvicola, deleteriouseffectson forestresource
Chrotopterusauritus,& Peropterixkoppleri,Emballonuridae) (Fenton productivity(Soriano& Ochoain press)
et al. 1992,Brossetet al. 1996,Utrera1996,reviewin Soriano& Ochoa
in press)
> Reduced guildcomplexity & kcalextinctionsafterlogging(5.8m3/ha,trees>
40cmdbh);effectsexacerbated byenrichment strips(FrenchGuiana:Brosset
inVenezuela:
similarfindirngs
et al. 1996; Ochoain press; reviewed inSoriano
& Ochoainpress)
> Open-forest, fast-flyingsppw/ long,narrowwingsdo bestb/cof
increased gaparea;dominated6-yr postsitew/ 6-7 trees/haharvested
(Australia:Crome& Richards1988)
> Spppreferringmediumto denseunderstorydisappeared afterbrushing
indicatinghighsensitivity to changes in foreststructure(Millerin press)
Birds > Nectarivore& generalist - nc or increasein most )
frugivore/insectivores Herbivory& frugivory- reducedfor fig-
spp(ohns 1989a,1992b;Lambert1991;Thiollay1992;Zakaria1994, eatingsppb/c of lower post-harvest
1996;Mason1996;reviewsin Mason& Thiollayin press,Zakaria& densityof Ficusspp(Lambert1991)
Francisin press,& Plumptrein press)
> Terrestrial& sallyinginsectivores & otherspp
(e.g.,Tyrannidae) > Predation- expectedto decreaseb/c of
depending on forestinterior (e.g.,thoseinmixed-sppflocks)decrease reducedinsectpreyabundance (b/c of
w/ logging(ohns 1989a,1991a, 1992a;Lambertet al. 1992;Thiollay hotter/drierconditions)(Mason1995)
1992;Zakaria1994;Bennet& Dahaban1995;Fanshawe 1995;Grieser
Johns1996;Mason1996;Owiunji & Plumptre1998;reviews inMason&
Thiollayin press,Zakaria& Francisin press,& Plumptrein press)
Birds > Predators, foliagegleaners responses
& largefrugivores- inconsistent in > Seeddispersal- reducedb/c of avoidance
aohns1991
differentsites/studies a; Lambertet al. 1992;Thiollay1992; of largeopeningsin forestcanopy(Stouffer
Grieserjohns1996;Zakaria& Nordin 1998;reviewsin Mason& & Bierregaard1995a,b)
Thiollayin press,Zakaria& Francisin press,& Plumptrein press)
> Understoryspp.- nectarivorous increased; decreased; > Colonizationrates-expectedto increase
insectivorous
latter trendworsenedby enrichment planting(Venezuela:Mason1996) whenrefugesprovidedw/in loggedareas
-(ohns1996)
 _____________________________ CommunityAttributes
Category Structure Composition Function

> Interiorunderstorysppdeclinedby 37-98%;generalists & spp

associated w/ densesecondgrowth,edges& largegapsincreased;
overallrichnessandabundance down27-34%(FrenchGuiana,1-17yrs
post,38%undergrowth& 63% canopyloss:Thiollay1997);(other
study:3 trees/ha,10yrs post)- lowerdiversityb/c of moreuniform
habitat(Thiollay1992)

> Flycatchers,
wren-warblers,
trogons& woodpeckersdeclined;
nectarivores
& somefrugivoresincreased
(Malaysia,
8 yrs post-logging:
Lambertet al. 1992)

9 Spp.richnesslowerthanunlogged(Indonesia: Marsden1998)
> 1/3sppexpectedto disappear(Liberia,loggedfragment:Kofron&
Chapman1995)

