THE ECOLOGY, MANAGEMENT AND MARKETING OF NON

The Pennsylvania State University
The Graduate School
Intercollege Graduate Degree Program in Ecology
THE ECOLOGY, MANAGEMENT AND MARKETING OF NON-TIMBER
FOREST PRODUCTS IN THE ALTO RIO GUAMÁ INDIGENOUS RESERVE
(EASTERN BRAZILIAN AMAZON)
A Thesis in
Ecology
by
James Campbell Plowden
 2001 James Campbell Plowden
Submitted in Partial Fulfillment
of the Requirements
for the Degree of
Doctor of Philosophy
December 2001
We approve the thesis of James Campbell Plowden.
Date of Signature
Christopher F. Uhl
Professor of Biology
Chair of the Intercollege Graduate Degree Program
in Ecology
Thesis Advisor
Chair of Committee
James C. Finley
Associate Professor of Forest Resources
Roger Koide
Professor of Horticulture Ecology
Stephen M. Smith
Professor of Agricultural Economics
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ABSTRACT
Indigenous and other forest peoples in the Amazon region have used hundreds of non-timber forest
products (NTFPs) for food, medicine, tools, construction and other purposes in their daily lives. As these
communities shift from subsistence to more cash-based economies, they are trying to increase their harvest and
marketing of some NTFPs as one way to generate extra income. The idea that NTFP harvests can meet these
economic goals and reduce deforestation pressure by reducing logging and cash-crop agriculture is politically
attractive, but this strategy’s feasibility remains in doubt because the production and market potential of many
NTFPs remains unknown. The need to obtain this sort of information is particularly important in indigenous
reserves in the Brazilian Amazon since these comprise the country’s largest category of protected forests, and
indigenous people are actively seeking economic means to support community development.
Between 1996 and 2000, I spent a total of two years in the Alto Rio Guamá Indigenous Reserve in
eastern Pará state, Brazil studying production ecology of five regionally important NTFPs: oleoresin from
copaiba trees (Copaifera spp.), resin from breu trees (Protium spp.), aerial roots from titica vines (Heteropsis
spp.), latex from amapá trees (Parahancornia amapa and Couma guianensis), and seed oil from andiroba trees
(Carapa guianensis). My studies were primarily based in the Tembé Indian village of Tekohaw that is a one or
two day journey by river and road from the regional center of commerce in Belém. The objectives of NTFP
case studies on copaiba, breu and titica were to: 1) quantify the amount of marketable product that could be
obtained per plant and per area of forest, 2) identify key factors that influenced variation in these amounts, 3)
estimate the amount of harvest that would be possible on a repeated basis and the length of this harvest cycle,
4) estimate the amount of time a harvester needs to spend to find, harvest and process a unit of product and
how much income they would earn for this time invested.
Results showed that a variety of factors contributed to the low economic returns that could be
obtained by harvesting the NTFPs investigated in this project. Copaiba oleoresin had a relatively high unit
price, but the availability of the product was severely restricted by the low density of the trees (<1 tree/ha), the
limited percentage (15%) of trees that yielded more than 50 ml of oleoresin when drilled, and a sharp drop-off
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in oleoresin yield in trees that were retapped one year after the initial drilling. While the likelihood of
obtaining oil from these trees was greatest in trees in the 55-65 cm Diameter at Breast Height (DBH) group,
there was drop-off in oleoresin harvest in even larger trees because most were hollow and had apparently lost
their capacity to store oleoresin in enclosed trunk cavities.
Breu trees had a density of 10 trees per ha, and about 40% of trees yielded some resin. The amount of
resin increased with tree size and was related to the degree of attack by a bark-boring weevil whose larvae
develop inside the resin lumps that form on the trunk. The larvae of a syrphid fly are also found in resin lumps
from some types of breu trees, but it is unlikely these flies stimulated resin flow. Breu resin loses about 17%
of its weight while drying, but some of this may be due to resin removal by stingless bees. The low market
price for the resin makes it difficult for harvesters to regularly earn the Brazilian minimum wage (about $US
3.50 per day in 1999), but since it will take about four years for resin levels to rebound to initial harvest levels,
it is a resource that if managed wisely can provide a periodic source of income.
Titica vines are very common in some forests and can be found on as many as several hundred host
trees per ha. To meet the demands of wicker furniture makers, harvesters only take relatively thick mature
aerial roots. While harvesters may gather 10 to 40 kg of roots per day, the removal of bad pieces, stripping of
root cortex, and drying reduces the commercial product to 20% of the harvested weight. This processing of
roots can take as much time as the harvesting. Among the products studied, titica offers the best short-term
revenue, but since many harvested roots die or fail to regrow after being broken from the main plant, it may
take decades for a titica population in an area to return to a harvestable condition.
These case studies showed that it was difficult to find NTFPs that met all the conditions (high density,
high yield, low percentage of loss during processing, high price, quick resource renewal) to make their
commercial collection worthwhile to Tembé harvesters on an ongoing basis. Better understanding NTFP
production capabilities and the factors that influence them will help communities realistically assess how much
each resource can contribute to their annual income and how harvesting can be managed to preserve each
source population as a long-term resource. An increase in demand for consumer goods, however, will
probably necessitate greater investments in intentional NTFP planting and other economic activities.
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TABLE OF CONTENTS
LIST OF FIGURES
LIST OF TABLES
…………………………………………….………………………………………..
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LIST OF APPENDICES
ACKNOWLEDGEMENTS
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CHAPTER 1 INTRODUCTION
………………………………………………………………...........
1
INTRODUCTION ……………………………………………………………………………….........
SELECTION AND BACKGROUND OF STUDY AREA ………..…………………………………
SELECTION OF STUDY SPECIES ………………………………………………………………….
REFERENCES .......…………………………………………………………………………….
2
5
7
10
CHAPTER 2 THE ECOLOGY OF COPAIBA (COPAIFERA SPP.) OLEORESIN HARVEST
IN THE AMAZON …….……………………….………………………………………………………
13
ABSTRACT .………………………………………………………………………………………….
INTRODUCTION: REVIEW OF COPAIBA PRODUCTION AND USE ….……………………….
RESEARCH OBJECTIVES ….……………………………………………………………………….
STUDY SITE ……………….………………………………………………………………………...
METHODS …………………….……………………………………………………………………...
RESULTS ……………………………….…………………………………………………………….
DISCUSSION ……………………………………….………………………………………………...
SUMMARY AND CONCLUSIONS ………………………….……………………………………...
REFERENCES ………………………………………………………………….…………………….
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CHAPTER 3 THE HARVEST OF BREU RESIN FROM BURSERACEAE TREES IN THE
EASTERN BRAZILIAN AMAZON AND THE ROLE OF WEEVILS AND BEES IN ITS
FORMATION AND MARKETING ………………….………………………………………………..
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ABSTRACT ………………………………………….………………………………………………..
82
INTRODUCTION …………………………………….………………………………………………. 83
STUDY AREAS ………………………………………………………………………………………
85
METHODS ……………………………………………………………………………………………
86
RESULTS ……………………………………………………………………………………………..
90
DISCUSSION ………………………………………………………………………………………… 104
CONCLUSIONS ……………………………………………………………………………………… 118
REFERENCES ………………………………………………………………………………………... 142
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CHAPTER 4 THE ASSOCIATION OF AN ALIPUMILIO FLY (DIPTERA: SYRPHIDAE)
WITH BURSERACEAE TREE RESINS IN THE EASTERN BRAZILIAN AMAZON ………….
153
ABSTRACT …………………………………………………………………………………………...
INTRODUCTION ……………………………………………………………………………………..
STUDY SITES ………………………………………………………………………………………...
MATERIAL AND METHODS ……………………………………………………………………….
RESULTS ……………………………………………………………………………………………...
DISCUSSION …………………………………………………………………………………………
REFERENCES ………………………………………………………………………………………..
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CHAPTER 5 THE ECOLOGY, HARVEST AND MARKETING OF TITICA VINE ROOTS
(HETEROPSIS SPP.: ARACEAE) IN THE EASTERN BRAZILIAN AMAZON …………………
167
ABSTRACT …………………………………………………………………………………………...
INTRODUCTION ……………………………………………………………………………………..
STUDY AREA ………………………………………………………………………………………...
METHODS …………………………………………………………………………………………….
RESULTS ……………………………………………………………………………………………...
DISCUSSION …………………………………………………………………………………………
CONCLUSIONS: CHALLENGES FOR SUSTAINABLE TITICA HARVEST ...…………………..
REFERENCES ………………………………………………………………………………………...
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CHAPTER 6 SUMMARY CONCLUSIONS
………………………………………………………… 220
FACTORS THAT AFFECT NON-TIMBER FOREST PRODUCT HARVEST AMOUNTS AND
PROFITABILITY ………………………………………………………………………………….
RESEARCH IMPLICATIONS FOR THE ROLE OF NON-TIMBER FOREST PRODUCTS IN
FOREST COMMUNITY DEVELOPMENT ……………………………………………………...
REFERENCES ………………………………………………………………………………………...
EPILOGUE
221
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………………………………………………………………………………………………
241
CHALLENGES AND LESSONS STUDYING NON-TIMBER FOREST PRODUCT ECOLOGY
IN AN INDIGENOUS COMMUNITY ……………………………………………………………
REFERENCES ………………………………………………………………………………………..
242
252
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LIST OF FIGURES
FIGURE 2.1 CROSS-SECTION ILLUSTRATION OF SOLID, OLEORESIN BEARING AND
HOLLOW COPAIBA TREES ……………………………………………………………………….
FIGURE 2.2 PERCENT OF HOLLOW AND FIRE DAMAGED COPAIBA TREES AT TEKOHAW
FIGURE 2.3 HEARTWOOD AND HOLLOW PROPORTION OF TRUNK AREA IN COPAIBA
TREES AT TEKOHAW SITE ……………………………………………………………………….
FIGURE 2.4 OLEORESIN HARVEST FROM COPAIBA TREES AT TEKOHAW BY SIZE CLASS
FIGURE 2.5 PATTERNS OF OLEORESIN FLOW FROM COPAIBA TREES AT TEKOHAW …..
FIGURE 3.1 RELATIONSHIP OF DISTANCE FROM VILLAGE TO BREU RESIN HARVEST ...
FIGURE 3.2 DISTRIBUTION OF WEEVIL LARVAE HEAD CAPSULE WIDTHS FROM BREU
RESIN LUMPS ………………………………………………………………………………………
FIGURE 3.3 ILLUSTRATION OF BREU RESIN LUMP AND ASSOCIATED INSECTS ………...
FIGURE 3.4 ESTIMATED GROWTH RATE OF INDIVIDUAL BREU RESIN LUMPS AT
TEKOHAW …………………………………………………………………………………………..
FIGURE 3.5 BREU RESIN WEIGHT LOSS IN OPEN OUTSIDE DRYING AT TEKOHAW ……..
FIGURE 3.6 BREU RESIN WEIGHT LOSS WITH AND WITHOUT STINGLESS BEES ………...
FIGURE 3.7 ESTIMATED FIRST TIME RESIN HARVEST AT DIFFERENT DENSITIES OF
BREU TREES ………………………………………………………………………………………..
FIGURE 3.8 ESTIMATED DAILY HARVEST AND SALE VALUE OF DRIED RESIN AT
DIFFERENT DENSITIES OF BREU TREES
FIGURE 5.1 ILLUSTRATION OF TITICA SEEDLING AND ADULT ON HOST TREE WITH
PRINCIPAL PLANT PARTS ………………………………………………………………………..
FIGURE 5.2 ILLUSTRATION OF TITICA ABSORBING ROOT AND PRINCIPAL PARTS ……..
FIGURE 5.3 DENSITY OF TITICA HOST TREES AND TITICA ROOTS IN LIGHTLY TO
HEAVILY USED FOREST SITES IN THE ALTO RIO GUAMÁ INDIGENOUS RESERVE …...
FIGURE 5.4 NUMBER AND TYPE OF TITICA ROOTS PER HOST TREE AT MEDIUM AND
LIGHT-USE FOREST SITES AT TEKOHAW AND CAJUEIRA …………………………………
FIGURE 5.5 RELATIONSHIP BETWEEN HOST TREE DBH AND THE NUMBER OF TITICA
ROOTS PER HOST TREE AT THE TEKOHAW MEDIUM-USE SITE …………………………..
FIGURE 5.6 DEVELOPMENT TIME AND MORTALITY OF IMMATURE TITICA ROOTS AT
THE TEKOHAW MEDIUM-USE SITE …………………………………………………………….
FIGURE 5.7 CONDITION OF TITICA ROOTS 7 MONTHS AFTER CUTTING AT TEKOHAW
MEDIUM-USE SITE ………………………………………………………………………………...
FIGURE 5.8 LOSS OF TITICA ROOT LENGTH AFTER NODE REMOVAL FROM 37 HOST
TREES HARVESTED AT CAJUEIRA ……………………………………………………………..
FIGURE 5.9 WEIGHT LOSS DURING PROCESSING OF TITICA ROOTS ………………………
FIGURE 5.10 WEIGHT LOSS OF TITICA ROOTS FROM DRYING AFTER CORTEX REMOVAL
FIGURE 5.11 LOCATION OF TITICA HOST TREES HARVESTED IN 4 DAYS BY A
COMMERCIAL COLLECTOR AT CAJUEIRA ……………………………………………………
FIGURE 5.12 LOCATION AND ORDER OF TITICA HOST TREES HARVESTED BY A
COMMERCIAL COLLECTOR AT CAJUEIRA IN ONE DAY ……………………………………
FIGURE 5.13 TITICA ROOT STEM STRIPPING RATE BY AGE OF PROCESSOR ……………..
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LIST OF TABLES
TABLE 2.1 COPAIBA TRUNK DIMENSIONS OF ALL DRILLED, OLEORESIN YIELDING AND
HOLLOW TREES AT TEKOHAW...........................................................................................................58
TABLE 2.2 CORRELATION BETWEEN COPAIBA TREE DIAMETER AND OTHER TRUNK
DIMENSIONS AT TEKOHAW STUDY SITE.........................................................................................59
TABLE 2.3 HARVEST OF OLEORESIN FROM COPAIBA TREES AT TEKHOHAW STUDY SITE
BY TREE TYPE.........................................................................................................................................60
TABLE 2.4 HARVEST OF OLEORESIN FROM COPAIBA TREES AT TEKOHAW STUDY SITE
BY SIZE CLASS ........................................................................................................................................61
TABLE 2.5 CHANGES IN OLEORESIN HARVEST FROM COPAIBA TREES AT TEKOHAW
STUDY SITE BY YEAR AND HARVESTING ORDER .........................................................................62
TABLE 2.6 DRILLING TIMES IN COPAIBA TREES AT TEKOHAW........................................................63
TABLE 2.7 OLEORESIN PRODUCTION FROM C. MULTIJUGA NEAR MANAUS, BRAZIL
64
TABLE 2.8 COPAIBA OLEORESIN HARVESTING MODEL BY AREA FOR LONG-TERM AND
SHORT-TERM PROJECTIONS
65
TABLE 2.9 COPAIBA OLEORESIN HARVESTING MODEL FOR TWO COLLECTION METHODS
AT FIRST-TIME HARVEST.....................................................................................................................66
TABLE 3.1 SUMMARY OF RESEARCH OBJECTIVES AND METHODS ...............................................121
TABLE 3.2 DENSITY OF BREU TREES IN DIFFERENT FOREST TYPES AT TEKOHAW ..................122
TABLE 3.3 BREU RESIN HARVEST PER TREE BY STATUS, TREE TYPE, LIVE TREE SIZE
(DBH), FOREST TYPE, AND DISTANCE FROM TEKOHAW ...........................................................123
TABLE 3.4 DIMENSIONS OF WEEVILS FOUND IN BREU RESIN LUMPS ..........................................124
TABLE 3.5 BREU RESIN LUMPS WEIGHT CORRELATION TO WEEVIL LARVAE AND BORE
HOLE DIMENSIONS ..............................................................................................................................125
TABLE 3.6 WEEVIL LARVAE DIMENSION CORRELATION TO BORE HOLE NUMBER AND
DIMENSIONS..........................................................................................................................................126
TABLE 3.7 RESIN LUMP NUMBER AND WEIGHT FOUND ON BREU TREES DURING
FOLLOW-UP HARVESTS......................................................................................................................127
TABLE 3.8 CORRELATION OF TREE SIZE AND HARVEST HISTORY WITH FOLLOW-UP
BREU RESIN HARVEST........................................................................................................................128
TABLE 3.9 PROJECTION OF BREU RESIN YIELD AFTER INITIAL HARVEST ..................................129
TABLE 3.10 BREU RESIN HARVEST MODEL - PART 1. ESTIMATED AVERAGE RESIN PER
TREE BY SIZE CLASS ...........................................................................................................................130
TABLE 3.11 BREU RESIN HARVEST MODEL - PART 2. DENSITY OF RESIN YIELDING TREES,
RESIN HARVEST AND HARVEST PER VALUE PER HECTARE ....................................................131
TABLE 3.12 DENSITY OF BURSERACEAE RESIN YIELDING SPECIES IN BRAZILIAN
AMAZON INVENTORIES......................................................................................................................132
TABLE 3.13 BREU RESIN HARVEST - PART 3. RESIN HARVESTING RATES AND VALUE BY
AREA AND TIME ...................................................................................................................................133
TABLE 4.1 BODY DIMENSIONS OF IMMATURE ALIPUMILIO FLIES FROM BREU RESIN AT
TEKOHAW
163
TABLE 5.1 SUMMARY OF TITICA STUDY OBJECTIVES, METHODS AND ANALYSIS ...................201
TABLE 6.1 PRODUCTION ECOLOGY SUMMARY FOR CASE STUDY NON-TIMBER FOREST
PRODUCTS AND MANIOC...................................................................................................................231
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LIST OF APPENDICES
APPENDIX 2-A COMMON NAMES FOR COPAIFERA TREES IN THE AMAZON REGION
80
APPENDIX 3-A SCIENTIFIC SPECIES AND COMMON NAMES OF RESIN YIELDING SPECIES
OF NEOTROPICAL BURSERACEAE ...................................................................................................150
APPENDIX 5-A GLOSSARY OF TERMS FOR TITICA VINES AND ROOT HARVEST ........................219
APPENDIX 6-A PRODUCTION ECOLOGY STUDIES ON AMAPÁ LATEX AT TEKOHAW ...............234
APPENDIX 6-B PRODUCTION ECOLOGY STUDIES ON ANDIROBA OIL AT TEKOHAW ...............236
APPENDIX 6-C MANIOC AND FARINHA PRODUCTION STUDIES AT TEKOHAW ..........................238
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ACKNOWLEDGEMENTS
I would like to thank the Rainforest Alliance (Kleinhans Fellowship), National Science Foundation
(Graduate Fellowship), Food, Conservation and Health Foundation, and the Hotchkiss School Students for
Environmental Action for their financial support of this project. I express my deep gratitude to my major
adviser Dr. Christopher Uhl who provided expert guidance, moral and financial support to me and this project
from beginning to end. I appreciate my committee members Dr. James Finley, Dr. Roger Koide and Dr. Steve
Smith for always being available to help as was my scientific counterpart in Brazil Prof. Francisco de Assis
Oliveira. Other advice, assistance, identification of specimens, and translations were graciously provided by
Dr. William Balée, Dr. Steve Beckerman, Dr. Scott Camazine, Dr. Jason Clay, Sérgio Antonio da Silva, Dr.