Herps > Heliothermiclizards- increaseddensityof largelizards& alteredcomm > in

Competition& Predation- increases
structure(Amazonian treefallgaps:Vitt et al. 1998) generalistspp(Vittet al. 1998)
> No changein reptilecommunitycompor abundance
(< 10%damage,2- > Colonization- generalistspp(e.g,Hyla
Bloxamet al. 1996)
12yrs post-logging: increase(Caldwell1989)
geographica)
> Loweramphibian diversity(Amazonia,
highintensityloggingor logging&
& roads:Pearman1997,Vitt et al. 1998)
fragmentation
> Sppthat livein leaflitter mostseverelyaffectedespeciallyafterhigh
intensitylogging(e.g.,monocyclicharvests); w/ lessintenseloggingmost
spppersistin shortterm,but invasion of 2° forestby largepredatory
lizardsmayextirpatesmallfrogs& lizards;somefrogsppincreaseafter
loggingb/c of pondinginskidtrail rutsor in streamimpoundments
_._____________ (reviewedinVitt & Caldwellin press)
Insects > Communitycompof Araneae,Hymenoptera, Heteroptera,Homoptera,> Shortgenerationtimes& highfecundity
differedin 1°forest,logged
Coleoptera,Orthoptera& Lepidoptera allowrapidrecoveryof effected
forest& plantations Nummelin1996)
(Uganda: populations(temperatezone studies:
Huhta 1976,Heliovaara& Vaisanen1984;
Spp.restrictedto understory,leaflitter & soilmostvulnerable(reviewed tropics:Ghazoul& Hill in press)
in Ghazoul& Hill in press)
Geometridmothdiversitylowestin abandoned clearcut,highestin > Negativecascading
effectson insect-
forestloggedw/ MUS(Malaysia: Intachatet al. 1997;Ghazoul& Hill in nutrientcycling)are likely
decomposition,
Press) (Ghazoul& Hill in press)
> Lowerlepidopteran mothdiversityafterlogging+ conversionto
dung& carrionbeetleslessaffected(Malaysia:
plantation;Coleopteran
Hollowayet al. 1992;Ghazoul& Hill in press)
> Dungbeetlespp.compositionin RILarearesembles10forestmorethan
Daviesinpress)
loggedarea(Malaysia:
conventionally
> Butterflies,flyingbeetles,canopybeetles,canopyants,leafeaterants,
termitesandsoilnematodes all declinedalongdisturbancegradient
(Lawtonet al. 1998)
> Butterflyrichness, abundance & evenness (Indonesia,
alldecreased 5 yrs
post-logging:Hill et al. 1995)
> Cassidinaebeetlerichness vs. 1°
greaterin loggedforestandplantations
forestandloggedforestcommunitycomppositionmorelikeplantations
(Uganda:Nummelin& Borowiec1991)
> afterinvasionof pioneerplantsat
Opportunisticspeciesincrease
expenseof closedforestspp(Hollowayet al. 1992)
> similarin Ioforestand MUSforest40 yrs after
Ant spp.richness
treatment(Malaysia:
Bolton1998)
 CommunityAttributes
Category Structure Composition Function

Soilorganisms > Fruitingbody(sporome) ) Mycorrhizalfungispp.richness higherin MUStreatedareathanin 1°

densityof mycorrhizal
fungi forest(Malaysia: Watlinget al. 1998);wood-rottingorganisms maybe
higher40 yrs.After MUS better impactindicatorsfor logging& othersilviculturaltreatments
treatmentthanin I' forest > Changes to soilpropertiesdirectlyinfluences
soilfaunadiversity(review
(Malaysia:
Watlinget al. in Camilo& Zou in press)
1998)
> Endogeic & anecicearthwormcommunities littleaffectedby smallscale
clearcuts& low frequency(>50 yr intervals)largescale(reviewedin
Camilo& Zou in press).