Douglas Daly, Dr. James Duke, Dr. B. A. Foote, Maria Graça-Zogbhi, Dr. Joe Kiesecker, Dr. Jean
Langenheim, Judith Lisansky, Dr. James Marden, Dr. Denny Moore, Dr. Charles O’Brien, Dr. William
Overall, Dr. Graham Rotheray, Rafael Rueda, Noêmia Pires de Sales, Dr. Jack Schultz, Dr. Patricia Shanley,
Dr. F. Christian Thompson, Dr. Giorgio Venturieri, Frank von Willert, Museu Goeldi staff botanists, and the
entire staff of IMAZON. Tom Nagel did the life-life illustration of the insects associated with the breu resin
lump shown in Chapter 3. Frederico Miranda de Oliveira, Francisco Potiguara, José Maria Galvão, Claudionor
Diaz, Jescelino Bessa and other staff members from FUNAI (National Indian Foundation of Brazil) provided
crucial logistical support in Belém and the field. Jim Lockman and Edson Vidal from IMAZON kindly
arranged access to their research sites at Mojú and Fazenda 7.
I am indebted to the community leaders Lourival Tembé, Veronica Tembé, Muxi Tembé, Waldeci
Tembé, Augustin Tembé, Petroni Ka’apor and the people of Tekohaw and other villages in the Alto Rio
Guamá and Alto Turiaçu Indigenous Reserves for their support of the project and my family. This project
would not have been possible without the efforts of Livindo Tembé, Xarope Tembé, Afonso Tembé, Bubute
Tembé, Caroço Tembé, Emídio Tembé, Kapara’i Tembé, Manoel Tembé, Mukuim Tembé, Moreira Tembé,
Reginaldo Tembé, Pachik Tembé, Pedro Tembé, Preto Tembé, Raimundinho Tembé, Santana Tembé, Xina’i
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Tembé, Xuxa Tembé, Zeca Tembé, Zezinho Tembé, Zu Tembé, Aloisio Munduruku, Caetano Munduruku,
Luçiano Munduruku, Lourival Munduruku, Geraldo Ka’apor, Makú Ka’apor, Xu’i Ka’apor, Godo, Bruce
Hoeft, Haley Mitchell, Delores Coan, Danielle Szabo, Silvana, Cleber, César, Damião Farias, Manoel Lopes
Farias and Elaelson who worked with me in the village and forest between 1996 and 2000.
Special thanks are due to my children Marissa and Luke Plowden for their adventurous spirit and
adaptability during our year together in Brazil and for their incredible grace dealing with my absence for
almost another whole year when I completed my field work. My parents Theodore and Suzanne Plowden gave
my family and me unflagging love and support throughout several challenging years. I dedicate this
dissertation to my wife Yuri Kusuda Plowden who has shared six years of the joys and heartaches of this
journey and demonstrated more courage, patience, love, perseverance and creative support than one man could
hope to see and receive from a partner in a lifetime.
CHAPTER 1
INTRODUCTION
2
INTRODUCTION
My connection to the idea that people living in tropical forests could make a decent living by
sustainable harvesting and selling non-timber forest products (NTFPs) began in 1989 when I stayed in a small
house in Rio Branco, Brazil formerly occupied by the rubber tapper and union activist Chico Mendes. Only
four months before my visit to a forest reserve that now bears his name, he had been standing on his back
porch when a local rancher shot and killed him. This man and other large land owners were vehemently
opposed to Mendes’ promotion of collective land rights for rubber tappers who had worked in servitude to
others in these forests for many decades. The global outrage at his assassination propelled the Brazilian
government to move forward with creation of Extractive Reserves throughout their Amazon region. These
reserves were intended to help collectors of NTFPs such as rubber, Brazil nuts, and palm hearts improve their
standard of living and practice sustainable forest management (Fearnside, 1989; Allegretti, 1998). The political
appeal of a concept that could simultaneously reduce rural poverty and conserve endangered rain forests was
irresistible. The economic viability of this strategy, however, seemed untested until a paper co-authored by
Charles Peters, Alwyn Gentry and Robert Mendelsohn was published in the journal Nature (Peters et al.,
1989). Their study showed that forest peasants in the Peruvian Amazon could potentially make more money
from a hectare of forest in the long run by selling an assortment of NTFPs than selling the rights to cut down
the best trees to local loggers.
Case studies of NTFPs from the Amazon, Asia, and Africa (Anderson, 1990; Nepstad and
Schwartzman, 1992; Plotkin and Famolare, 1992) provided additional examples of forest peoples who were
making a living selling NTFPs without destroying the forest. Panayotou and Ashton (1992) popularized the
idea that NTFP harvesting could be integrated with timber management to generate revenue from the full range
of forest resources. Other books followed that provided lists of promising NTFP species and recommended
strategies for how to implement NTFP sustainable harvest and marketing (Bursztyn et al, 1993; Clay and
Clement, 1993; Peters, 1994; Warner and Pontual, 1994; Oliveira et al, 1995; SEBRAE, 1996; Wollenberg and
Ingles, 1998). Even NTFP promoters, however, acknowledged that making NTFP marketing the focal point of
3
tropical forest community development and forest conservation cannot succeed unless certain biological, social
and economic conditions are met. Long-term commercial harvests of NTFPs are problematic if 1) the resource
base is too small to be sustainably harvested, 2) the demand for the raw material is too low to generate a
reasonable price for collectors, 3) marketing logistics are too complicated, 4) demand is so high that production
from wild populations is replaced by plantation sources or other substitutes or 5) benefits from NTFP sales are
not distributed equitably within a community (Browder, 1992; Pendleton, 1992; Survival International., 1992;
Homma, 1993; Dove, 1994; Stiles, 1994).
Some of the most optimistic studies of NTFP economic viability have been done in seasonally flooded
(varzea) forests with dense concentrations of a few economic species in areas that have relatively easy access
to urban markets (Peters et al, 1989; Anderson, 1990). It has been harder to demonstrate NTFP commercial
viability in upland (terra firme) forests where the natural density of economic species is often low (Peters,
1994) and transporting products to market can be difficult and expensive (Homma, 1993). In theory, a
community with a diverse base of NTFPs could avoid some of the pitfalls described above by selling moderate
amounts of a variety of products (Clay and Clement, 1996) - an idea being explored in several Extractive
Reserves in Brazil (ELI, 1995; Murrieta and Rueda, 1995).
The goal of developing NTFP marketing in the context of community economic development with
forest conservation has so far received little attention in indigenous reserves (Rueda and Lisansky, personal
communication 1995). These efforts are especially needed in Brazil where such reserves cover 94.6 million ha more than three times the size of other protected areas in the country combined (Schwartzman, 1996). The
potential conservation value of these areas is enormous, but it cannot be assumed that these values will be
maintained in the long term without concentrated attention. Many indigenous lands have been destructively
exploited for mineral and timber resources or usurped for conversion to non-Indian pastures and farms. Even
where indigenous people have relatively secure control of their territory, traditional land use practices are changing
in response to groups’ desire to earn more income (Baksh, 1995; Stearman, 1995). Since increasing production of
agricultural cash crops can increase deforestation (Baksh, 1995), developing multiple NTFPs as a source of
revenue for indigenous people seems like an attractive option since many groups already use hundreds of rainforest
plant and animals species for subsistence purposes (Balée, 1994; Phillips et al, 1994).
4
It cannot be assumed, however, that every NTFP that has been collected for consumption in the
village will be worthwhile to harvest for sale. Making this determination will require information about the
plant’s basic ecology, handling requirements, sustainable harvest levels (La Frankie, 1994; Peters, 1994; Boot and
Gullison, 1995), economic returns (Godoy et al, 1993; Godoy and Bawa, 1993; Warner and Pontual, 1994; Shanley
et al, 1996), and conditions that favor plant regeneration (Martini et al, 1994; Laird, 1995). Studies that combine
traditional knowledge with an applied scientific approach could provide valuable information to forest peoples
that wish to evaluate the economic viability of specific NTFPs and the role that they can play in community
development (Shanley et al, 1996).
The overall aim of my research was to conduct production ecology studies on several economic
species of NTFPs in an indigenous community in the Brazilian Amazon. I secondly wished to quantify how
much income people were making or could make by harvesting and selling modest amounts of various NTFPs.
The specific objectives of each case study were to: 1) quantify the amount of marketable product that could be
obtained per plant and per area of forest, 2) identify key factors that influenced variation in these amounts, 3)
estimate the amount of harvest that would be possible on a repeated basis and the length of this harvest cycle,
and 4) estimate the amount of time a harvester needs to spend to find, harvest and process a unit of product and
how much income they would earn for this time invested. In some cases, I also examined the impact of current
harvesting techniques on harvested plants and their populations and suggested changes that might reduce
negative impacts. I avoided making a commitment to determine the “sustainable” harvest level since this
implied an even more comprehensive approach that would assess what impact a particular harvest level could
have on the target population, its dependent fauna and other members of the biotic community (Hall and Bawa,
1993). The results of three NTFP case studies will be presented in Chapters 2 (Copaiba oleoresin), 3 (Breu
resin), and 5 (Titica vine roots) of this dissertation. Some of the general conclusions will be discussed in
Chapter 6 – Summary Conclusions.
Apart from the scientific oriented goals described above, I wished to grow from the experience of
being a foreign researcher working on an applied ecology research project with a group of indigenous people.
I felt that there was so much I could learn about tropical forest ecology and peoples’ relationship to it by living
and working with people who had grown up in the forest. I also believed that I had knowledge, skills and
5
passions I was ready to share that would make a positive contribution to whatever community was willing to
take me in. The challenges that I encountered trying to balance the demands of doing high-quality scientific
work and being a good citizen in my host community are also discussed in the Epilogue. These experiences
provided even more powerful lessons for me than the results of the NTFP case studies themselves, and I hope
they will provide food for thought for other researchers.
SELECTION AND BACKGROUND OF STUDY AREA
The Alto Rio Guamá Indigenous Reserve is a 278,000 ha federal indigenous reserve in the Brazilian
Amazon (FUNAI, 1975). It is the main homeland for about 800 Tembé Indians who live in villages along the
Guamá and Gurupi Rivers in eastern Pará state (Sales, 1993). The Tembé use hundreds of forest products for
subsistence purposes (Balée, 1987) and have a long history selling a few forest and agricultural products (Arnaud,
1982; CEDI, 1985; Sales, 1994). The high biodiversity and deforestation threat in this and five other contiguous
indigenous and biological reserves have made the region a top conservation priority of the entire Amazon region
(Conservation International, 1991; Fearnside and Ferraz, 1995; IBGE, 1995).
In 1993, I visited several Tembé villages along the Guamá River to discuss my desire to study the
ecology and market potential of NTFPs in cooperation with an indigenous community. The Tembé leaders
were very cordial and frank with me. Their biggest concern was that ranchers, colonists and loggers had
illegally occupied large areas of their reserve, and day by day these incursions were getting closer to their
villages. They told me a few elders would take me around the small section of forest they still controlled, but
they said it would be futile to study its NTFP commercial potential. Invaders had deforested and taken over so
much of their territory that they could no longer feed themselves by hunting and fishing, and other forest
products were becoming scarce. They urgently needed new sources of income to buy food and basic items, but
they most needed help to regain control of their land. I said I would think about how I could assist with the
land issue and gratefully accepted their offer to let me tour their forest.
For two days my Tembé guides walked with me through the forest and pointed out more than a dozen
trees whose boiled bark treated various ailments. They pulled the plug from a drilled copaiba (Copaifera spp.)
6
tree to collect some liquid oleoresin, showed me how to get drinking water from a machete split vine, and
introduced me to the sweet aroma of resin on a fallen breu tree (Protium spp.). After tripping over a log, I
discovered that rubbing andiroba (Carapa guianensis Aubl.) seed oil on my bruised shin actually reduced the
pain and swelling as well as it kept mosquitoes off my exposed skin. It was exciting to finally get a first hand
look at the living plants that were the source of forest remedies I had only previously seen in bottles and string
tied bundles in the Belém city folk market. This enthusiasm was tempered by the realization that my hosts had
not exaggerated their plight. We reached an area of forest that had been burned by colonists after less than an
hour’s walk from the village. The copaiba tree we had taken oil from was the only productive tree the elders
knew of that was left in that part of the forest.
On my last night in São Pedro village, I was fortunate enough to witness and briefly take part in a
traditional Tembé style celebration. I finally climbed into my hammock late at night after the day’s events and
a relentless stomping dance left me exhilarated and exhausted. I drifted to sleep listening to the hypnotic
shaking of maracas and a chant about a giant macaw. The tune of this song and the half–learned lyrics kept
repeating themselves in my mind long after I came home. I did not know how it would happen, but I had a
strong feeling I would hear that chant again.
My next opportunity to connect with the Tembé came three years later in the spring of 1996 when my
high school invited me to give a presentation on Earth Day and also agreed to my request to sponsor the visit of
an indigenous leader from the Amazon. It seemed like a long-delayed chance to make good on my wish to
help the Tembé with their land rights’ concern. This time I made a contact with people working with villages
along the Gurupi River. Unlike their cousins in the northern section of the reserve, the Tembé in this area were
surrounded by large expanses of primary forest. They had abundant land available for hunting, fishing,
farming, and collecting forest products for use in the home and occasionally bringing to market. Gurupi
communities were also very aware of the problems caused by non-Indian invaders and were trying to confront
them from a position of strength. During a ten day trip to the U.S. in April 1996, the Tembé leader Muxi and I
gave presentations to twenty-four government agencies, conservation and human rights groups, schools and
churches. He enthusiastically endorsed my proposal to study marketable NTFPs in the southern section of the
7
reserve because villages there were anxious to increase their income to improve health care, education and
security of their land. I would be welcome to stay at his village called Tekohaw.
SELECTION OF STUDY SPECIES
The process of selecting candidate species for this research began in the markets of Belém. Located
near the mouth of the Amazon River, Belém is the main commercial center for the state of Pará and is the
transit point for many Amazonian products en route to the rest of Brazil and foreign ports. I first became
familiar with the Ver-o-Peso market that hosts the region’s largest concentration of forest product dealers. The
folk market section has primarily medicinal products that include barks, plant seeds and exudates (oil, resin,
latex), leaves, and various animal products. Other retail vendors sold an assortment of wild and cultivated
fruits, and a wholesale market devoted exclusively to açai palm (Euterpe oleracea Mart.) fruit was held every
morning. I later visited the CEASA wholesale produce market where businesses handle large volumes of
cultivated fruits and minor amounts of regional forest fruits collected from wild populations and small-scale
plantings.
When I arrived at Tekohaw village for the summer of 1996, Tembé leaders told me they had many of
the trees whose products I had seen in the market. The challenge was identifying which ones looked most
promising as species that I could study and that they could profitably harvest. While their forest had most of
the medicinal bark trees being sold in the market, I ruled out these products since dealers seemed to have all
the supply of these they needed. Learning how much bark could be removed from a tree without causing it too
much damage would be an interesting study, but the ready availability of massive quantities of bark obtained
from felled trees would leave little incentive for developing careful harvesting techniques. All of the medicinal
leaves sold in the market were cultivated, so these offered no opportunities as marketable forest products.
Some fruits had interesting commercial prospects, but the potential for damage or rotting during uncertain river
and overland transport from the reserve to the market was worrisome and gaining reliable data on fruiting
patterns could take many years of continuous observation (Wheelwright, 1986). The class of products that
initially seemed most feasible from a marketing and research standpoint was plant exudates including the latex
8
from amapá (Parahancornia amapa (Hub.)Ducke), seed oil from andiroba and trunk oleoresin from copaiba.
Their advantage was that they did not spoil in a short time, and dealers seemed willing to buy them with no
prior arrangement for $US 3 to 8 per liter with the highest prices offered for copaiba. Copaiba was also the
only one of these products that had an international market.
Since the Tembé said that copaiba trees were reasonably abundant in their forest and they had
experience collecting its oleoresin, I chose to focus on this tree for the summer of 1996 and did follow-up work
with it in subsequent years. The details of this case study are presented in Chapter 2. The initial season of
work with copaiba had two concrete results. First it showed me that I thoroughly enjoyed working with these
people and their forest, and they seemed to enjoy working with me. I was invited to come back for a longer
time and bring my family. Secondly, the research showed that copaiba trees were less common and yielded
less oleoresin than the Indians had thought. The next phase of the research would need to include some other
species.
We tentatively decided to explore amapá latex , andiroba seed oil, cumaru (Dipteryx odorata L.) seeds
and seed oil, açai fruit, bacuri (Platonia insignis Mart.) fruit, and cupuaçu (Theobroma grandiflorum (Willd. ex.
Spreng.)Schum.) fruit as potential new market products. These products were all routinely marketed in the
region (Clay and Clement, 1993; Shanley, 1994) and commonly used by the Tembé as food or medicines
(Balée, 1987). The research would also quantify production ecology information on resin produced by breu
trees (mostly Protium spp.) and aerial roots from the titica vine (Heteropsis spp.) that were the two most
important NTFPs already being sold by the Tembé in the Gurupi region.
I returned to the reserve with my wife and two children in the fall of 1997 to take up this ambitious
research agenda. Cumaru was removed as a candidate species quickly because it was relatively rare, trees
fruited at different times, and fruit fall was spaced over many months. Bacuri trees were supposedly abundant
near the village of Rabo de Mukura, but studying them from a base at Tekohaw was not practical. Although
people from Tekohaw called one large section of forest near the village the “cupuzal” in supposed recognition
of its high density of cupuaçu trees, our surveys of the area found only two of these trees. Whether forest fires
that had passed through there in the early 1980s had killed many of these trees or it was just less naturally
common than they thought, it was not a large enough population to study.
9
Of all the marketable fruits, açai palm trees were by far the most abundant and important to the
Tembé. When these trees reached their peak fruiting from October through December, every Tembé family
devoted considerable time gathering the hard berries and preparing açai juice. I started production studies in
several plots, but the work was thwarted by two main factors. The first was that areas that were far enough
from the village to be left alone when fruits were abundant in the beginning of the fruiting season became too
attractive for some families not to harvest a few months later when fruits were harder to find. The second
problem was that a fire that escaped from a farm field swept through one research plot and caused all the fruit
in it to abort. Other challenges conducting ecological research in the heart of a community forest will be
discussed in Chapter 6 – Summary Conclusions. Some production ecology work was also conducted with
amapá and andiroba. These studies were not complete enough to present as full case studies in this
dissertation, but a few highlights from work with these species will be discussed in Chapter 6.
It seems somewhat ironic that my research did not end up focusing on new commercial NTFPs for the
Tembé but studying the two forest products that already earn them the most income – breu resin and titica vine
roots. As Chapters 3, 4, and 5 will demonstrate, these efforts that sought to blend indigenous knowledge and
beliefs with a rigorous data gathering approach provided the most new insights and hopefully most useful
results for ecology as a field of scientific inquiry and for people who are looking for practical means to make a
simple living in a forest.
10
REFERENCES
Anderson, Anthony B. 1990. ed. Alternatives to Deforestation: Steps toward Sustainable Use of the Amazon
Rain Forest. Columbia University Press, New York.
Baksh, Michael. 1995. Changes in Machigeunga quality of life. pp. 187-205 in Leslie E. Sponsel (ed.)
Indigenous Peoples and the Future of Amazonia: An Ecological Anthropology of an Endangered
World. The University of Arizona Press, Tuscon.
Balée, William. 1987. A etnobotânica quantitativa dos índios Tembé (Rio Gurupi, Pará). Boletim do Meseu
Paraense Emilio Goeldi: Botanica, Agosto (1):29-50.
Balée, William. 1994. Footprints in the Forest: Ka’apor Ethnobotany - the Historical Ecology of Plant
Utilization by an Amazonian People. Columbia University Press, New York.
Boot, Rene and Gullison, R.E. 1995. Approaches to developing sustainable extraction systems for tropical
forest products. Ecol. Applications 5(4): 896-903.
CEDI. 1985. Tembé. pp. 177-209 in Povos Indígenas no Brasil, No. 8: Sudeste do Pará. Centro Ecumênico de
Documentação e Informação, São Paulo.
Clay, Jason W. and Clement, C.R. 1993. Selected species and strategies to enhance income generation from
Amazonian forests. FAO Forestry Paper (Final Draft). United Nations Food and Agricultural
Organization, Rome.
Conservation International. 1991. Workshop 90: Biological Priorities for Conservation in Amazonia.
Conservation International, Washington, D.C.
Dove, Michael R. 1994. Marketing the rainforest: “green” panacea or red herring? Analysis from the EastWest Center No. 13, May.
Environmental Law Institute (ELI). 1995. Brazil’s Extractive Reserves: Fundamental Aspects of their
Implementation. ELI Research Report. ELI, Washington, D.C.
Fearnside, Philip M. 1989. Extractive reserves in Brazilian Amazonia. BioScience 39(6):387-393.
FUNAI. 1975. Situação Geográfica das Terras do “Posto Indígena Alto Rio Guamá” (map). Ministério do
Interior, Fundação Nacional do Índio (FUNAI), 2o Delegacia Regional, Belém, Pará.
IBGE. 1992. Atlas Naçional do Brasil. Fundação Instituto Brasileira do Geografia e Estatistica (IBGE), Rio de
Janeiro.
Godoy, Ricardo A. and Bawa, K.S. 1993. The economic value and sustainable harvest of plants and animals
from the tropical forest: assumptions, hypotheses, and methods. Economic Botany 47(3):215-219.
Godoy, Ricardo A., Lubowski, R. and Markandaya, A. 1993. A method for the economic valuation of nontimber tropical forest products. Economic Botany 47(3): 220-233.
11
Hall, Pamela and Bawa, K. 1993. Methods to assess the impact of extraction of non-timber tropical forest
products on plant populations. Economic Botany 47(3):234-247.
Homma, Alfredo Kingo Oyama. 1993. Extrativismo Vegetal na Amazônia: Limites e Oportunidades.
EMBRAPA-SPI, Brasília.
La Frankie, James V. 1994. Population dynamics of some tropical trees that yield non-timber forest products.
Economic Botany 48(3): 301-309.
Laird, Sarah. 1995. The natural management of tropical forests for timber and non-timber products. OFI
Occasional Papers No. 49. Oxford Forestry Institute, Oxford.
Murrieta, Julio R. and Rueda, Rafael P. 1995. Extractive Reserves. IUCN Forest Conservation Programme,
Gland. pp. 133.
Nepstad, Daniel C. and Stephan Schwartzman. 1992. eds. Non-Timber Products from Tropical Forests:
Evaluation of a Conservation and Development Strategy. Vol. 9. Advances in Economic Botany. The
New York Botanical Garden, Bronx.
Panayotou, Theodore and Peter S. Ashton. 1992. Not by Timber Alone: Economics and Ecology for Sustaining
Tropical Forests. Island Press, Washington, D.C.
Pendleton, Linwood H. 1992. Trouble in paradise: practical obstacles to nontimber forestry in Latin America.
pp. 252-262 in eds. Plotkin, Mark and Famolare, L. Sustainable Harvest and Marketing of Rain Forest
Products. Island Press, Washington, D.C.
Peters, Charles M., Gentry, A.H., and Mendelsohn, R.O. 1989. Valuation of an Amazonian rainforest. Nature
339:655-656.
Peters, Charles M. 1994. Sustainable harvest of non-timber plant resources in tropical moist forest: an
ecological primer. Biodiversity Support Program c/o World Wildlife Fund, Washington, D.C.
Plotkin, Mark and Lisa Famolare. 1992. Sustainable Harvest and Marketing of Rain Forest Products. Island
Press, Washington, D.C
Rodrigues, Roberto M. 1989. A Flora da Amazônia. CEJUP. pp. 463.
Sales, Noêmia Pires de. 1993. Pressão e Resistência: Os Índios Tembé-Tenetehara do Alto Rio Guamá e a
Relação com o Território. União das Escolas Superiores do Pará, Diretoria de Pesquisa, PósGraduação e Extensão, Depto. de Ciências Socias. Relatório Final.
Sales, Noêmia Pires de. 1994. Os Tembé no Alto Rio Guamá: Reelaborações Étnicas - Identidade e Território.
UNAMA - Universidade da Amazônia, Pró-Reitoria de Pesquisa, Pós-Graduação e Extensão, Depto.
de Ciências Socias. Relatório de Pesquisa.
Schwartzman, Steve. 1996. Brazilian indian lands threatened. Environmental Defense Fund electronic mail
alert.
Stearman, Allyn MacLean. 1995. Neotropical foraging adaptations and the effects of acculturation on
sustainable resource use. pp. 207-224 in Leslie E. Sponsel (ed.) Indigenous Peoples and the Future of
Amazonia: An Ecological Anthropology of an Endangered World. The University of Arizona Press,
Tuscon.
12
Stiles, Daniel. 1994. Tribals and trade: a strategy for cultural and ecological survival. Ambio 23(2):107-111.
Survival International. 1992. “Rainforest harvest” projects harm, not help, Indian communities. Survival
International Press release June 15. London.
Warner, P.D.III. and Pontual, A.C. 1994. Manual de Comercialização de Produtos Florestais. GENESYS
(Gênero em Sistemas Econômicos e Socias), The Futures Group, Washington, D.C.
Wheelwright, Nathaniel T. 1986. A seven-year study of individual variation in fruit production in tropical birddispersed tree species in the family Lauraceae. pp.19-35 in eds. Estrada, Alejandro and Fleming, T.H.
Frugivores and Seed Dispersal. Dr. W. Junk Publishers, Dordrecht
Wollenberg, Eva and Andrew Ingles. 1998. eds. Incomes from the Forest: Methods for the Development and
Conservation of Forest Products for Local Communities. Center for International Forestry Research,
Bogor.
13
CHAPTER 2
THE ECOLOGY OF COPAIBA (COPAIFERA SPP.) OLEORESIN HARVEST IN
THE AMAZON
14
ABSTRACT
Oleoresin extracted from the trunk of many species of copaiba (Copaifera spp.)(Caesalpiniaceae) trees
in the Amazon basin and other parts of South America has been used as a medicine by indigenous people and
westerners for more than 400 years. The oleoresin is rich in volatile and non-volatile terpenoid compounds
produced in parenchyma cells in many parts of the tree. The material is deposited into resin canals in the
sapwood and can accumulate large amounts in cavities when these canals break down in the heartwood. Trees
have been harvested by cutting into the trunk with an axe and by drilling into the heartwood. Popular accounts
indicate that the best producing species can yield a range of 2.5 to 24 liters of oleoresin per tree with some
trees yielding as much as 60 liters at one time. The only previous systematic production study of a copaiba
species (C. multijuga Hayne), however, showed that average yields were 0.2 liters per tree during the initial
harvest, and that this average steadily approached zero on each of four successive harvests.
I studied copaiba production ecology in the Alto Rio Guamá Indigenous Reserve, Pará, Brazil to
evaluate the harvesting economics of this product as a means to increase income for Tembé Indians in the
reserve. The study involved drilling three varieties of copaiba trees one meter above ground level and
measuring oleoresin yields at the time of the initial harvest and two more times. The average oleoresin yield
for the first harvest was 0.073 liters per tree for all 57 trees that were drilled. The average first harvest yield
among trees that yielded some resin during the study was 0.230 liters per tree, but this amount rapidly
dwindled during successive harvests one and two years later. There were some differences among tree
varieties, but trees in the upper-middle size classes (45 - 65 cm DBH) were found to yield the most resin while
small trees (<45 cm DBH), very large trees (>65 cm DBH), and hollow trees of any size yielded little or no
oleoresin. Internal measurements showed that the size at which oleoresin yield first occurred corresponds well
to the size at which heartwood began to form. Resin yield began to drop off in very large trees in which
hollowness became increasingly common. Some small trees seemed to be prematurely hollow due to burn
damage sustained in an accidental fire that passed through the forest in the 1980s.
The natural low density of copaiba trees (e.g. 0.1 to 2 trees/ha) means that much more time may be
needed to initially locate trees than to harvest them. Where density and average production are both low,
15
commercial harvesting may not be viable. Knowledge of specific copaiba features and preferred habitats can
improve search efficiency for these trees that are widely dispersed in a diverse forest. Drilling two holes in a
tree can take one person 40 minutes and waiting to capture all oleoresin escaping from a large cavity could take
hours or days. With large flows, it would be more efficient to leave a collecting device in place and return
later. Unless densities, average production, and the number of potential harvests per tree are high, though,
copaiba oleoresin extraction will not yield more than a few dollars per day for one person in a 1000 ha area of
forest. Plantations of copaiba trees could dramatically increase harvesting efficiency, but it seems unlikely that
the price for the oleoresin alone would justify the expense of maintaining these cultivated trees for 70 years or
more until resin is potentially extractable in sizeable amounts. The existence of local markets for unprocessed
copaiba oleoresin and small businesses that distill and repackage it into several forms, however, shows that
favorable economic conditions for harvesting and marketing copaiba from wild populations do exist in some
parts of the Amazon. Additional studies are needed to document what species, densities, harvesting
techniques, and other conditions make these operations profitable and if they are extracting the raw material on
a repeated basis.
16
INTRODUCTION: REVIEW OF COPAIBA PRODUCTION AND USE
OLEORESIN PRODUCTION IN COPAIFERA TREES
Copaiba trees belong to the genus Copaifera in the family Caesalpiniaceae (or Fabaceae sub-family
Caesalpinoideae). The scientific name is derived from a combination of the Tupi Indian word for the resin
from the copaiba tree, “copai”, and the Latin verb, “fero”, meaning to bear or produce (Marafioti, 1970; Allen
and Allen, 1981). Authors normally attribute 28 to 40 species to this genus that includes both shrubs and trees
(Record and Mell, 1924; Allen and Allen, 1981; Dwyer, 1951). Most Copaifera plants are native to the
neotropics with at least 25 species represented in the Amazon region (Gentry, 1993). Another four or five
species are found in the African tropics (Record and Mell, 1924; Langenheim, 1973). Neotropical Copaifera
species produce liquid oleoresins while material found in African species can harden to a copal like substance
(Anderson, 1955; Langenheim, 1973).
Copaiba oleoresin is produced in the tree’s trunk, stems, and leaves. It consists of large but varying
amounts of volatile oils (primarily composed of sesquiterpene hydrocarbons usually including caryophyllene),
non-volatile resinous substances and small quantities of acids (Furnemore, 1926; Fernandes, 1949;
Langenheim, 1973; Maia et al. 1978; Leung, 1980; Herres et al, 1986; Duke, 1986; Figliuolo et al., 1987;
Budavari, 1989; Duke and Cellier, 1993; Braga et al., 1998; Monti et al., 1999; Cascon and Gilbert, 2000).
The combination of the sesqui- and diterpenes along with phenolics may be responsible for the plant’s
resistance to many insect and microbial attacks (Figliuolo et al., 1987; Feibert and Langenheim, 1988; Macedo
and Langenheim, 1989a, 1989b; Langenheim, 1990; Schultes and Raffauf, 1992). The compounds are
synthesized in parenchyma cells that line rounded pockets, cysts or elongated canals (Langenheim, 1973). The
resin is secreted into these spaces that are formed by schizogeny and/or lysigeny. Schizogeny leads to the
separation of cells, which increases their intercellular space to create thin capillaries. Canals up to two meters
long have been observed (Correa and Bernal, 1989). In Copaifera, these canals are located in concentric
circles in the secondary wood that may delineate seasonal growth (Record and Mell, 1924; Langenheim, 1973;
17
Alencar, 1982; Vetter and Botosso, 1989). In addition to its presence in these schizogenous ducts, the
oleoresin sometimes accumulates in lysigenous cavities formed by the breakdown of secretory cells. In the
center of the trunk these cavities may enlarge to the point where they contain gallons of oleoresin (Youngken,
1943; Osol and Farrar, 1947; Poucher, 1950).
About twenty Copaifera species from South America are known to be sources of oleoresin. The main
commercialized species are: C. reticulata Ducke (80% in the Brazilian Amazon), C. guianensis Desf. (10% in
Brazilian Amazon); C. multijuga Hayne (5% in Brazilian Amazon), C. officinalis L. (3-5% in Brazilian
Amazon), C. langsdorffii Desf., and C. martii Hayne (Allen and Allen, 1981; Sampaio, 1993; MMA/SCA et
al., 1998). Oleoresin has also been harvested in the region from: C. bijuga Hayne, C. canime Harms, C.
confertiflora Benth., C. coriacea Mart., C. cuneata Tul., C. duckei Dwyer, C. glycycarpa Ducke, C. longicuspis
Ducke, C. longifolia Huber, C. marginata Benth, C. pubiflora Benth, C. spruceana Benth, and C. venezuelana
Pittier and Harms (Osol and Farrar, 1947; Howes, 1949; Dwyer, 1951; Mors and Rizzini, 1966; Allen and
Allen, 1981; Duke, 1986; Sampaio, 1993; Cascon and Gilbert, 2000), and C. camibar Poveda and Sanchez
(Berry et al., 1997). The wide distribution of the genus and its broad utility is reflected in the numerous
common names it has throughout the continent (Appendix 2-A).
The chemical composition, color and viscosity of oleoresin varies widely from species to species.
The “Para” variety of copaiba oleoresin comes from several species including C. reticulata, C. multijuga and C.
martii (Osol and Farrar, 1947). These are most common in Pará and Amazonas states in Brazil. Since C.
duckei is closely related to C. reticulata (Dwyer, 1951), its oleoresin is probably also included in this group.
The oleoresin of these species in general has the highest volatile oil content (35 - 90%) and greatest fluidity
among the commonly harvested copaiba trees (Youngken, 1943; Osol and Farrar, 1947). C. reticulata
oleoresin emerges from the tree as a clear, colorless and thin liquid, but on exposure to the air it turns a thick
yellowish brown and has a disagreeable smell (Le Cointe, 1947; Allen and Allen, 1981; Duke, 1986). C.
multijuga is fluid, pale-yellow to transparent and has a light agreeable smell (Le Cointe, 1947; Allen and Allen,
1981) with 58 - 78% essential oils (Furnemore, 1926). A recent analysis found that total sesquiterpenes
(hydrocarbons and oxides) accounted for 80 - 97% of several samples in this species (Cascon and Gilbert,
2000). The botanical characters of the broadly defined species C. langsdorffii are often hard to distinguish
18
from C. reticulata and other Copaifera (Dwyer, 1951). This species including its many varieties is considered
the principle source of the more viscous copaiba oleoresin commonly known as the “Maranham” or “Rio de
Janeiro” variety. It has the consistency of olive oil with a range of 30 - 75% volatile oils (Sayre, 1906;
Kraemer, 1907; Furnemore, 1926). The “Maracaibo” variety of copaiba oleoresin principally comes from C.
officinalis and the related C. guianensis in the northern Amazon countries of Venezuela, Columbia and the
Guianas (Sayre, 1906; Osol and Farrar, 1947; Dwyer, 1951). It is a thick, dark, and coumarin scented product
that has a 40 - 72% range of volatile oils in Surinam and 20 - 40% volatile component in Columbia (Sayre,
1906; Furnemore, 1926; Le Cointe, 1947).
The groupings of Copaifera species that have progressively denser oleoresin are not geographically
distributed in the same order. C. langsdorffii with its many variants is found in subtropical dry through wet
forest life zones from northern Argentina, through Paraguay and Brazil up into Venezuela and Guyana (Dwyer,
1951; Duke and Cellier, 1993). Of the major oleoresin producing species, it is the species most commonly
found in dry woodland areas (USDA, N.S.; Duke and Cellier, 1993). C. reticulata is found primarily in moist
tropical forests of Brazil and Bolivia (Dwyer, 1951). It grows best in flood-free areas of forest on sandy soils
(Duke, 1986). Farther to the north C. officinalis and C. multijuga likewise tolerate a variety of dry and wet
conditions. C. multijuga can be found in terra firme (non-flooded rainforest), varzea (flooded forests), sandy
margins near lakes and streams and even the cerrado (savannah) of central Brazil (Duke, 1986; Sampaio,
1993). It is most often found on extremely nutrient poor soils (Figliuolo et al, 1987). The main home range of
C. officinalis is in Venezuela, but it is also found in the West Indies and the Brazilian state of Bahia (Dwyer,
1951). Its relative C. guianensis is found in Brazil and its range also includes the Guianas, Columbia and
Panama (Dwyer, 1951). Although some leguminous non-woody plants and trees are known for their ability to
fix nitrogen through root nodules, Copaifera and other genera in its Detarieae tribe apparently do not possess
this capability and still persist well in nutrient poor soils (Figliuolo et al, 1987; Moreira et al., 1992).
COPAIBA ETHNOBOTANY, MODERN USE AND TRADE
Many Amazonian indigenous and traditional forest peoples including the Kaw, Tupi, Puinaves and
Makunas have used this chemically potent oleoresin as a traditional medicine (Record and Mell, 1924; Allen
19
and Allen, 1981; Schultes and Raffauf, 1990; Plotkin et al., 1991). The first western record of these trees was
in a 1534 report to Pope Leo X that referred to them as “copei.” A Portuguese monk writing about the natural
products of Brazil in 1625 mentioned a drug called “cupayba”. A tree yielding the substance was first
described and illustrated in 1648, and the London Pharmacopoeia listed the drug in 1677 (Youngken, 1943).