Aquatic
systems > Lossof instream& riparianzonediversity;decreased abundanceof > Alterationsin light& temperatureregimes,
fishes& benthicinvertebratesintolerantof (a)highsedimentation,(b) streamsedimentloads,turbidity,stream
shiftsin photosynthesis/respiration
ratios,(c) invasive
spp(reviewin chemistry,hydrology& fluvial
Pringle& Benstead in press) geomorphologymaycausecascading
effectson aquaticfoodwebs(Pringle&
Benstead in press)
Appendix IV-
Bird Responses (densitybased on
inter-site comparisons) to Logging
in Tropical Forests, Organized by
Feeding Guild

Biodiversity Series - Impact Studies 51

01
Bird responses(density' basedon inter-site comparisons)to loggingin tropical forests,organized by feeding guild

Ulu Ulu
Budongo, Kibale, Tekam, Segama, Segama, Imatoca, PistedeSt -Elie,
Site Uganda Uganda Malaysia Sabah Sabah Venezuela FrenchGuiana
Logging
intensity 20-80m3 /ha 2! m3lha 18trees/ha 79m3 lha 90m3 lha 5-8 m3lha 1Om3lha 10m3/ha
Recovery(yearssinceharvest) 25-50 25 6-7 3-4 9-10 1-6 1 10
Hunting no no unknown no no no yes yes

Guild
Frugivore + nc + - nc nc - -
Frugivore/insectivore
(arboreal) nc + + + + + nc nc
Frugivore/insectivore(terrestrial) nc + - - nc nc na na
Frugivore/granivore nc nc na na na na na na
Granivore nc + na na na + na na
Granivore/insectivore nc + na na na na na na
Insectivore(sallying) - - - nc nc
Insectivore(terrestrial) nc nc
Insectivore(arboreal foliagegleaner) nc nc - nc na
Insectivore(understory foliagegleaner) nc + nc - nc
Insectivore (bark gleaning) nc - - nc
Insectivore/nectarivore nc nc nc + + +
Predator nc + nc + - na
Predator/frugivore na na - nc nc na

Source Owiunjiand Dranzoa Johns1989a Nordinand Lambert Mason1996 Thiollay Thiollay

Plumptre 1995 Zakaria et al. 1992 1992,1997 1992,1997
1998 1997
Notes:
Density(calculated
by samemethodwithina study,but differedbetweenstudies).+ = higherfollowinglogging,- = lower followinglogging,nc = no change,
PB 0.05. (Adaptedfrom MasonandThiollay,in press;Plumptreet al.,in press;andZakariaand Francis,in press).
na= informationnot available;
Appendix V
Impacts of Logging (and other
silviculture treatments where noted)
on the Species Component of
Biodiversity in Tropical Forests