Linnaeus formally described the genus Copaifera in 1762 (Woodson and Schery, 1951). The oil was included
in the United States Pharmacopoeia from 1820 to 1940 when it was admitted to the National Formulary
(Youngken, 1943).
Copaiba was traditionally used in the Amazon region and abroad as a cicatrizant, anti-inflammatory,
and a treatment for chronic gonorrhea and skin ailments including herpes, sores, eczema and psoriasis. Internal
uses include treatment for a variety of ailments involving mucous membranes such as sore throat, bronchitis,
ulcers and various conditions of the genito-urinary tract (including cystitis, leucorrhea, syphilis and urinary
incontinence). It is also used as anti-rheumatic, antiseptic, anti-bacterial, diuretic, expectorant, hypotensive
agent, laxative, purgative, vermifuge and vulnerary (Youngken, 1943; Uphof, 1968; Leung, 1980; van den
Berg, 1984; Duke, 1986; Basile et al., 1988; Schultes and Raffauf, 1990; Duke and Vásquez, 1994; Viera,
1992; Fleury, 1997). Both C. coriacea and C. multijuga have been noted for various medicinal properties. The
species most consistently mentioned for effectiveness in treating tough urinary tract, pulmonary and skin
ailments are C. langsdorffii, C. officinalis, and C. reticulata. As mentioned, oleoresins from these trees are
thicker than other Copaifera species and have an acrid bitter taste (Le Cointe, 1947; Uphof, 1968; Rodriguez,
1989; Sampaio, 1993). To cut the bitter taste in internal preparations, it is sometimes mixed with andiroba oil
(from Carapa guianensis Aubl.), honey, santal oil, and cubebs (Viera, 1992).
While the advent of anti-biotics decreased foreign demand for copaiba as a treatment for venereal
disease (Osol and Farrar, 1947), modern studies have demonstrated the folk medicine’s effectiveness as an
anti-bacterial (Verpoorte and Dahl, 1987), anti-oxidant (Desmarchelier et al., 1997) and anti-inflammatory
agent (Basile et al., 1988). Today, copaiba oil is still commonly sold in folk medicine markets such as Ver-oPeso in Belém. It is also sold in various commercial preparations such as gel caps and included in several
Brazilian brands of “natural” cough syrup (MMA/SCA et al., 1998; Amazon Ervas, N.S.; Lab oratorio São
Lucas, N.S.).
20
Apart from medicinal uses, copaiba oleoresin has been widely used in other applications. The raw
material, distilled essential oils, and resinous compounds have made copaiba useful in the preparation of
cosmetics, lacquer, paints, varnishes and tracing paper (Youngken, 1943; Fernandes, 1949; Balsam and
Sagarin, 1974; Leung, 1980; Duke, 1986; MMA/SCA et al., 1998). Cosmetic preparations includes soaps,
detergents, creams, lotions, massage oil, and perfumes where copaiba oil or resin serves as a fragrance and
odor fixative (Poucher, 1950; Leung, 1980; Duke, 1986). Both copaiba oleoresin’s (referred to in the trade as
copaiba balsam) and copaiba oil’s ongoing use by cosmetic manufacturers is demonstrated by its inclusion on
product lists of eleven U.S. raw material suppliers to this industry (DCI, 1992). Although the term balsam is
often used with reference to copaiba, this oleoresin does not fit balsam’s true chemical definition because it
lacks the requisite benzoic or cinnamic acid or their esters (Leung, 1980).
The high concentration of sesquiterpene hydrocarbons in C. multijuga (Cascon and Gilbert, 2000) led
to the discovery that it could be used directly in diesel engines as a fuel (Record and Hess, 1943; Uphof, 1968;
Wang and Huffman, 1981; Calvin, 1983; Duke and Cellier, 1993). Experimental plantations of the tree were
started in the early 1980s near Manaus, Brazil with the idea of producing copaiba oil as an alternative energy
source to fossil fuels (Calvin, 1983; MMA/SCA et al., 1998). Even if these could not produce sufficient
supplies directly, Calvin (1983) speculated that sesquiterpene producing genes from copaiba might be
transferred to a suitable prolific Euphorbiaceous plant to maximize biosynthetic production of these chemicals.
As these plantations approached their 16th year, greater emphasis was being placed on the trees as a source of
oleoresin for medicinal and industrial products and wood for pulp and timber (MMA/SCA et al., 1998).
Trade statistics show that Brazil and Venezuela have been the major commercial producers and
exporters of copaiba oil. During the late 19th and entire 20th century, the U.S. was the largest importer.
France, Germany, United Kingdom and Japan are other important foreign markets (Duke, 1986; MMA/SCA et
al., 1998). Between 1883 and 1920, the U.S. bought at least 15 to 28 tons of the Maracaibo variety of copaiba
oil per year from Venezuela (Pittier, 1926; Furnemore, 1926; Dwyer, 1951). U.S. imports picked up shortly
before World War II and have fluctuated around 70 to 100 tons per year since (USDA, N.S.; Osol and Farrar,
1947; MMA/SCA et al., 1998). Measuring actual production in the region is difficult, but the Brazilian
Amazon exported about 120 tons to other parts of Brazil and foreign countries in 1994. Amazonas state is the
21
largest exporter accounting for 89 tons per year worth about $US 215,000. Rondônia and Amapa states also
exported some of the product (MMA/SCA et al., 1998). The Pará variety is the type of copaiba oil most
commonly traded in Brazil (Dwyer, 1951).
OLEORESIN HARVESTING PRACTICES AND YIELD
The first people to use copaiba may have taken advantage of oleoresin that occasionally exudes to the
outside of the tree, but all accounts of its use mention two main methods of harvesting the liquid. The first
technique involves chopping a large hole or wedge near the base of the tree with an axe. This action may
expose an oleoresin-filled cavity and creates a drainage reservoir into which the oleoresin ducts will eventually
drain their fluid (USDA, N.S.; Pittier, 1926; Furnemore, 1926; Howes, 1949; Poucher, 1950; Youngken, 1943;
Spruce, 1970). The technique may also kill the tree or at a minimum preclude any potential future harvest
(Pittier, 1926; SEBRAE, 1995).
The second, more recent method of harvesting copaiba is to drill a 1.5 to 5 cm diameter hole 0.6 to 1.0
meter from the ground into the heartwood center of the tree and collect the resin that drains out over successive
hours or days in a container. Once a tapping session is concluded, the hole is tightly plugged with a piece of
wood (Pittier, 1926; Calvin, 1983; Duke and Cellier, 1993; SEBRAE, 1995). Harvesters sometimes bore two
holes into the trunk. The first hole is drilled less than a meter above the ground; the second one is bored three
to seven meters higher - sometimes after the flow has stopped in the bottom hole (Osol and Ferrar, 1947; Mors
and Rizzini, 1966; Correa and Bernal, 1989; Duke, 1986; Duke and Cellier, 1993). A variation on the double
drilling technique involves breaking off a large branch from the tree (Plowden, in press). Fires are sometimes
made at the base of the trunk to hasten resin flow (Duke, 1986; Shanley et al., 1998). A pump was
occasionally used to accelerate oleoresin extraction in Venezuela (Pittier, 1926). Several accounts indicate
that some species are most often tapped in the dry season to maximize oil yield (Herrera, 1921; Le Cointe,
1947; Rodriguez, 1989; Duke and Cellier, 1993) while other species may yield more in the rainy season
(Dwyer, 1951; Alencar, 1982; SEBRAE, 1995; MMA/SCA et al., 1998). Recommended harvest frequency
varies from twice a year (Calvin, 1983) to once every two years (SEBRAE, 1995; MMA/SCA et al., 1998).
22
Different accounts portray widely varying oleoresin yields from copaiba trees. At one extreme are
reports that the pressure of the liquid can be so high that it sometimes bursts out of the trunk with a loud
explosion (Sayre, 1906; Pittier, 1926). Various authors state that harvesting one tree can produce 40 - 62 liters
of oleoresin at a time (Sayre, 1906; Pittier, 1926; Osol and Ferrar, 1947; Poucher, 1950; MMA/SCA et al.,
1998). Other reports indicate that average oleoresin yield per tree is 2.5 liters (SEBRAE, 1995; MMA/SCA et
al., 1998), 4.5 liters (Fernandes, 1949), or 20 - 24 liters (Calvin, 1983; Correa and Bernal, 1989; Duke and
Cellier). Even these so-called averages may be based on anecdotal accounts since they do not reference
specific studies. They are all much higher than the results of the one previous systematic study of copaiba
oleoresin production conducted near Manaus, Brazil (Alencar, 1982). In this study, 82 C. multijuga trees
yielded an average of 0.2 liters of oleoresin in the first of five successive tappings. While yields increased for
some trees during the second harvest, production dropped to an average 0.03 liters during the final tapping two
and a half years after the start of the experimental harvest. The largest amount that ever came out of a tree at
one time was 3.5 liters.
Several factors may explain the large discrepancy between the apparent abundant oleoresin in the
anecdotal accounts and the modest amounts harvested in the Alencar study. It’s likely that different species in
other areas produce more or less than the C. multijuga population near Manaus. Spruce (1970) notes that C.
martii trees near Santarem were relatively common but yielded so little oil they were not worth tapping. While
C. multijuga is considered a major oleoresin producer, other populations of this species or other species in
more favorable environments could generate more resin. Copaiba trees that were harvested with an axe
probably account for some of the highest reported yields; this method would predictably collect more oil than
boring one or two holes drilled near the ground. Another factor may be that earlier accounts are based on
second-hand stories of high yielding trees that underplay or ignore trees that yield little or no oleoresin.
Local lore indicates that successful harvest of copaiba oleoresin is far from guaranteed. In the eastern
Brazilian Amazon, forest peasants and Indians commonly say that looking up into the canopy before drilling or
chopping into a tree will cause the liquid to get sucked up to the top of the tree (Shanley et al., 1998; Plowden,
in press). The other stipulation is that a collector (presumably a man) should not have sexual relations with his
wife for several days before setting out to harvest copaiba. Folklore also suggests that oleoresin will not leave
23
the tree if a pregnant woman is present beneath it (Shanley et al., 1998). The central dilemma regarding
copaiba is that it has been assumed that trees regularly produce oleoresin in live tissues throughout their lives,
but there has been tremendously unexplained variability in the success people have had in harvesting this
product.
Due to the uncertainties about “average” oleoresin production, it is hard to estimate the number of
trees that have been harvested with an axe or a drill to produce more than 100 tons of copaiba oil per year in
the Brazilian Amazon alone. During the peak of the trade in the early 20th century, it seems likely that
thousands of copaiba trees were destructively harvested with an axe. Since copaiba has a low density that
varies from 0.1 to 2 trees per hectare (Ramirez and Arroyo, 1990; Sampaio, 1993; SEBRAE, 1995; MMA/SCA
et al., 1998; ter Steege and Zondervan, 2000) harvesting with the destructive harvesting method may have
taken a serious toll on copaiba populations (Pittier, 1926; Dodt (1939) cited in Sales, 1994). The vulnerability
of Copaifera to logging (Martini et al., 1993) reinforces the caution that will be needed to maintain current
populations if adult trees are killed through logging or destructive resin harvesting. Copaiba oleoresin is
frequently cited as a potential income generating non-timber forest product (NTFP) for forest peoples in the
region (Sampaio, 1993; Warner and Pontual, 1994; SEBRAE, 1995; MMA/SCA, 1998), so there is a strong
need to better understand the factors that affect its production and realistically assess harvest potential using
minimally destructive methods.
RESEARCH OBJECTIVES
In spite of the long history of copaiba oleoresin use as a medicinal product and cosmetic by
indigenous people and modern pharmaceutical companies throughout the Amazon region, there have been
almost no systematic efforts to quantify the yield of oleoresin from various types of Copaifera trees. There is
even less understanding of what factors related to the tree and its environment influence the amount oleoresin
that is produced and stored and how various harvesting techniques and other factors influence the amount that
is harvestable. The main goal of this study was to investigate the production ecology of a diverse group of
copaiba trees in the Alto Rio Guamá Indigenous Reserve in the eastern Brazilian Amazon and compare results
to the Alencar (1982) study and popular accounts. The specific objectives were to combine field observations
24
with low impact harvesting to learn how factors such as tree type, tree size, tree condition (fire damaged,
hollow or undamaged), internal trunk component dimensions (amount of sapwood and heartwood), bore hole
number, and drilling frequency relate to oleoresin harvest. Since oleoresin flow seemed to vary dramatically
between different parts of the tree (sapwood, sound heartwood, rotten or hollowed heartwood), special
attention was also paid to elaborating these differences as well to gain a better understanding of the timing and
location of oleoresin production and storage in the trees. A secondary objective was to evaluate the harvesting
efficiency and economics of copaiba oleoresin extraction with the drilling technique and recommend criteria
for optimizing its operation.
STUDY SITE
Between 1996 and 1998, a series of experimental harvests of copaiba trees were conducted in the Alto
Rio Guamá Indigenous Reserve located in eastern Pará state of the Brazilian Amazon. This was part of a
larger study to assess the production ecology of a five different commercial NTFPs in the reserve. The main
study site was an area of about 500 ha of closed tropical rainforest near the Tembé Indian village of Tekohaw
(020 37.7’ S; 460 33.1’ W) located on the Gurupi River. Underlying soils are on the border of two major soil
types classified by the Brazilian system as yellow latossols and red-yellow podzols - the latter called spodosols
in the American system (Projeto Radam, 1973; Kalpagé, 1974). The site has three principal forest types:
upland (“terra firme”) forest, lowland forest that is occasionally flooded (“baixo”), and seasonally flooded
forests near streams and the main river (“igapó”). Near this and other villages, patches of mostly terra firme
forest are cleared and burned to plant manioc and other crops (Sales, 1993 & 1994). This land use pattern
creates a mosaic of forest in various stages of current farm use, fallowed secondary forest and old forest. Other
important aspects of the landscape are swaths of secondary forest that resulted from unintentional forest fires
that burned through non-agricultural sections of the area in the early 1980s. The terra firme forest that hosts
copaiba is dominated by trees in the Lecythidaceae, Burseraceae, Leguminosae, Sapotaceae, and
Euphorbiaceae families (Balée, 1987).
In the early 20th century, harvesting copaiba oleoresin was one of the principal economic activities of
the Tembé in the region (CEDI, 1985; Sales, 1993 & 1994). Itinerant river traders (called “regatões”) were the
25
primary buyers who exchanged basic commodities with the Indians for copaiba oil and other forest products.
The terms of trade were so unfavorable to the Tembé, though, that extended families often moved in response
to the exhaustion of local copaiba resources (CEDI, 1985). This predatory harvest became so widespread that
one anthropologist who was concerned about the depauperate situation of the tribe in 1939 recommended that
the government use “energetic means to stop the complete destruction of the copaiba trees” (Dodt, 1939 cited
in Sales, 1994).
METHODS
In the summer of 1996, a four person team of Tembé elders searched the forest in the vicinity of
Tekohaw village with the author for any type of adult (>10 cm DBH) copaiba tree. Searches were conducted
primarily in patches of intact or mostly intact terra firme forest sometimes bordered by agricultural fields,
young secondary forest, or “baixo” (occasionally flooded) forest in June and July. Searches were concentrated
in terra firme areas where the Tembé believed they were most likely to find the widely dispersed trees.
Additional trees located in the area during a systematic inventory of part of the study area were later added to
the study.
When trees were located, their diameter was measured at two or more locations within 20 cm of
breast height. The measuring points were noted in relation to their distance directly below two nails put in
opposite sides of the trunk. This distance was measured on a pair of metal tape measures hung on each nail.
Common tree names were noted according to Tembé classification. Using sight and sound, it was determined
to the extent possible if the tree had a large hollow. Other observations included noting the presence of wound
scars on the bark that indicated past axe harvesting, recording the presence of burn wounds, and noting if the
tree was alive or dead or had any other circumstances that might affect the tree, such as damage from another
fallen tree. Tree positions were initially mapped using paces and compass direction; they were later mapped
with a GPS unit. Leaf samples were collected when possible to assist with identification. Each tree was
marked with a numbered aluminum tag.
The first round of the oleoresin harvest was conducted in the local dry season in July and August,
1996. Depending on tree size and condition, one to three holes were bored toward the center of the trunk with
26
a drill that was 19 mm in diameter and approximately 0.5 meter long. With the exception of two trees (around
28 cm DBH), no tree less than 34 cm DBH was drilled. If more than one hole was bored, subsequent holes
were bored about 60o to either side of the initial hole. The first hole was generally made about 1 meter from the
ground; other holes were started about 20 cm higher or lower than the first hole to avoid intersecting bore
holes. While some trees thought to be hollow were drilled in the first few weeks of the study, obviously
hollow trees were not subsequently drilled.
As the drill bore passed into the tree, the distance from the outside of the bark to the following points
was determined by inserting a stick into the bore hole when the drill bit was withdrawn and measuring the
length of the stick to that point: 1) the beginning of the heartwood (as noticed by the transition from light
colored sapwood to dark reddish wood, 2) the place where any oleoresin started flowing out, 3) the beginning
and end of any hollow or low density rotten wood, and if reached 4) the transition from heartwood to sapwood
on the opposite side of the tree. The angle of the drill with respect to a plane perpendicular to the trunk was
also recorded. The cosine of this angle was multiplied times the actual distance the drill bit traveled before
reaching these points to estimate the distances from the bark edge to that wood type on a plane perpendicular to
the trunk’s vertical orientation. If any oleoresin emerged, it was collected and measured in graduate cylinders
or beakers. If no oil came out within several minutes of drilling, a hardwood plug was banged into the drill
hole. Observations were also noted for any other liquids or sounds that accompanied the drilling. Some trees
in good condition (i.e. not rotten or hollow) that did not produce oil immediately after drilling had the plug
briefly removed within the next few days to collect any small amount of oil that may have accumulated.
The second round of harvesting took place in November, 1997, approximately 14 months after the
initial drilling. Tree diameters were remeasured and bore-hole plugs were removed to measure oleoresin flow.
No new drilling was conducted except to drill through plugs that broke off during attempts to pull them out.
New plugs were made if necessary to replace original rotted ones. Where oleoresin had leaked onto the trunk
in between harvesting events, this was noted. A third round of harvesting was conducted in May, 1998. In this
round, trees drilled in 1996 were again checked in the same manner as described for the second round. A few
trees not drilled in the first round and other trees subsequently found in the study area were drilled for the first
27
time with the same protocols. This round, the amount of time required to drill to different parts of the tree was
also recorded along with the amount of oleoresin that had accumulated at successive five minute intervals.
RESULTS
TREE NUMBERS, TYPES, SIZE, AND CONDITION
The initial search for copaiba trees in the study area in 1996 found 55 copaiba trees. Another 15 trees
were found in 1997 and 1998 in the course of doing work with other tree species in the area. A systematic
survey of 7 hectares of terra firme forests located 6 copaiba trees (i.e. 0.86 trees per ha). This estimated
density, however, may not represent the study area as a whole since only 70 trees were found in an intensively
traveled area of two to three hundred hectares.
The Tembé recognize four types of copaiba trees in the nearby forests. The most common and
smallest variety was the white copaiba tree (“copai ching”) that had an average DBH of 47.3 ± 2.5 cm (n=43).
It usually had a slightly tapering cylindrical trunk, smooth grey bark and whitish sapwood. Its oleoresin is pale
yellow and flows readily. The mid-size copaiba tree in the area is called “copai kuru;” this is best translated as
the knobby copaiba tree. These trees had an average DBH of 59.3 ± 4.0 cm (n=14). It also has a grayish
cylindrical trunk and light colored sapwood, but it is covered with round eruptions on the bark that are just
under 1 cm across and 0.5 cm tall. Its oleoresin is grayish and very viscous. The largest copaiba tree in the
area is the red copaiba tree (“copai pirang”) whose trunk flares out into a square base. Red copaiba trees at the
site had a mean DBH of 70.2 ± 5.5 cm (n=13). The tree’s bark is usually ridged and brownish red, and its
sapwood and medium viscosity oleoresin are also reddish-orange. A fourth type of copaiba tree called “copai
hu,” the “large copaiba” tree, is occasionally found close to main rivers. Its characteristics are a blend of those
described above. Since only one specimen of this type of tree was found several kilometers from the other
study trees in this very different habitat, it was not included in this study.
The common Tembé names for the copaiba trees in their area were applied consistently by different
Indians, but there were variations that made placing some trees into one of these three categories more
28
difficult. Some trees classed as white copaiba trees had a square base like the red copaiba and/or a few knobs
typically found on the knobby type. It is not known if the variants of these three types represent separate
varieties, separate species, or occasional hybrids between closely related species. Where some doubt existed
about how to classify a particular tree, the name designated by the elder during the initial survey has been used.
Attempts were made to gather flowers from study trees to aid in their identification, but since these trees
generally only reproduced once every couple of years, collecting the diminutive and short-lived flowers
remained an unfulfilled task. A botanist from the Museu Goeldi reported that leaf samples collected from
study trees came from at least three possible species of Copaifera, but it was not possible to match species
names with the Tembé common names.
A relatively high percentage (21.4%) of the copaiba trees observed in this study had been damaged by
fire. The percentage was greatest among the white copaiba trees that were most often found in areas subject to
the forest fires of the 1980s. Relatively few knobby and red copaiba trees were burned in this unintentional
fire, but several trees were burned or lost in the course of the study in forest areas converted to manioc
cultivation.
The other notable feature of the copaiba study trees was that 48% of them had some hollow or rotten
wood in their trunk. The percentage of trees with a hollow space increased significantly with tree size class (χ2
= 18.6; df=6; p=0.01) and was significantly higher in fire damaged trees than unburned trees (χ2 = 8.0; df=1;
p=0.01)(Figure 2.2). The proportion of tree types with a hollow was highest in red copaiba, but the percentage
of hollow trees was not significantly different between the types.
TRUNK DIMENSIONS
During preliminary copaiba drilling, Tembé elders stated that oleoresin was only obtained when the
heartwood (“miolo”) of the tree was exposed with a drill or an axe. Consequently this study recorded the
dimensions of sapwood, heartwood, oleoresin bearing zones and hollows in each tree and the amount of
oleoresin obtained. When drilling into a tree, the transition from white sapwood to dark dark heartwood was
also accompanied by an increased difficulty in drilling. The distance from the outer bark to the beginning of
29
heartwood represents the width of the bark plus sapwood. This distance had a mean of 14.2 ± 0.4 cm (Table
2.1; Figure 2.1) and was not significantly related to overall tree size (Table 2.2). Heartwood diameter was
estimated by subtracting two times the distance from the bark to the beginning of the heartwood from the total
diameter. The ratio of heartwood diameter to overall tree diameter (at drill height) and percentage heartwood
area to total cross-sectional area were both highly significant (Table 2.2). In the smallest size class (25 - 35 cm
DBH), heartwood area accounted for about 5% of total area (Figure 2.3). This percentage steadily increased
with tree size and exceeded 40% in trees over 75 cm DBH.
When trees yielded oil in the process of drilling, oil flow commenced when the drill had gone an
average of 18.7 cm into the trunk (Table 2.1, Figure 2.1). In most cases this point occurred when the drill had
passed 4 - 5 cm beyond the start of the heartwood; the distance from the estimated center of the tree also varied
widely and averaged 6.6 cm To efficiently capture escaping oil, the drill was withdrawn immediately. With
trees that had more than one hole drilled at a time, copious oleoresin flow sometimes emerged from one or
more holes (Table 2.3), so oleoresin rich sections were not apparently are all centered in the pith, regularly
shaped or connected to each other. Apart from always existing in the heartwood, the unpredictability of
oleoresin rich locations is reinforced by the lack of significant correlation between either its starting point, its
distance from the tree center or tree diameter (Table 2.2).
Drilling measurements showed that hollows also occurred primarily in heartwood (Figure 2.1), but
they may have sometimes occurred in sapwood as well (Table 2.1). When hollows were present, their volumes
were quite extensive since 11 out of 14 trees with hollows that were drilled more than once during a harvest
had hollows at more than one drilling location. Estimates of these hollow areas showed they occupied a mean
of 55% of the heartwood area and 15% of the entire trunk area. The amount of hollow area and ratio of hollow
area to total cross sectional area of the trunk were significantly correlated with tree size in trees that had
hollows (Table 2.2).
OLEORESIN YIELD WITH COPAIBA TREE DRILLING
When trees were drilled there were a number of possible outcomes once the drill bit made it past the
sapwood. In most cases, drilling produced no oleoresin at all (Table 2.3), and the drill frass was solid but dry.
30
In some cases, as drilling proceeded into the heartwood, the frass was highly scented and oily. Drilling further
then unleashed a steady outpouring of oleoresin. Most of the oleoresin drained from the presumed cavity came
out within the next hour, but depending on the size of the oleoresin rich section, the material continued to
slowly drip out for another day or more. A slightly larger percentage of trees yielded a very small amount of
resin (≤ 10 ml.) during one harvest, but this sometimes was not discovered until a day later.
The other common occurrence was hitting a tree with a hollow. There was only one case where even
a small amount of oleoresin was collected from a tree with a hollow even though intact heartwood was usually
encountered before hitting the hollow, and hollows were sometimes confined to a limited section of the tree.
The third condition found was rotten wood. Depending on its condition, these sections were either very easy to
drill through or had to be penetrated in stages as crunchy sections were encountered. The drill frass from
rotting sections was dry and sometimes powdery. Hollow and rotting sections did not yield oleoresin, but
termites often emerged from the bore hole of trees with these conditions. Several times, drilling yielded 0.5 to
1.3 liters of water tinged with tiny bits of oil. Its red color also indicated the presence of phenolics. A small
amount of water also emerged prior to the arrival of oleoresin in one knobby copaiba tree.
Another interesting phenomenon that sometimes accompanied drilling was a sucking sound that
seemed to be a vacuum attempting to pull material up into the tree. This only occurred after the drill had
reached the heartwood. This sound was associated with one each of the following events: before a minor
oleoresin flow; after a minor oleoresin flow, no oleoresin flow, and before the drill hit a hollow. One other
observation indicated that oleoresin flow in the lower part of the trunk may sometimes be connected with the
canopy. Shortly after oleoresin had begun to flow out of the first hole drilled in one tree, a rain storm began.
As the wind periodically got stronger and swayed the upper branches, the resin flow rate temporarily increased
as well. When the wind abated, so did the resin flow.
OLEORESIN HARVEST VARIATION BY TREE AND SITE VARIABLE
Oleoresin production among copaiba types showed that a higher percentage of white and red varieties
yielded oleoresin more often and in greater amounts than the knobby variety (Table 2.3), but comparing yield
in all drilled trees did not reveal significant differences. Looking just at trees that yielded some oleoresin
31
(Oleoresin Harvest Classes 2 &3) accentuates the poorer harvest of the knobby variety, but the preponderance
of modest yields in all trees still fails to generate significant differences between these types. The likelihood
that the white copaiba tree is a more prolific resin producer is indicated by the fact that its average total harvest
and average yield per harvest are higher than both the red and knobby varieties even though the average
number of times white trees were harvested and the average number of holes drilled per harvest were lower
than the other two types (Table 2.3).
The size of the copaiba tree has a stronger relationship to oleoresin yield than types examined in this
study. No tree smaller than 35 cm DBH yielded any oleoresin at all (Oleoresin Harvest Class 1), and no tree
smaller than 45 cm DBH yielded more than 50 ml. of oleoresin in any one harvest (Oleoresin Harvest Class
3)(Table 2.4). Oleoresin yield peaked sharply in the 55 - 65 cm DBH size class with a mean total harvest of
305 ml. per tree. This size class also accounted for 63% of the trees that produced >50 ml of oleoresin during
any drilling event. Trees greater than 75 cm DBH never yielded more than a trickle of oleoresin flow (Figure
2.4). A One-way AOV comparison of the mean total amounts of oleoresin harvested between size classes of
all trees that were drilled had a p-value of 0.071 (F=2.17). The harvest peak in the 55 – 65 cm DBH size class
can not be attributed to variation in the order of harvests or number of holes drilled per harvest since these had
no significant differences between the size classes (Table 2.4). The dominance of oleoresin harvest by trees
that are 45 - 75 cm DBH is reinforced when examining only trees that yielded some oleoresin (Oleoresin
Harvest Classes 2 & 3). The 55 - 65 cm DBH size class again surpasses the others with an average total yield
of 566 ml. per tree. Probably due to the small number of oleoresin trees in this sample, however, the
differences comparing size class to the production variables are not statistically significant.
Copaiba trees that were hollow yielded very little or no resin when they were drilled. Only 5 out of
27 (18.5%) trees drilled that had even a partial hollow yielded a small amount of oleoresin. The largest amount
that came out of any hollow tree at one time was 10 ml. When the distribution of the three oleoresin harvest
classes were compared between hollow and non-hollow trees, there was a significant difference (χ2 = 8.5;
df=2; p=0.014). The means of the total amount of resin produced by hollow and non-hollow trees during all
harvests was significant at the 94% confidence level (AOV: df=56; F=3.67; p=.061) while differences in mean
amount of oleoresin per harvest were not significant. Tree size was again shown to be important when resin
32
yields were examined for hollow and non-hollow trees by size class. For the three size classes that include
trees between 45 and 75 cm DBH, 53% of the non-hollow trees yielded at least 50 ml. in one harvest and
another 20% yielded at least a little oleoresin at some point.
OLEORESIN PRODUCTION VARIATION WITH THE ORDER AND TIMING OF HARVEST
Oleoresin yield sometimes varied according to the order of harvest. The average oleoresin yield for
the first harvest was much greater than the second amount that was slightly larger than the third harvest (Table
2.5). These differences, however, were not significant comparing all drilled trees or all trees that yielded some
oil (Oleoresin Harvest Classes 2 &3). The first harvest was significantly greater than the second harvest when
comparing just those trees (Oleoresin Harvest Class 3) that yielded more than 50 ml. of oleoresin in any one
harvest (Mann-Whitney Test; p=.03). This difference could not be attributed to tree size since both groups had
a very similar range. The difference did not carry over from the second to the third harvest when average
amounts were both very low. The change from the first to second harvest was almost but not always a
decrease. In 4 out of 18 oleoresin producing trees that were harvested more than once, the second harvest
yielded more than the first. Only one of these four trees, however, yielded more than a very modest increase.
The average amount of oleoresin from the first harvest of trees initially harvested in the dry season
(July and August) of 1996 was smaller than those only harvested in the rainy season (May) of 1998. Since this
difference was not significant, no effect of seasonal or annual differences on oleoresin production due to
climatic differences can be asserted.
OLEORESIN LEAKAGE
These descriptions of oleoresin harvest may have implied that oleoresin production and storage in
these copaiba trees was equivalent to the amount of oleoresin harvested, but it is important to report a few
observations that showed that the amount of oleoresin produced, stored and accessible for harvest may be quite
different. While oleoresin must be produced in living tree tissues, the drilling study indicated that all major
oleoresin flows came from accumulations in the heartwood. The origin of the minor flows that occurred hours,
33
days or months after the initial drilling could not be identified. The leakage of oleoresin onto the trunk from
several trees occurred from places other than bore holes, however, this indicated that at least some of this
material produced in the sapwood can be released to the outside. First, oleoresin was observed leaking from
nail wounds in three trees. Of the two trees that were drilled from this group, one yielded some resin during a
harvest, and the other did not. Oleoresin stains were also found around burn wounds of two trees that did not
yield any resin when harvested. Another tree was found with copious oleoresin leakage on one side of its
buttress root. It was the only case where such leakage was not associated with a visible wound. This large tree
had a partial hollow near the side of the root, but unlike most such trees, it did yield a small amount of
oleoresin on the opposite side of the tree with heartwood that was intact as far as the drill could reach.
HARVESTING EFFICIENCY AND ECONOMICS
In the summer of 1996, ten days were devoted almost exclusively to searching for copaiba trees.
During this time, three to four men collectively found one to nine trees each day with an average of five trees
per day. The lowest numbers occurred during days when forest that was occasionally flooded (“baixo”) was
crossed in transit between separate areas of terra firme forest. The overall number showed that copaiba tree
density was low even in its highest local concentration. The average number of trees found per day could have
been considerably higher if 20 - 30 minutes had not been taken at each tree to measure various tree dimensions
and site characteristics. Known trees close to Tekohaw were located quickly. Others were found in patches of
terra firme forests that were up to 3.5 km from the village. Actual travel distances to the farthest trees were
much longer since there were no straight line trails to those areas of the forest. When the study moved into the
harvest phase, one to five trees were drilled each day giving an average of 3.3 trees drilled per day. This
number was lower than would be possible doing drilling only because an average of 2.6 other trees per day
were found and measured.
Time keeping of some harvesting operations showed that it usually took almost five minutes to drill
through the bark and sapwood to reach the heartwood (Table 2.6). Another 20 minutes on average was spent
drilling through the heartwood. In spite of its apparent greater hardness, the drilling rate going through
heartwood was significantly faster than drilling through sapwood. The time needed to reach a hollow was less
34
than the time needed to reach the oil rich zone in those trees where oleoresin flowed out at the time of the
drilling. This matches the shorter average distance from the outside of a tree to a hollow than to an oil rich
zone (Table 2.1).
When a good oleoresin flow commenced, it took a considerable amount of time to collect it. There
were two principal patterns of oleoresin flow rate out of bore holes (Figure 2.5). In trees that yielded the most
resin (>350 ml from one hole), the majority of resin initially come out at a fast steady rate. Once about 93% of
the oleoresin rich zone had apparently been drained, the flow rate sharply dropped off to less than 4 ml. per
minute. This point was reached within half an hour in holes that ultimately yielded less than one liter, but it
took an hour and a half to reach for one hole that produced 1225 ml. With trees that produced smaller
amounts of resin, oleoresin slowly trickled out until it stopped.
A waterproof collection system was set up in one of the first trees tapped to capture oleoresin flow for
several days. Considering the rapid drop off in flow rate within several hours even with the most copious
flows, this practice was not continued since it was time consuming to construct and make repeated visits to
trees that were a considerable distance from the village. This strategy for collecting oleoresin that came out
quickly after drilling was necessary for research efficiency, but it may not be the best one for optimizing
commercial harvest. In sum, it took about 25 minutes to drill one hole to and through the heartwood of an
average tree with intact heartwood. It took an average of 16 minutes to discover if a tree was hollow by
drilling if the tree was not so hollow that banging on the trunk revealed its condition. Combined drilling and
waiting time for oleoresin producing trees varied from half an hour to three hours to capture more than 90% of
the material from an oleoresin yielding hole.
ESTIMATES OF COPAIBA OLEORESIN HARVESTING BY AREA AND TIME – IMPLICATIONS
FOR HARVESTERS
The efficiency and economic viability of harvesting oleoresin from a wild population of copaiba trees
depends on a number of factors. The first is population density (Peters, 1994; Warner and Pontual, 1994).
Since this may range from 0.1 to 2 trees per ha for copaiba trees (MMA/SCA et al., 1998) at a small scale, the
number of copaiba trees in an area of forest will have a direct effect on both the size of the potential resource
35
and the efficiency with which that resource may be harvested. In order to predict oleoresin yields and gross
revenue on a per tree and per area basis under different conditions, a simple model was constructed using a
range of minimum, medium, and maximum values from this and other studies for density, percentage of
oleoresin yielding trees in the population, resin yield per tree, the number of times a tree may be harvested
during its productive period and the time horizon that is used to assess the output estimates of production and
value (Table 2.8).
Density estimates in the model use lower and upper limits for Copaifera trees that ter Steege and
Zondervan (2000) found in Guyana since these represent one of the best data sets from relatively large (100 ha)
plot inventories of diverse forest types in the Amazon region. The medium value of 0.3 trees/ha is close to
both the mean density of copaiba trees from these plots and the 0.27 trees/ha density figure for the genus in the
Brazilian state of Acre (SEBRAE, 1995). The second key factor in assessing harvest potential and designing
an optimal harvesting strategy is to estimate the ratio of productive (or at least potentially productive) trees to
the total number of trees by determining which size classes (if any) yielded the largest amount of product
(Peters, 1994). Both the Tekohaw and Manaus copaiba studies showed that only trees in the middle to upper
middle size classes of their respective populations were usually worth harvesting (Table 2.4; Alencar, 1982) so
the model calculates yields assuming that 15 – 50% of the trees in a population will be in these harvestable size
classes.
The two time horizons used for the harvesting by area model are 40 years and 6 years. The longer
period corresponds to the estimated amount of a time a tree would have in its prime oleoresin yielding years
around Tekohaw or a similar forest. If the diameter growth rate of the trees is 0.5 cm per year, it would take
about 40 years for a tree to grow from 45 cm DBH when substantial oleoresin yields first seem possible to 65
cm DBH when the probability of losing oleoresin production and/or storage potential is very high. Amounts of
potential resin yield and value are, therefore, based on exploiting the full capacity of a tree during its oleoresin
yielding life and estimate harvests under assumptions of minimal harvest potential (one or two harvests) and
annual harvests for the entire period. The six year time-line presents a shorter time-span of predicted resin
yield where the medium set of criteria allowed for a second and final harvest of productive and the maximum
set of criteria allowed for three harvests that occurred on alternate years during the same period.
36
The model shows that under minimum values for overall copaiba tree density, percentage of
productive trees in the population, amount of resin harvested per tree, and number of harvests possible per tree,
the average yield by area would be less than 1 ml oleoresin per ha per year in both the 40 and 6 year time
horizon with a value less than $US 0.01 per ha per year. For a 1000 ha area of intact forest (comparable to the
area of forest within easy access of a Tembé village), these equivalent values are about $US 1.58 per year with
the 40 year horizon and $US 10.50 per year with the 6 year horizon. The per year values are higher in the
shorter time horizon because all of the benefits are reaped in the first and only year the trees are harvested.