Biodiversity Series - Impact Studies 53

 Impacts of logging (and other silviculture treatments where noted) of the speciescomponent of biodiversity in tropical forests
Taxa Structure
Vegetation > Smallersizeclasses mostaffected>
by felling,skidding,androad
construction
> 62% seedling& 59%sapling
mortality(Malaysia: Borhanet al.
1987)
S I1I%of treesuprooted(Brazil:
Uhl & Guimaraes1989)
17treesdamaged for everytree
cut (E.Kalimantan:
Abdulhadiet
al. 1981)
0 >50% of youngtreeskilled(N.
Queensland, Australia:Cromeet
al. 1992)
)0- - 60-70%of residualtrees
damagedby logging(Fox1968;
Nicholson1979)
> Largeandsmalltreeskilledwith
equalprobabilityJohns1988;
Cromeet al. 1992)
> Speciesw/ type IIl population
structuremaybeeasily
eliminated(C. Peterspers.
comm.)
> Spatiallyclumpedpattern of
conspecifics shiftsto dispersed
or
randompatterns4 reproductive
biologyandlogisticsof future
silviculture
Primates > >
Species
Attributes
Composition
Highgradinglargecommercial trees
causedlocalextirpationof Caesalpinia
echincata,
Ceibapentandra, andAniba
duckei;Amazonia(Gentry& Vasquez
1988,GrieserJohns1997)
> Harvesting
figspromotestheir
regeneration& maynot havewildlife
impactsdue to scarcity of well-formed
trees(Bolivia;Fredericksen et al. 1999) >
> Populationsof wild fruit treesare often )
decimatedby destructiveharvesting (Peru:
Vazquez& Gentry1989)
SeeAppendixVI
Function
> Reduced seedavailability
& reducedseeddispersalby rodentsof
largeseededcommercialspeciesafterlogging(Guyana: Forget&
Hammondin press,Hammond& Thomasin press)
> Fordioeciousspecies,seedcrop decreases with increasing
distanceof malesto females(Mack1997)
) Higherseedlingmortality(esp.invery disturbedareas)in Sabah
(Pinardet al. 1996)
(iade l 96
Lowerseedling growth rates(Nussbaum et al. 1995)
Reduced densityof adults(i.e. breedingindividuals)4 fewer
flowersper unitarea(C. Peters,pers.comm.)
> 1% increasein tree mortality3-4 yr post-logging (Silvaet al. 1995,
F1 & Camacho1999)
Fegan
) 4-fold increaseintree mortality2 yrs post-logging (Thiollay1992)
) Moreresourcesavailableto residualtreesb/c of reductionin
canopycoverandstem density(Maitre1991);but damaged trees
mayneedto useresourcesto healw/ netresultof increasein
abortedfruits(Stephenson1981)
> Potentiallyhigherseedpredationlevels(Schupp1990,C. Peters,
pers.comm.)
> Temporaryincreaseingrowthratesdependingon extentof
canopyopening(WanRazali1989)
> Meandbh increment2-3x higherin loggedvs.unloggedJonkers
1987)
> Lower growth rates after thinning + selective loggingin (E.
Malaysia:
Primacket al. 1989)
> Lowergrowthratefor Araucaria
cunninghmamii
(Australia:Enright
1978)
Non-rodent > Civetdensitiesin loggedforestwere only
mammals 20%of 1°(Malaysia, 2-12yrs post
selectivelogging,no hunting:Heydon&
Bulloh1996);butincreased when<33-
50% forestimpacted(Malaysia, 5-10yrs
postlogging:Johns1983b)
> Tree shrewdensititesgreater40 yrs post
MUStreatmentthan 10 forest(Malaysia:
Laidlaw1998)
> Giantforesthogpopulations declineafter
logging(Davieset a].in press)
> Elephantpopulations increasein response
to logging(Africa& Asia:GrieserJohns
1997)
> Mousedeer (Tragulus) reduced
populations
afterlogging(80-100m3/ha,Malaysia:
Heydon& Bulloh1997)
&
Rodents > showedvariable
Squirrel& rat populations
Marsupials responses
& species-specific to MUS
Laidlaw
40 yrs post-treatment:
(Malaysia,
1998)
Bats > Vampirebatdensitiesincreaseif cattle
introducedto loggedareasaohns et al.
1985,Wilkinson1985,Johns1988,1992a,
Fentonet al. 1992)
Birds > SeeAppendixIV
Herps
Insects
Soilorganisms
Appendix VI
Primate Responses (by feeding guild)
to Logging in Tropical Forests

Biodiversity Series - Impact Studies 57

 Primate responses(by feedingguild) to loggingin tropicalforests;O=no change;- = decreaseafter logging;+ = increaseafter logging
GUILD/ SPECIES UluSegamaNangaGaat Tekam Sg.Lalang Kemasul Pta da Kibabe Budongo Kalinzu Kirindy Lope
Castanha
Logging Intensity(trees/ha) 20 10 18 > 20 >20 5.1 or 7.4 6-25 ? . I
Damagelevels(% tree loss) 32-58 54 5 38 55 60 25or 50% >50 ? II
Years since logging 6-12 0-4 0-12 20-30 20-30 11 12-28 1-50 4-20 1-15
Omnivores
Cebus apella +
Cebus albifrons +
Lophocebus albigena 0/- 0
CercopithecusIhoesti O_0
Cercopithecusascanius +1- + + I
Cercopithecuscephus 0
Cercopithecusmedius +
mitus
Cercopithecus _ 0/- + 0
Cercopithecuspogonias 0
Cercopithecus
nictitans . 0
Ga/ago s_p .
Gorilla gorila 0
Macaca spp _
Macaca SPP _
Macaco fascicularis 0 + .
MacOca nemestrina 0 0 + 0
Mandrillus sphinx 0
Microcebus spP +
Pan trogtodytes 0/- 0 +
Perodicticus potto .
Phaner furcifer _ +
Pongo pygmaeus +
Saguinus mystax +
Saimiri sPP _ +
Folivores/frugivores
Alouatto seniculus _
Ateles paniscus 0
Callicebus moloch +
Callicebus torguatus __ __ +
Procolobus badius 0/-
Colobus guereza . + + 0
Colobus satanus 0
Presbytis hosei + .
Presbytis melalophos + _
Presbytis obscura +
Presbytis rubicunda . .
 Frugivores/folivores I I l_l
Hylobates tars +
Hylobates muelleri 0
_ Lagothrix lagotricha
Pithecia albicans + _
References Johns Dahaban Johns Laidlaw 1994 Laidlaw 1994Johns 1991b Skorupa 1986, Plumptre & Howard Ganzhorn White 1992,
1989a,b 1996 1989a,b 1988;Weisenrseel Reynolds 1991; 1995; 1994
et al. 1993; 1994 Hashimoto Ganzhorn et
Chapman et al. 1995 al. 1990