Medium estimates for copaiba tree and harvest factors that either match or are slightly better than conditions
found at Tekohaw generate oleoresin harvest rates of 6.8 liters per year in the 40 year projection and 45 liters
per year in the six year projection from a 1000 ha area. The oleoresin sales value would range from $US 0.07
to $US 2.94 per ha per year with the two time horizons. Maximum estimates are derived from a population
with moderate density and the highest proportion of productive trees that biannually yield 2.5 liters of oleoresin
per harvest (an amount cited in various popular accounts as the “average” yield of oleoresin per tree). Under
these optimistic circumstances, the rate of oleoresin harvest would be 0.9 liters per ha per year worth $US 6.56
per ha per year under both time horizons.
Indians or other forest dwellers traversing the woods to hunt or collect other forest products could
easily harvest copaiba trees on an opportunistic basis for many years without approaching the limit of the
resource. This mode, however, does not represent the potential upper range of income if the intent was to
maximize copaiba oleoresin harvesting for commercial purposes. This model, therefore, assumes that the
operation would be divided into a search phase and a harvest phase. In the search phase, a systematic effort
would be made to locate all of the adult copaiba trees in a particular area of forest. For ease of calculation and
illustrative purposes, this model uses 1000 ha as a base area that would generally be accessible within an
hour’s walk from the center of a circular area. During the initial phase, trees would be quickly measured and
marked, but little other time would be spent with each tree other than noting if the tree was fire damaged or
appeared to be hollow. In the harvest phase, only those trees deemed likely to yield appreciable amounts of
oleoresin based on their size and condition would be drilled. Beyond wasting effort, another reason to avoid
drilling small trees is that doing so may interfere with normal heartwood development (Hillis, 1987) and
37
increase the chance of early pathogen attack. Each tree would be drilled twice unless the first bore hole
showed the tree was hollow. Estimates assume one person does all activities. Drill times could be reduced if
two people drilled at the same time, but potential revenues per person would then be reduced.
The second copaiba oleoresin harvesting model (Table 2.9) attempts to estimate the amount of
oleoresin that could be harvested and the amount of income earned from selling that product per time invested
depending on minimum and maximum assumptions about tree density, resin harvest per tree depending on two
different harvesting strategies that are often employed. In both methods, harvesters search for trees and drill
any tree two times that he believes will likely yield oleoresin. In the first harvesting strategy (Method A), a
harvester collects any oleoresin that comes out of each hole until the flow has almost or completely stopped.
He then plugs the holes and moves on to look for another tree. The model assumes this procedure would be
followed every day for six days per week. In the alternate strategy (Method B), the harvester prepares a device
to catch oleoresin that avoids collecting rainwater and debris. After drilling trees and determining that resin
flow will continue for awhile, he leaves the collection device in place and moves on to drill the next tree. The
harvester looks for and drills any productive trees he finds for five days. On the sixth day, all of the trees
drilled during the previous two days are revisited to collect accumulated oleoresin and plug the trees. This
method makes for more efficient use of time on drilling days, but it requires extra time to return to trees to
complete the harvesting.
This model shows that a harvester using Method A would gather 0.13 to 2.36 liters per day under the
minimum to maximum conditions and 0.12 to 8.15 liters per day using Method B (Table 2.9). These would
translate into $US 0.90 to $16.53 per day with Method A and $US 0.85 to $57.03 per day with Method B.
Under minimum conditions, all of the potential productive copaiba trees in a 1000 ha area could be drilled one
time in 70 to 74 days. With higher density and yield values, it would take 230 to 794 days to complete an
initial round of harvesting.
38
DISCUSSION
TREE NUMBERS, TYPES, SIZE, AND TAXONOMY
The low density of copaiba trees found in the Tekohaw study area is typical for the genus in other
Amazonian forests. Calculating precise densities of uncommon trees, however, from inventories based on
plots that are 1 ha or smaller is not very precise. Ethnobotanical surveys of seven 1 ha plots near the
Tambopata Reserve in Peru found one copaiba tree (Phillips et al., 1994). Balée (1987 & 1994) did similar
detailed surveys in one ha forest plots near a Tembé village not far from Tekohaw and seven other areas in the
adjoining Alto Turiaçu Indigenous Reserve in Maranhão state and did not find a single copaiba tree. A census
of C. pubiflora trees in 277 hectares of forest in Venezuela showed a density of 1.04 trees/ha (Ramirez and
Arroyo, 1990). Broader scale inventories of forests in Guyana showed that all species of Copaifera combined
had between 1 and 174 trees per 100 ha in areas where they were found at all, resulting in an average density of
0.36 trees/ha (ter Steege and Zondervan, 2000). Although he did not conduct a systematic inventory, Alencar
(1982) found 82 C. multijuga trees in an extensive search of about 200 ha giving a rough density of 0.41
trees/ha. While the inventory of 7 ha at the Tekohaw site indicated copaiba density was 0.86 trees/ha, the
figure for the area as a whole is probably smaller since an intensive search of several hundred hectares of terra
firme forest only found 70 copaiba trees.
One notable feature of the copaiba populations in the Tekohaw area was that the mid-range size
classes (46% in the 40 – 60 cm DBH size classes) were better represented than the smaller ones (24% in the 20
– 40 cm DBH size classes). Alencar (1982) found a similar size class distribution among C. multijuga trees
near Manaus. This lack of trees in the smallest size classes could either mean that younger trees were not
found as readily as older ones that were easier to identify and/or that the populations were in phases of
relatively poor recruitment of younger trees. This pattern has been observed in demographic studies of other
tropical trees (Clark and Clark, 1992) and does not necessarily reflect a lack of propagules. Field observations
of fruiting copaiba trees near Tekohaw and Manaus (Alencar, 1981 &1984) show that trees can produce
thousands of fruits at a time and that the seeds readily germinate. Like most tropical trees, however, both and
39
seedlings are heavily preyed on by insects and mammals (Alencar, 1984; Ramirez and Arroyo, 1987a, 1987b),
and those that survive have special requirements for light conditions to compete with other seedlings (Alencar,
1984; Mostacedo et al., 1998; Ramirez and Arroyo, 1990; Fredericksen et al., 2000). It may just be that the
conditions that most favor copaiba establishment occur at long intervals.
For young trees that become established, growth is still slow. The average copaiba tree diameter
growth rate at the Tekohaw site was estimated to be 0.65 cm per year. This is greater than the average growth
increment of 0.26 cm per year carefully measured in a C. pubiflora population in a drier forest region of
Venezuela (Ramirez and Arroyo, 1990). These growth rates are consistent with the finding that medium age
trees (40-70 cm DBH) in the Amazon have lived an average age of 200 years or more (Vetter and Botosso,
1989). This could mean that the trees in the demographic peak of the copaiba populations around Tekohaw
were established during conditions that existed more than a century ago.
The problem of identifying Copaifera trees to species in areas of high diversity is a problem both in
the field and herbarium. The genus was last described forty years ago by Dwyer (1951) who commented that
leaf characteristics alone could not be used to distinguish the species. This finding was echoed by legume
taxonomists at both the New York Botanical Garden and the Museu Goeldi in Belém who could not identify
the copaiba species from the Tekohaw study area based on leaf samples, although C. reticulata, C. langsdorffii,
C. martii, C. guianensis, and C. duckei were mentioned as possibilities (Sergio Antonio da Silva, personal
communication 1998). An analysis of oleoresin samples from several of the study trees, however, did not
match the chemical profiles of previously studied species (Vera Cascon, personal communication 1999). For
the third time in twenty years, a Brazilian systematist is now taking up the challenge of revising Copaifera
taxonomy in the region (Regina Celia, personal communication 2000). Fertile specimens would no doubt help
to resolve this dilemma, but since flowering occurs on average once every two to four years (Alencar, 1984 &
1988; Fredericksen et al., 2000; Table 4), the field challenges of collecting these materials and resolving the
genus’ taxonomy will require an extremely dedicated effort.
40
TREE CONDITION AND SITE PREFERENCE
The higher percentage of burned white copaiba trees may be connected to site preference since they
were located primarily in areas the Tembé described as “areia” (sandy) while the red copaiba trees were found
more often in areas described as “barro” (clay) soil. Soil texture measurements of samples from both types,
however, showed that each had high percentages of both components (Plowden, unpublished data). Whether
or not copaiba type specific site preferences exist or explain differences in their exposure to the forest fire of
the 1980s, the expansion of agriculture around Tekohaw led to fire damage and loss of several copaiba study
trees and forest habitat. Copaiba does have some resiliency dealing with burn damage, however, since new
sprouts were often seen coming out of the lower trunk of trees that had suffered such damage.
The linkage between tree size and hollowness found in this study has been found in other groups of
both temperate and tropical trees (Fearnside, 1992; Lindenmayer et al., 1993, 2000; Ball et al., 1999;
Greenberg and Simons, 1999). The occurrence of hollowness also varies by species (Fearnside, 1992;
Lindenmayer et al., 1993; Greenberg and Simons, 1999). In a study of a mixed group of trees near Manaus,
27% of trees greater than 40 cm DBH were hollow (Fearnside, 1992). This is less than the 56% comparable
figure for copaiba trees in the Tekohaw study area, but these trees are apparently more resistant than another
tropical legume angelim pedra (Dinizia exelsa Ducke) that is almost always hollow (Fearnside, 1992).
The high proportion of hollowness in red copaiba trees (69%) is apparently related to the large
number of its trees in the largest size classes (69% were >60 cm DBH). The relatively high proportion of
hollow white copaiba trees (38%) likely reflects the relatively high proportion of trees damaged by fire (28%).
In a forest area unaffected by intense fire, the proportion of hollow trees in the smaller size classes would
probably be much less than found in this study area. Beside fire, other hollow trees in the study area had been
hit by falling trees or were growing with their trunk abutted against another large tree. Many hollow trees had
large colonies of termites living in or near them. Other studies have verified that fire, wind induced damage,
successional series of fungal pathogens, termites and other insect attacks can all be agents of heartwood rot and
hollow formation (Boddy and Rayner, 1981; Lindenmayer et al., 1993, 1997, & 2000; Mattheck et al., 1994;
Greenberg and Simons, 1999). Once tree rot begins, termites are efficient at excavating decayed wood
(Lindenmayer et al., 1993).
41
The only hollow trees found in the smallest size class (20 - 40 cm DBH) of copaiba trees at Tekohaw
were all fire damaged, so the onset of hollowness in undamaged trees normally seems to commence in larger
trees. It is interesting to note that among the three trees that had apparently been axe harvested in the past (as
indicated by a circular area of scar tissue near the base of the trunk that was about 45 cm in diameter), only one
of them was hollow. This tree had also been severely fire damaged. These observations showed that axe
harvesting does not necessarily lead to early hollowness or mortality.
TRUNK DIMENSIONS
The importance of the relative size of copaiba’s sapwood, heartwood, and hollow sections of a trunk
is that they determine the spatial limits for the tree to produce and store oleoresin. Many trees that make
copious resin in the cambium quickly release stored material and may make additional amounts to protect
living tissues from physical injury or attacks from other organisms (Langenheim, 1973; Phillips and Croteau,
2000). Some trees make phenolic compounds in the tree’s internal zone where senescing sapwood is
undergoing its transformation to dead wood. There are a few other trees that store resins in their heartwood,
but the site of their synthesis and mechanisms for transferring them to this inner zone are not know (Hillis,
1987; Langenheim, personal communication 2001). The mechanism that links production of oleoresin by
Copaifera parenchyma cells, their deposit in secretory canals, and their accumulation in the heartwood of trees
at this study site is not understood. There are other types of Copaifera that apparently yield oleoresin without
needing to drill as far into the trunk as the heartwood (Langenheim, personal communication 2001). While
oleoresin deposits in the heartwood may stave off microbial attacks, the existence of empty hollows in
damaged or older copaiba trees seems to signal an end to the tree’s ability to store substantial amounts of resin.
The finding that copaiba sapwood width is not correlated to tree size once a tree reaches a certain
minimum size can be combined with tree growth rate to estimate the age at which a tree may begin to
accumulate harvestable quantities of resin. If copaiba diameter increases at an average rate of 0.5 cm per year,
the average sapwood width that is 14 cm represents 56 years worth of growth. Since heartwood formation was
not observed in the copaiba trees less than 30 cm DBH, the initial point at which resin may begin to
accumulate in the tree would take at least this long and probably longer if growth is slower in the younger
42
trees. There are wide variations between trees, but this size for the initiation of heartwood is typical of many
(Hillis, 1987).
Once heartwood formation begins, the close relationship found between heartwood size and overall
tree size in these copaiba trees (Table 2.2; Figure 2.3) is common to many other species (Hillis, 1987). While
overall tree diameter is very responsive to environmental conditions, the extent of heartwood formation is less
affected by these influences and is more determined by genetic factors (Fries, 1998; Ericsson and Fries, 1999).
The conversion of sapwood with live cells and starch reserves to heartwood composed of fully lignified nonliving tissues is a natural process in the life of a tree (Hillis, 1987). It involves the active translocation of
nitrogen and other mineral nutrients from dying sapwood and its infusion with a range of extractives that
include tannins, other phenolics and sometimes terpenoids (Andrews et al. 1999; Bergstrom et al., 1999). The
factors that trigger these anatomical, biochemical and physiological changes in the tree’s interior are still
largely unknown, but one benefit of the process is that they protect the heartwood for many years from
microbial and insect attacks (Bergstrom et al., 1999; Fujita et al., 1999). Keeping heartwood intact is
advantageous because trees that surpass a certain threshold of hollowness become vulnerable to breakage
(Mattheck et al., 1994; Lindenmayer et al., 1997). The red color of copaiba’s heartwood indicates it contains
substantial phenolics (Hillis, 1987), and the presence of oleoresin in the tree’s interior presumably acts as an
additional defense against decay causing organisms.
The key observation about the origin of substantial oleoresin flows is that it was usually well inside
the outer limits of the heartwood. This indicates that whatever conditions created these oleoresin rich areas
occurred close to the pith and did not exist until a tree had grown well beyond initial heartwood formation.
Since major oleoresin flows were not obtained in trees less than 50 cm DBH (Table 2.4), such trees were
probably established for at least 100 years.
Like oil rich zones, tree hollows in these copaiba trees were generally located in the heartwood zone.
As found with other trees, the size of these cavities corresponded well to overall tree size (Fearnside, 1992;
Greenberg and Simons, 1999; Lindenmayer et al., 1999), although the extent to which a hollow consumed a
tree’s inner volume varied considerably. Estimates of hollow area size in this study’s copaiba trees (Table 2.1;
Figure 2.3) showed that when they were present, hollows consumed an average of 15% of the total tree cross
43
sectional area and 55% of the heartwood. These are likely underestimates of the true extent of hollowness in
all trees, however, since these data were based only on trees that were drilled. A number of other large
obviously hollow trees (based on the “hollow” sound produced when banging on the tree) that were not drilled
probably contained a higher percentage of hollow space than the areas found in the harvested trees which only
revealed their partial hollows upon drilling. These results are comparable to two studies in the Amazon which
showed that 20 - 30% of the stem volume of hollow trees was filled with air or light debris from termite
activity (Fearnside, 1992). Greenberg and Simons (1999) found that hollows first appeared in turkey oaks
(Quercus laevis Walt.) and sand post oaks (Q. margaretta Ashe) an average of 69 and 106 years after
establishment. Rot was found in these respective trees with as few as 35 and 27 growth rings. The average
size or age at which hollows began in this study’s copaiba trees could not be calculated, but the smallest size at
which a fire damaged tree had a hollow was found was 31.8 cm DBH, while the smallest non-fire damaged tree
was 43.2 cm DBH. These trees were at least 60 - 80 years old if it is assumed they grew at an average
diameter growth rate of 0.5 cm per year.
COMPARISON OF OLEORESIN HARVEST IN TEKOHAW COPAIBA TREES TO C. MULTIJUGA
NEAR MANAUS
This study of several species of copaiba trees near in the eastern Brazilian Amazon had similar results
in some aspects to those found by Alencar (1982) who studied oleoresin harvest of 82 C. multijuga trees in the
central Brazilian Amazon near Manaus. The most important common finding was that the mean first harvest
was greater than subsequent harvests (Table 2.5; Table 2.7). At Manaus the percentage of trees that yielded
>25 ml. per harvest significantly declined from the first to the fifth harvest (Pearson’s Correlation. Coefficient.
= -0.926; p=.024). As found at Tekohaw, the second harvest of several trees was larger than the first, but this
effect was not repeated in subsequent harvests where the third, fourth, and fifth harvests from these trees
spread about a year apart almost all went to zero. These results suggest the following scenario. The strong
initial harvest resulted from draining oleoresin that was produced over many years and had accumulated in
cavities in the heartwood. Once this material was removed, these cavities did not fill up again. The second
round of harvesting from the same holes sometimes yielded a small amount of resin by draining off material
44
that had dripped into the bore hole from active resin canals in live tissue. This fluid may have built up as a
result of normally produced material coming from severed resin ducts. It may also have been the result of
enhanced oleoresin production stimulated by the drill wound. The drop off by the third and subsequent
harvests might have occurred because woundwood had formed around the drill wound. This type of discolored
wood whose physical and chemical properties resemble heartwood (Hillis, 1987; Shigo, 1994) could have
sealed off the drill site from active resin ducts in the area. An examination of drilled trees could verify if
copaiba sapwood and heartwood react similarly to injuries seen in many other trees. The drop off in third and
later harvests might also have occurred if any oleoresin production response to the initial wound was only
short-lived.
This phenomenon of rapid harvest drop-off is apparently not shared by all Copaifera species. Apart
from many popular accounts that refer to multiple harvests without details, Cascon and Gilbert (2000) report
that they removed 300 - 550 ml. of oleoresin from the same holes in ten consecutive tappings at one to four
month intervals from the same C. duckei tree in Amapá state, Brazil. While they did not attempt to drain the
tree during these harvesting events, the large changes in the ratio of the main oleoresin volatile components
during the harvests showed that the material collected in later harvests was not the result of a large previous
accumulation.
The second important common finding between the Manaus and Tekohaw copaiba studies was that
the mid-size copaiba trees were the best oleoresin producers. Alencar (1982) found a positive but nonsignificant correlation between tree size and resin harvest, but young trees and very large trees (59 - 78 cm
DBH) yielded little if any oleoresin. It, therefore, seems that the major oleoresin producers in this population
were between 40 and 60 cm DBH. This reinforces the idea that the conditions for maximizing oleoresin
harvest are somehow related to tree age and condition. Oleoresin tappers of Dipterocarpus alatus Roxb. ex G.
Don in Laos also found that yields peaked in mid-size classes because young trees did not produce much, and
older trees were more likely to be hollow (Ankarfjärd and Kegl, 1998). Oleoresin harvest also peaked in the
40-50 and 50-60 cm DBH size classes of Benguet pine (Pinus kesiya Royle ex Gordon) in the Philippines, but
in this case the drop off in the largest 60-70 cm DBH size class was not due to hollowness (Orallo and
Veracion, 1984). This is an important finding because it shows that trees past a certain age may actually lose
45
oleoresin production capacity. Rather than viewing hollowness as the cause for the loss of oleoresin
production capacity, it may be that the loss of oleoresin production capacity and its defensive properties is a
natural result of aging that leads to the increasing likelihood of hollowness in older copaiba trees.