* Modified from Plumptre and Grieserjohns (in press)

u-
Appendix VII
Impacts of Logging (and other
silviculture treatments where noted)
on the Genetic Component of
Biodiversity in Tropical Forests

Biodiversity Series - Impact Stutdies 61

 Impactsof logging(and other silviculturetreatments where noted) on the geneticcomponentof biodiversityin tropical forests

GeneticAttributes
Taxa Structure Composition Function
Vegetation P Geneticdiversity decreasedby ' Outcrossingrates unchangedin Shorealeprosula(Chan 1981,
5.0-23.4% (logged I yr prior; Lee et al. 1996,Wickneswari et al. 1997)& Dryobalanops
regeneratedstandsshowed no aromatica(Kitamura et al. 1994)
significantloss);effect on dioecious Outcrossingrates decreasedin Shoreamegistophylla(Murawski
speciesperhapsunderestimated et al. 1994a,b);density of mature individualslikely an important
(Wickneswariet al. 1997) factor

> Reducedcross-pollinationrates in Shoreasiamensisafter logging

(Ghazoulet al. 1998)and reduced pollination successof
Dipterocarpusobtusifoliusafter loggingS. siamensis(Ghazoul
1999)
P Outcrossing ratesof Carapaspp. were lower after controlled
experiments in loggedvs. unloggedforests in FrenchGuiana
(Doligez& Joly 1997) but were not significantlydifferent in
Carapastands in Costa Rica(Hall et al. 1994)possiblybecause
of very high stand densitiesin Costa Rica.
P Studiesciting genetic erosion of Swieteniamahoganiifrom
logginghave not been substantiatedin follow-up studies
(reviewed in Lugo 1999)
Primates
Non-rodentmammals
Rodents& Marsupials
Bats
Birds
Herps
Insects
Soilorganisms _s_ _
Glossary'
Advanced regeneration: seedlings or saplings
present in the understory prior to logging or
other silvicultural treatment
Aerial logging: a timber yarding system using
suspended cables, helicopters, or balloons to lift
logs
Afforestation: establishment of forests in areas
that did not historically support them (e.g., in
savannas or drained marshes)
Bole: trunk or main stem of a tree
Buffer zone: vegetation maintained on the
borders of roads, wetlands, or other ecologically
sensitive areas to mitigate the impacts of
management activities
Bulk density: the weight per unit volume of a
material (often soil)
Carbon sequestration: carbon storage
Coppice: sprouting of trees from stumps or
roots
Cutting cycle: the time (years) between stand
entries for timber extraction; there can be more
than one cutting cycle in a "rotation"
Dendrology: the study of trees including their
identification
Discounting: adjusting a future value by
dividing by a function of the (generally
BiodiversitySeries- ImpactStudies
compound) interest rate; used to calculate the
"net present value"
Edaphic: related to or caused by particular
conditions in the soil
Enrichment planting: increasing the density of
desirable species by interplanting, planting in
gaps, or planting along cleared lines
Fragmentation: the process of breaking once
continuous expanses of an ecosystem type
(typically forest) into island-like patches in a
matrix of other land uses
Gap: generally refers to a hole in the canopy
created by the death of a canopy tree, but there
are also understory gaps and belowground gaps
Girdle: to make an incision around a tree trunk
at least as deep as the cambium with the
intention of killing the tree; if combined with
arboricide application then referred to as
"poison girdling"
Group selection: harvesting of trees in clusters
generally no wider than twice the height of the
mature trees to promote regeneration of
moderately light-demanding species (a
"polycylic" system)
Hauling: carrying logs by truck (lorrie) from the
forest to processing or exporting facilities
Heartrot: decomposition of the central
stemwood of living trees typically due to fungal
63
Biodiversity Conservation in the Context of Tropical Forest Management
infection after mechanical or fire-related