If first harvests primarily tap oleoresin that has accumulated in heartwood cavities over many years,
there should be no seasonal effect on its yield. On the other hand, seasonal differences in yield seem quite
possible if harvesting primarily drains oleoresin recently deposited in canals from living tissues. C. multijuga
yields were higher in harvests conducted in the rainy season than the dry season, although comparison of these
group means were not statistically significant (Table 2.5) and harvest order could have been responsible for the
higher rainy season yields observed. My study found that first harvest oleoresin yields were higher in a group
of trees drilled in the late rainy season in 1998 than ones drilled in the middle of the 1996 dry season. These
differences, however, were not statistically significant. Dwyer (1951) reported that oleoresin harvests from C.
venezuelana and C. pubiflora in Venezuela also peaked during the rainy season between December and April.
In contrast to these accounts of greater oleoresin harvest in the rainy season are reports that other
copaiba species yield (and possibly produce) most oleoresin in the dry season. Peak harvesting of C.
officinalis in Venezuela and Peru occurs in the summer months between July and November (Herrera, 1921;
Duke and Cellier, 1993) while C. reticulata trees are most often tapped from August to October (Le Cointe,
1947). Resin production has been shown to be higher in summer months in several pine species perhaps
because lower water availability in this season allows a tree to devote more photosynthate to production of
defensive chemicals than growth (Lorio and Sommers, 1986; Blanche et al., 1992). In copaiba’s case, peak
flowering and fruit production occur during the rainy season (Alencar, 1988) so this might introduce
competition for carbon resources during this period. Given the uncertainty about a seasonal effect on oleoresin
harvest, the generic recommendation by some Brazilian agencies to harvest copaiba trees in the rainy season
(SEBRAE, 1995; MMA/SCA et al., 1998) seems premature.
There are a few notable differences in the results and conclusions between the Manaus and Tekohaw
studies. The first is that the maximum and average oleoresin yield for first time harvest from the C. multijuga
trees (3500 ml. max; 212.7 ml. ave.)(Table 2.7) were larger than obtained from the Tekohaw copaiba trees
(2028 ml. max; 72.5 ml. ave.)(Table 2.5). The most obvious possible difference is that different species of
46
Copaifera were being harvested at the two sites, but there is not enough information available yet to make such
a comparison. The difference in the size class distribution between the two study areas may account for some
of the greater production at Manaus. At that site, 60% of the copaiba trees were in the optimal oleoresin
yielding size range while only 49% of the Tekohaw trees were in the 45 - 75 cm DBH range that included their
maximum oleoresin yielding trees. Tekohaw copaiba trees that were drilled contained both a higher
percentage of small trees that were presumably too young to accumulate much resin and a higher percentage of
hollow trees that had lost their capacity to store the material.
One other difference that may be related to a difference in the harvest at the two study areas is the
content and viscosity of the oleoresin. At the Tekohaw site the knobby and red copaiba trees both have fairly
viscous oleoresins. The white copaiba trees resin was less viscous than the other two, but even it seemed less
fluid than samples of “copaiba preta” (presumably C. multijuga) oil observed for sale in the Belém market. As
mentioned earlier, there is a wide variation in the percentage of essential oils in copaiba oleoresin at both the
intra and inter-specific levels (Sayre, 1906; Kraemer, 1907; Furnemore, 1926; Ossol and Farrar, 1947; Leung,
1980). While no research has examined a potential link between copaiba oleoresin composition and
productivity, there is a known inverse relationship between the ratio of volatile to non-volatile chemicals and
an oleoresin’s fluidity and viscosity (Langenheim, in press). It has been observed in other oleoresin producing
trees that the volatile and other light portions of the material (principally monoterpenes and sesquiterpenes)
give it the ability to flow through resin canals and deliver the heavier and potentially more toxic defense
compounds (mostly diterpene resin acids) to the site of injury or attack (Langenheim, 1990; Phillips and
Croteau, 1999). Since heavier resins flow more slowly than lighter ones, it is understandable why some
harvesters have set a fire at the base of some copaiba (Duke, 1986; Shanley et al., 1998) and some Dipterocarp
trees in Asia (Ankarfjärd and Kegl, 1998) to increase the flow rate of some heavy resin producing species.
Many types of terpenoids can be energetically expensive for a tree to produce (Gershenzon, 1994), and the
diterpenes found in copaiba might be costlier to make than the simpler sesquiterpenes. If this is true, a tree
might be able to produce a higher volume of lighter weight oleoresin than a heavy one for the same investment
in carbon and other plant resources.
47
Based on the results of drilling two holes into each tree, Alencar (1982) indicated that all of the
available oleoresin may come out of the first hole drilled, and thus implies all of the secretory canals are
connected to each other. He acknowledged this runs counter to an assertion of an earlier Brazilian authority
(Pio Corrêa, 1932) who stated that copaiba’s longitudinal secretory canals form long pockets and that the
canals of some zones do not connect with those in others. Alencar seems to assume that most of the oleoresin
that is tapped comes from material draining from severed active resin ducts so drilling only one bore hole is
sufficient to tap this interconnected network of canals. The Tekohaw study was not designed to strictly test the
effect of bore hole number on oleoresin yield, but results lend more support to Pio Corrêa’s (1932) original
view. There were six trees where the first hole drilled yielded more oleoresin than the second hole, 13 trees
where the second hole’s yield was greater than the first and 2 trees where both holes’ yield was the same. In
the 13 cases where only one bore hole yielded any resin, the second bore hole was the source of the resin 77%
of the time. Other findings stated above support the idea that initial oleoresin harvest primarily taps old
accumulations in heartwood cavities, although subsequent harvests may draw from resin that has dripped out
of recently severed resin ducts in the cambium and sapwood.
Alencar’s (1982) indication that none of the nine hollow trees at his site yielded any oleoresin points
to another potential difference between the studies regarding the degree of compartmentalization of modified
wood types in copaiba trees. The average oleoresin yield for hollow trees drilled at Tekohaw was less than 1
ml. per harvest, but 18.5% of such trees did yield a small amount of resin. Heartwood rot and hollowness may
consume most of a tree’s heartwood, but at least a minor capacity to produce an oleoresin apparently
sometimes remains. The importance of confirming the negative association between hollow trees and
oleoresin presence, though, is that it corrects an apparent earlier misconception about this relationship. Spruce
(1970; p. 162), for example, described the mechanism leading to copaiba harvest in the following way: “In old
trees the trunk becomes hollow at the core, and there the oil accumulates and is extracted by boring with an
auger.”
There are differences in oleoresin harvest levels between the Manaus and Tekohaw copaiba trees, but
these are negligible compared to the popular references that claim copaiba trees yield an average of 2.5 to 24
liters per harvest with maximum amounts up to 60 liters. Since these accounts are not based on systematic
48
research, it is impossible to identify specific factors that might be responsible for this large perceived gap.
Tree type, size, condition, and season have been discussed as potential influences in comparing the Manaus
and Tekohaw studies. Potential differences that might result from radically different harvesting methods could
also influence harvest levels. As mentioned before, axe harvesting would be the surest way to expose oleoresin
cavities and canals; it would also be the most destructive. The two height harvesting scheme referenced in
popular accounts and mentioned by Tembé elders deserves some consideration. It is an interesting coincidence
that the upper height mentioned as the prime spot for drilling (or chopping) a second time is around the area
where the heartwood to total cross-sectional area ratio is known to be greater than at the base. Depending on
the species, this ratio reaches its peak some 2 - 7 meters above the ground and then tapers inward higher up
(Hillis, 1987; Stokes and Berthier, 2000). Whether this heartwood bulge exists to give the tree some special
structural properties or maintain a particular sapwood width optimized for water transmission (Berthier et al.,
2001), piercing it higher up potentially accesses a greater volume of stored oleoresin. Since boring into a tree
near ground level sometimes produced a sucking sound that might pull some preformed oleoresin higher up in
the trunk, making a second cut higher up could conceivably break the pressure tension in the canals and more
easily permit the resin to flow down to the lower hole through the force of gravity. Breaking off an upper limb
as was suggested by one harvester near the Tembé reserve could have a similar effect. In this study the
observation that resin flow increased from one tree when the wind was blowing hard could also be explained
on this principle. A controlled experiment that compared a lower hole harvesting method to a lower and upper
hole harvesting strategy would be needed to properly discern any real differences that result from these two
techniques.
THE INTERNAL LIFE OF A COPAIBA TREE
The three main questions implicit in this discussion that remain to be answered are what conditions
stimulate the production of oleoresin in copaiba trees, what conditions lead to the formation of resin rich
cavities, and what conditions lead to the loss of oleoresin formation and storage. As mentioned earlier,
anatomical evidence shows that copaiba trees produce longitudinal ducts arranged in concentric rings (Record
and Mell, 1924; Langenheim, 1973; Alencar, 1982; Vetter and Botosso, 1989). Each new ring is presumably
49
created once a year in the cambium. While it has been assumed that the parenchyma cells lining these ducts
regularly produce resin and deposit it into the canals, there is no direct evidence that this always occurs. The
Tekohaw study has shown that wounding a copaiba tree with a nail, a drill or a fire may lead to the outflow of
a small to moderate amount of oleoresin, but in most cases these external injuries do not stimulate any resin
flow at all. While insect or pathogen attacks may elicit greater resin production than purely mechanical injury
in other oleoresin producing trees (Phillips and Croteau, 1999), their response to any external wound is much
more consistent than copaiba’s. While copaiba oleoresin is always found in leaves (Langenheim, personal
communication 2001), oleoresin formation in the trunk may be mostly made only when the tree is induced to
do so under particular circumstances.
One precondition for copaiba trees having harvestable quantities of oleoresin seems to be a minimum
age or size. The smallest tree in the Tekohaw study to yield any resin was 27.5 cm DBH - a tree that is over 50
years old if its diameter grew at average rate of 0.5 cm per year. Given the wide use of copaiba oleoresin as an
anti-microbial medicine and the demonstration that leaf resins have anti-fungal properties (Arrhenius and
Langenheim, 1983, 1986), a fungal or bacterial pathogen is a likely candidate for the agent needed to stimulate
oleoresin production in the tree. Fungal agents have been found responsible for generation of resins collected
from infected Dragon tree (Draceana draco) and gharuwood trees (Aquilaria spp. and Gonystylus
bancanus)(Gianno, 1984; Langenheim, in press). Part of the process might be linked to a genetically controlled
stimulus.
One speculative scenario for production and storage of oleoresin in copaiba is as follows. When
oleoresin is produced but not released to the outside, it may remain in the longitudinal resin ducts. As the tree
grows, the older ducts formed in the cambium spend several decades in progressively senescing sapwood. At
the sapwood-heartwood transition zone, additional starch resources may be mobilized to create extra amounts
of oleoresin before the living cell machinery collapses. It has been shown that heartwood specific phenolic
compounds have been created in this zone (Hillis, 1987), but the process for terpenoid resin deposition in the
heartwood has yet to be explained (Langenheim, personal communication 2001). As a copaiba tree reaches the
50 cm DBH size (probably after 100 years of age), the oldest resin ducts near the center of the tree seem to
disintegrate through lysigeny or some other process and form larger spaces. This might be caused by a specific
50
pathogen or be linked to age related erosion of inner heartwood phenolics. In Scots pine (Pinus sylvestris L.),
the concentration of the stilbene pinosylvin decreases from the outer to the inner heartwood, possibly due to
oxidation or polymerization of this decay resistant chemical (Bergström et al., 1999). If there is a similar loss
of potency in copaiba heartwood phenolics in the oldest section, the release of oleoresin that has been shown to
have anti-oxidant properties (Desmarchelier et al., 1997) could provide a second line of defense against
microbial and insect attack. No defense is perfect or lasts forever, and it seems that copaiba trees in which
hollows have formed have lost their oleoresin defense perhaps because newly formed oleoresin can not be
channeled to these areas and older accumulations of resin have drained out through crevices of deteriorating
wood. If this scenario is correct, harvesting oleoresin that has been building up in the trunk of a tree for many
years that is not replenished could hasten the internal demise of the tree.
It seems possible that drilled copaiba trees that yield with copious flows of water instead of oleoresin
may have been infected by some type of “wetwood” infection that surpasses or takes hold after oleoresin
defenses have deteriorated. Accumulations of water have been observed in the heartwood of other trees such
as Cryptomeria japonica D. Don, although its cause was unknown (Nakada et al., 1999). Extensive work on
anatomical, physiological, and biochemical aspects of copaiba resin and canal formation and dissolution would
need to be done to assess various aspects of this scenario.
ESTIMATES OF COPAIBA OLEORESIN HARVESTING BY AREA AND TIME – IMPLICATIONS
FOR HARVESTERS
The density of copaiba trees, the average amount of oleoresin that can be harvested per tree, and the
number of harvests that can be obtained from a tree are the most significant factors that will determine the
viability of a commercial copaiba operation since their range of potential values may vary by 15 fold between
the minimum and maximum likely values from one area to another. The results of the Manaus (Alencar, 1982)
and this study show that potential harvest of oleoresin and revenue are likely to be in the minimum to medium
range. At this level commercial harvesting would not be worthwhile. In a case where copaiba trees were
relatively abundant and they actually could yield 2.5 liters per tree per harvest, commercial harvesting could
generate two to seven times the Brazilian daily minimum wage for someone with access to a large amount of
51
intact forest. While this optimistic scenario has not yet been documented in a specific population, the readily
available supply of copaiba oleoresin in various markets indicates that harvesting is commercially viable for at
least some people.
The effect of fire on the health of trees and resin production for current and future cohorts is decidedly
negative. The presence of burned trees translates into a higher percentage of hollow trees that lowers the ratio
of productive to total trees. In areas where people hope to harvest copaiba in the future, those trees, and if
possible that area, should be protected from intentional agricultural and accidental fires.
The longer term prospects for copaiba oleoresin supply may depend on how people treat forests in
general. The overall density of copaiba trees and the percentage of those trees that yield oleoresin will directly
decline in relation to the amount of forest that is burned. How copaiba trees are harvested can also make a
large difference in the yield and health of the trees. Axe harvesting may yield the most oleoresin, but this
practice is undoubtedly bad for the individual trees (Pittier, 1926; SEBRAE, 1995) and may have been
responsible for a region-wide population decline of copaiba trees (Dodt, 1939 cited in Sales, 1994). The
lethality of the technique probably depends on the severity of the wound and resiliency of the tree. Oleoresin
has been harvested from several species of dipterocarp trees in Southeast Asia for many decades by carefully
chopping holes into the trunk and setting the hole on fire (Gianno, 1984, 1990; Ankarfjärd and Kegl, 1998).
Even improper drilling of copaiba trees, however, may kill a tree within three years (SEBRAE, 1995).
The harvesting time model (Table 2.9) demonstrated that the density of copaiba trees also has a strong
effect on harvest efficiency. In low density areas, the time required to find potentially productive copaiba trees
dwarfs the amount of time devoted to actual harvesting whereas in high density areas, the search factor is
almost irrelevant.
Although this variable was not included in the model, search efficiency also varies considerably with
a prospective harvester’s ability to correctly identify copaiba trees at a distance since this determines their
search rate. Elder Tembé who had a well honed search image for the subtle bark blemishes and canopy form
of copaiba trees spent much less time finding them in the midst of a diverse forest than younger men who
needed to carefully examine many trees up close to determine their identity. Detailed knowledge or sense
about a tree’s habitat preferences also influenced search efficiency. Even though overall copaiba density was
52
low at a large scale at Tekohaw, spending more time looking carefully in promising sites and skirting forest
patches where such trees were not generally found probably increased the number of trees found per day.
A comparison of the wait and collect oleoresin resin harvesting strategy (Method A) versus the come
back and collect oleoresin strategy (Method B) in the harvesting time model (Table 2.9) showed that these
strategies have similar daily harvesting rates under the least favorable circumstances. When trees are more
abundant and yield moderate to large amounts of oleoresin, however, it becomes increasingly advantageous to
spend the extra time setting up a weather resistant collection device and return later to collect the accumulated
material. The advantage of this method is indicated by a greater harvest efficiency (Trees and Resin Harvested
per Week), financial return for time invested ($/Day), and the total amount of time that would be needed to
harvest all of the trees in a given area (Days to Harvest Area). The best strategy is probably a common sense
hybrid of the two methods. If the initial resin flow was weak, it would be worth waiting to collect most of the
oleoresin, plug the tree and not devote further time to a follow-up visit. If the initial flow was strong and
seemed likely to continue for an hour or longer, setting up a collection device would probably pay off because
it would avoid wasting time waiting and allow collection of all the oleoresin - not just the majority that would
emerge in several hours.
Where only a few people harvested copaiba in a large area of forest, this activity would be as
remunerative as collecting many other forest products. As was seen in the area based model, though, the
potential revenue from copaiba harvesting would yield very poor economic returns to harvesters unless
conditions were at least as good as described in the medium set of criteria.
COPAIBA OLEORESIN MARKETING
As is true with most extractive products, the middle men fare considerably better than the collectors
(Pendleton, 1992). Small-scale merchants who sell copaiba oleoresin from medicinal plant shops in Belém
receive $R 15 - 40 per for a full liter bottle - two to six times more than the $R 7 they normally pay the
harvester. These vendors usually sell the liquid in small 20 ml. plastic containers for $R 2 which means the
resale price is $R 100 per liter. Companies that purchase larger quantities of copaiba oleoresin sometimes
filter the oleoresin to resell some in small containers for $US 2.80 (equivalent to $US 140 - 188 per liter).
53
Most of the product is distilled so the resin and purified oil can be sold separately. The resin is generally
packaged in 5 kg sacks whose minimum price is $US 2.60 per kg. The purified essential oil is sold to
industrial users in 200 liter barrels for a minimum of $US 10 per liter or packaged in bottles with 50 gel
capsules each containing 500mg of oil that sell for $US 12 each - equivalent to $US 480 per liter (MMA/SCA
et al., 1998). There are many equipment and labor costs associated with these operations (SEBRAE, 1995;
MMA/SCA et al., 1998), but it is clear that the profit margin of corporate copaiba sellers is still much higher
than the collector’s.
Proposals to attract investment in a small copaiba oleoresin processing and packaging factory in a
state such as Acre predict that the project could generate a net revenue of about 25% and pay back the initial
investment in less than two years (SEBRAE, 1995; MMA/SCA et al., 1998). These proposals are based on the
input of 4.4 tons (about 5000 liters) of raw copaiba oleoresin per year. This would be processed and sold as
150 sacks of resin, 3000 liters of purified oil in barrels and 25,000 bottles of capsules. Considering reports that
the state of Amazonas annually produces more than 100 tons and Rondônia production had reached 30 tons per
year by 1994 (MMA/SCA et al., 1998), this does not seem like an overly ambitious goal. This analysis,
however, is based on the unproven assumption that copaiba trees can yield an average of 2.5 liters of oleoresin
per year for many years (SEBRAE, 1995) so the production target could theoretically be met by 2000 trees.