damage
Heterozygous: having two different alleles at
the same locus on homologous chromosomes;
an index of genetic diversity
High-grading: removal of the best trees often
leaving a residual stand dominated by trees of
poor form and non-commercial species;
"creaming"
Highlead yarding: a yarding system in which
logs are skidded along the ground to a central
point by a cable passing through a block at the
top of a tower or spar tree (not to be confused
with "skyline yarding")
Liana: woody vine
Liberation thinning: a silvicultural treatment in
which future crop trees are released from
competition typically by girdling near neighbors
(note: term used differently by many North
American foresters)
Logging intensity: can refer to either the timber
volume or number of trees harvested per unit
area
Natural forest management: management of
forests for timber, non-timber forest products,
and environmental services by relying
principally on natural regeneration (compare to
plantation forestry)
Neotropics: American tropics
Net present value: the current value of some
future cost or benefit
Paleotropics: tropical areas in Africa, Asia,
Australia, and the Pacific Islands
Plantation: a stand of trees established by
planting; tree farm
64
Reduced-impact logging: a set of techniques
designed to avoid excessive damage to soil or to
the residual stand during and after timber
harvesting; typical components include pre-
planned skid trails, directional felling, and
engineering specifications for stream crossings
Reforestation: the recovery of forests in areas
that were deforested; compare to "afforestation"
Regeneration method: a timber harvesting
method that promotes development of a new
age class; the principal methods include
clearcutting, seed tree, shelterwood, selection,
and coppicing
Riparian: a terrestrial area bordering a water body
Rotation: the period between establishment of
regeneration and final harvesting in even-aged
stands
Shelterwood method: a stand regeneration
method that occurs in two phases, a first phase
in which enough timber is harvested to promote
development of a new age class in partial shade,
and a final felling of the remaining mature
timber after regeneration is established
Silviculture: the art and science of controlling
the establishment, growth, composition, and
health of forests and woodlands
Skidding: yarding logs with a rubber-tired self-
propelled machine or bulldozer ("snigging" in
Australia)
Skyline yarding: a log yarding system in which
logs are partially or entirely suspended from a
taut cable (not to be confused with "high-lead
yarding")
Stand: a contiguous group of trees with a more-
or-less similar history of disturbance, species
composition, size-class distribution, and
structure
Environment Department Papers
 Glossary

Succession: the gradual change in structure and Yarding: the process of extracting logs from the
composition as an area recovers from stump to the roadside, river edge, or other site
disturbance or on previously un-vegetated areas from which they are hauled

Thinning: a silvicultural treatment designed to

reduce stand density Note

Tractor: can refer to a bulldozer (=powered 1. Based heavily on Helms (1998).

vehicle with crawler tracks) or a farm tractor

Biodiversity Series - Impact Studies 65

Bibliography
Abdulhadi, R., K. Kartawinata, and S. Sugardjo.
1981. Effects of mechanized logging in the
lowland dipterocarp forest at Lempake, East
Kalimantan. Malaysian Forester 44:407-418.
Alexander, 1., N. Ahmad, L.S. See, A.G. Marshall,
and M.D. Swaine. 1992. The role of
mychorrhizae in the regeneration of some
Malaysian forest trees. Philosophical
Transactions of the Royal Society of London.
Series B, Biological Sciences 335:379-388.
Appanah, S., and M.R.A. Manof. 1991. Smaller trees
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