Since the proposal recommends that trees are harvested every other year, it really means that 1000 trees would
need to be harvested every year with an average yield of 5 liters per tree each time they are tapped. In Acre,
copaiba density is believed to be about 0.27 trees per hectare (SEBRAE, 1995) so the operation would need to
tap every copaiba tree in a 7407 ha area of forest. If the patterns of oleoresin harvest found at the Manaus
(Alencar, 1982) and the Tekohaw study hold true in Acre, only 25-50% of the trees will yield appreciable
amounts of oleoresin, the average yield from these trees will be much less than predicted even for first time
harvests, and the average yields will drop dramatically after the initial harvest. If these less favorable
conditions prevail, an area 10 to 1000 times greater would be needed to sustain the oleoresin production goals
for this one factory. Production studies will be conducted with copaiba trees in Extractive Reserves in Acre to
verify whether the optimistic yields included in the factory proposal are justified (Daisy A. P.G. Silva, personal
communication 2001).
54
WILL PLANTATIONS BE THE MAIN SOURCE OF COPAIBA IN THE FUTURE?
Studies in other copaiba rich areas in Brazil (Pará, Amazonas, and Rondônia), Venezuela and other
countries could help broaden the understanding of the true oleoresin production potential of wild populations.
Alencar (1982) concluded that the wild population of C. multijuga could supply limited amounts of oleoresin
for medicinal purposes, but that production levels were insufficient to consider its use as an alternative source
of energy to fossil fuels. Early projections were that one acre (0.4 ha) of 100 mature copaiba trees could
produce 25 barrels – about 52 liters per tree per year (Wang and Huffman, 1981). It was, therefore, logical that
experimental plantations of C. multijuga were initiated in the early 1980s to test its potential for oleoresin, pulp
and timber production (MMA/SCA et al., 1998). Related work has shown that copaiba seeds have good
germination rates (Alencar, 1981) and have good survival in a plantation setting. After 16 years, trees had
reached an average size of 9.2 cm DBH (MMA/SCA et al., 1998). This represents an annual diameter growth
rate of 0.575 cm per year. This rate for juveniles growing in full sun is not surprisingly faster than the average
equivalent growth rate 0.2595 cm per year for all size classes of a wild population of C. pubiflora (Ramirez and
Arroyo, 1990). Even if the plantation growth rate is steadily maintained, though, it will take 70 - 87 years for
these trees to reach the 40 - 50 cm DBH range - the likely minimum size that they could potentially yield
substantial amounts of oleoresin. Trees were planted at a 2.5 x 5 meter spacing (MMA/SCA et al., 1998)
giving a density of 800 trees per hectare. This plantation will inevitably be thinned over time so if survivors
achieve the same crown area as their counterparts in the wild population of C. multijuga (145 square meters at
45 cm DBH - based on data in Alencar, 1982), they will be reduced to a density of 69 trees per hectare. This
higher density could dramatically improve harvesting efficiency, but it will take until the middle of this century
to find out if a monoculture plantation of these trees will produce more or less oleoresin than their noncultivated relatives that have been exposed to the full array of biotic influences in a diverse forest.
Even if oleoresin yields are similar or better than found in wild populations, it seems unlikely that the
price for copaiba oil will be sufficiently high to economically justify more than 70 years of investment in a
copaiba plantation intended solely for oleoresin production. The experimental plantation could, however,
serve as a field site for certain types of controlled experimental research that would be difficult to undertake in
genetically diverse and dispersed wild populations. While trees are growing it could be worth examining the
55
effect of etherel (based on the phytohormone ethylene) on copaiba trees since it has been shown to stimulate
heartwood formation and increase production of various exudates in other trees (Hillis, 1987). Other chemicals
and specialized wounding techniques have been specifically used to stimulate resin production in a variety of
conifer and hardwood trees (Howes, 1949; Anderson, 1955; Hillis, 1987, Langenheim, in press). Trees that are
slated for thinning could also be used to carefully assess the extent that drilling induces wound wood, inhibits
heartwood formation and otherwise affects future oleoresin harvest. Other variations on harvesting techniques,
such as the high and low drilling scheme, could be tested under more controlled circumstances than would be
possible with a wild population.
SUMMARY AND CONCLUSIONS
This study has verified many of the results of Alencar (1982) that wild populations of copaiba trees do
not all have the oleoresin yielding capacity that is often described in popular accounts. It has become clearer
that trees that are too young, too old, or hollow at any age do not yield large amounts of oleoresin. There may
be one set of circumstances that induce some copaiba trees to produce above average amounts of oleoresin in
its live tissues during middle age and a different set of circumstances that lead to the creation of resin-filled
cavities in the heartwood later in life. While trees in the upper middle size classes are likely to yield the most
oleoresin, there is no certainty that all such trees will do so. This study also reinforces the Alencar (1982)
study’s result that the first harvest of a copaiba tree is likely to yield more resin than any successive one. This
finding, likewise, is contrary to many popular accounts that assert trees can be tapped many times without
declining yields. The combination of low density, limited productive trees in a population, low yield per tree,
and limited number of potential harvests found in the Tekohaw and Manaus studies lead to very low estimates
for potential oleoresin harvest and financial return on a per area basis and per time basis for a harvester. These
contrast with proposals that indicate a small factory that processes and repackages copaiba oleoresin for sale in
various forms can be reliably supplied by a typical Amazonian terra firme forest and operate with a healthy
profit margin. While this study has made some interesting findings about the biological and ecological
parameters of copaiba oleoresin production and harvest in one area of the eastern Brazilian Amazon that is rich
in Copaifera diversity, additional studies on the physiology of oleoresin production and harvesting protocols
56
will be needed before its findings can be generalized or shown to be anomalies in relation to populations of
other Copaifera species in different parts of the Amazon.
Where copaiba trees are found to only yield one or two good oleoresin harvests, local income could
be enhanced by culling older harvested trees. Copaiba trees are widely valued for their timber (Chichignoud et
al., 1990; Chudnoff, 1984; Martini et al., 1994), but this double use of copaiba trees would need to be done in
concert with a well-managed operation that preserved sufficient fruiting trees to avoid negative impacts on its
population and other serious pitfalls that frequently accompany logging in the region (Uhl et al., 1991, 1997;
Verissimo et al., 1992; Johns et al., 1996; Barreto et al., 1998). Resin harvest in young trees has been
successfully integrated with timber harvest of older trees with longleaf pine (Pinus palustris) and slash pine (P.
elliottii) in North America and several dipterocarp species in Southeast Asia (Langenheim, in press).
Copaiba trees that have passed their oleoresin producing prime, however, should not necessarily be
thought of as trees that have lost their value to the forest or its people. A variety of invertebrates utilize the
small amounts of resin that sometimes ooze onto copaiba bark from natural causes or human related activities.
Several species of Euglossa and Tetragona stingless bees were seen flying close to fresh oleoresin on a copaiba
tree trunk. They presumably collected some of the resin for nest building or other purposes (Roubik, 1989;
Nogueiro-Neto, 1997). Adults and several nymphal stages of at least one species of assassin bug
(Apiomerinae: Reduviidae: Hemiptera) were often seen hiding in bark crevices near resin leaks with bits of
resin on their forelegs - presumably waiting to capture a bee. This behavior of applying sticky resin to
raptorial legs to aid in the capture of stingless bees has been observed in several genera of this group in Costa
Rica and Brazil (Johnson, 1983; Adis, 1984). The Tembé are very familiar with this assassin bug and give it
the more distinguished titles “maranu’yw” (Tembé name for a forest spirit) and “don de copaiba” (Portuguese
phrase meaning owner or caretaker of the copaiba tree).
Even fire damaged and the oldest trees at Tekohaw and Manaus (Alencar, 1984, 1988) that yielded
little or no oleoresin continued to reproduce for many years. Beyond collecting resin, stingless bees also visit
copaiba trees to gather pollen from its flowers (Crestana and Kageyama, 1989), and aril coated seeds are
consumed by a variety of insects (Ramirez and Arroyo, 1987a, 1987b; Leal and Oliveira, 1998), birds
(Plowden, in press), and mammals (Alencar, 1984). Tembé hunters are well aware of these connections since
57
they searched for toucans in the canopy of fruiting copaiba trees and built “esperas” (an impromptu tree perch
made with stout young trees lashed several meters above the ground between two trees) to wait for game
animals such as agoutis, pacas, and deer that feed on fallen copaiba fruits at night. This resource is particularly
important in the dry season when fewer forest fruits are typically available (Alencar, 1988; Lourival Tembé,
personal communication 1996). Even non-reproducing copaiba trees may also have special value to wildlife.
Di Bitetti et al. (2000) showed that C. langsdorffii was one of four species of large trees frequently used as a
sleeping site by a group of capuchin monkeys (Cebus apella nigritus). The copaiba and other preferred trees
were all tall emergents with large crown diameters and many horizontal branches and forks that provided
stable night-time resting places.
The economic prospects of copaiba oleoresin harvesting may not always be attractive in many
situations, but it would be inadvisable to use purely economic rationale to argue for their preservation or
maintaining the forests that contain them. Copaiba oleoresin will be harvested and used by indigenous and
other forest dwellers for their own use whether or not it is worthwhile to sell. The Tembé in particular bemoan
the loss of copaiba and other trees valued by their culture to vast areas of forest burned by non-indians that
have occupied the Alto Rio Guamá reserve. On the other hand, the desire for sustainable economic
development among forest peoples is quite real, so these aspirations will probably need to be met through
means other than reversion to the intensive and probably predatory exploitation of copaiba and other similarly
scarce extractive forest resources.
58
TABLE 2.1 COPAIBA TRUNK DIMENSIONS OF ALL DRILLED, OLEORESIN
YIELDING AND HOLLOW TREES AT TEKOHAW
DIMENSIONS
N (#Trees)
(Mean Dia. Drill
Height* (cm))
MEAN ± S.E.
ALL DRILLED TREES
47 (58.3)
Distance to Heartwood (cm)
14.2 ± 0.4
47 (58.3)
Heartwood Diameter (cm)
30.0 ± 2.4
47 (58.3)
Heartwood Area/Total Area %
26.0 ± 2.2
OLEORESIN YIELDING TREES
8 (57.4)
Distance to Oil (cm)
21.3 ± 2.0
8
(57.4)
Oil Zone Radius (cm)
7.2 ± 2.0
DRILLED HOLLOW TREES
17 (67.3)
Distance to Hollow (cm)
18.7 ± 1.8
17 (67.3)
Hollow Radius (outer edge)(cm)
15.0 ± 2.4
13 (71.4)
Hollow Area/Heartwood Area %
55.1 ± 8.6
13 (71.4)
Hollow Area/Total Area %
15.0 ± 2.9
Hollow area estimation method: Where end point of hollow during drilling is on near side of calculated trunk
center, hollow area calculated as if beginning and end of hollow represent inner and outer points of a hollow
ring in entire cross sectional area of trunk. Where end point of hollow during drilling is on far side of
calculated trunk center, hollow area calculated as a circle with a radius based on the average of the beginning
and end hollow points.
*Dia. Drill Height: Average diameter of tree at heights where it was drilled. Drill height ranged from 0.8 to
1.2 meters above ground level. Due to tapering of the trunk, these diameters were larger than diameters
measured at 1.5 meters (DBH). They are included here for reference since dimension distances were also
measured at drill height.
59
TABLE 2.2 CORRELATION BETWEEN COPAIBA TREE DIAMETER AND
OTHER TRUNK DIMENSIONS AT TEKOHAW STUDY SITE
DIMENSION
N (# Trees)
Corr. Coeff.1
R-Sq. (%)2
p-value
47
.198
3.9
.181
Distance to Heartwood
47
.933
87.0
.000
Heartwood Diameter
47
.755
57.0
.000
Heartwood/Total Area %
8
.278
7.7
.505
Distance to Oil
8
.245
6.0
.559
Oil Zone Radius
17
.363
13.2
.152
Distance to Hollow
17
.709
50.3
.001
Hollow Radius (outer edge)
13
.691
47.8
.009
Hollow Width
13
.746
55.7
.003
Hollow Area
13
.202
4.1
.507
Hollow/Heartwood Area %
13
.568
32.2
.043
Hollow/Total area %
Notes: 1. Pearson’s Correlation Coefficient; 2. Linear regression R-squared value. Correlations compare trunk
dimension with tree diameter measured at average drill height (approximately 1 meter above ground).
60
TABLE 2.3 HARVEST OF OLEORESIN FROM COPAIBA TREES AT
TEKHOHAW STUDY SITE BY TREE TYPE
OLEORESIN HARVEST CLASS
Class 1 (0 ml)
Class 2 (1-49 ml max in one harvest)
Class 3 (>50 ml max in one harvest)
SUMMARY HARVEST DATA
(Mean ± S.E.)
ALL TREES
Ave. Total Harvest (ml)
Ave. Amount per Harvest (ml)
Ave. # Harvests
Ave. # Holes per Harvest
Ave. # Holes per Harvest with Oil
CLASS 2&3 TREES ONLY
Ave. Total Harvest (ml)
Ave. Amount per Harvest (ml)
Ave. # Harvests
Ave. # Holes per Harvest
Ave. # Holes per Harvest with Oil
WHITE
N = 34
23 (67.7%)
6 (17.7%)
5 (14.7%)
KNOBBY
N = 12
10 (83.3%)
1 (8.3%)
1 (8.3%)
RED
N = 11
6 (54.6%)
3 (27.3%)
2 (18.2%)
All
N = 57
39 (68.4%)
10 (17.5%)
8 (14.0%)
N = 34
103.3 ± 64.3
84.2 ± 60.5
1.9 ± 0.2
1.9 ± 0.1
0.5 ± 0.1
N = 11
319 ± 188
260 ± 181
2.0 ± 0.3
1.9 ± 0.1
1.5 ± 0.2
N = 12
22.9 ± 22.5
7.7 ± 7.5
1.8 ± 0.2
2.0 ± 0.2
0.2 ± 0.1
N=2
138.0 ± 133.0
46.2 ± 43.7
2.4 ± 0.5
2.5 ± 0.5
1.0 (no S.E.)
N = 11
65.0 ± 43.5
47.9 ± 39.1
2.2 ± 0.2
2.1 ± 0.3
0.8 ± 0.3
N=5
143.0 ± 87.3
105.4 ± 83.0
2.2 ± 0.4
2.4 ± 0.4
1.8 ± 0.2
N = 57
79.1 ± 39.5
61.2 ± 36.9
2.0 ± 0.1
2.0 ± 0.1
0.5 ± 0.1
N = 18
250 ± 117
194 ± 113
2.2 ± 0.2
2.2 ± 0.1
1.5 ± 0.1
TABLE 2.4 HARVEST OF OLEORESIN FROM COPAIBA TREES AT TEKOHAW STUDY SITE BY SIZE CLASS
DBH (cm) SIZE CLASS
OLEORESIN HARVEST CLASS
(Number and % of Size Class)
Class 1 (0 ml in every harvest)
Class 2 (1-49 ml max in one harvest)
Class 3 (>50 ml max in one harvest)
SUMMARY HARVEST DATA
(Mean ± S.E.)
ALL TREES
Ave. Total Harvest (ml)
Ave. Amount per Harvest (ml)
Ave. # Harvests
Ave. # Holes per Harvest
Ave. # Holes per Harvest with Oil
CLASS 2&3 TREES ONLY
Ave. Total Harvest (ml)
Ave. Amount per Harvest (ml)
Ave. # Harvests
Ave. # Holes per Harvest
Ave. # Holes per Harvest with Oil
25-35
N=6
35-45
N = 13
45-55
N = 11
55-65
N = 13
65-75
N=4
>75
N = 10
All
N = 57
4 (10.3%)
2 (20.0%)
0 (0.0%)
12 (30.8%)
1 (10.0%)
0 (0.0%)
7 (18.0%)
2 (20.0)
2 (25.0%)
6 (15.4%)
2 (20.0%)
5 (62.5%)
2 (5.1%)
1 (10.0%)
1 (12.5%)
8 (20.5%)
2 (20.0)
0 (0.0%)
35
10
8
N=6
2.7 ± 2.0
0.9 ± 0.7
2.3 ± 0.4
1.7 ± 0.2
0.5 ± 0.3
N=2
8.0 ± 4.0
2.7 ± 1.4
3.0 ± 0.0
2.0 ± 0.0
1.5 ± 0.5
N = 13
3.4 ± 3.4
1.1 ± 1.1
1.9 ± 0.3
1.7 ± 0.1
0.1 ± 0.1
N=1
44.0 (no S.E.)
14.7 (no S.E.)
3.0 (no S.E.)
1.3 (no S.E.)
1.0 (no S.E.)
N = 11
17.4 ± 11.6
13.8 ± 11.0
1.6 ± 0.3
2.0 ± 0.1
0.5 ± 0.2
N=4
47.7 ± 27.6
38.0 ± 28.3
1.8 ± 0.5
1.9 ± 0.1
1.3 ± 0.3
N = 13
305 ± 161
247 ± 155
2.2 ± 0.2
2.0 ± 0.2
1.0 ± 0.3
N=7
566 ± 267
459 ± 270
1.7 ± 0.4
2.0 ± 0.0
1.9 ± 0.3
N=4
71.0 ± 66.4
23.7 ± 22.1
2.3 ± 0.5
2.2 ± 0.6
0.8 ± 0.5
N=2
142.0 ± 128.0
47.4 ± 42.6
3.0 ± 0.0
2.9 ± 1.2
1.5 ± 0.5
N = 10
0.9 ± 0.6
0.5 ± 0.3
1.8 ± 0.2
2.1 ± 0.2
0.2 ± 0.1
N=2
4.5 ± 0.5
2.3 ± 0.3
2.0 ± 0.0
2.5 ± 0.5
1.0 ± 0.0
N = 57
79.1 ± 39.5
61.2 ± 36.9
2.0 ± 0.1
2.0 ± 0.1
0.5 ± 0.1
N = 18
250 ± 117
194 ± 113
2.2 ± 0.2
2.1 ± 0.1
1.5 ± 0.2
62
TABLE 2.5 CHANGES IN OLEORESIN HARVEST FROM COPAIBA TREES AT
TEKOHAW STUDY SITE BY YEAR AND HARVESTING ORDER
ALL DRILLED TREES
FIRST HARVEST
(All Trees)(ml)
1996
1998
FIRST HARVEST
(Oil Yielding Trees)(ml)
1996
1998
TREES DRILLED ≥2
TIMES (Oil Yielding Trees)
ALL HARVESTS BY
ORDER (ml)
Harvest 1
Harvest 2
Harvest 3
AMOUNT CHANGE BY
HARVEST (ml)
Harvest 1 to 2
Harvest 2 to 3
N
MEAN ± S.E.
MEDIAN
MINIMUM
MAXIMUM
57
72.5 ± 39.3
0
0
2028
33
24
18
35.1 ± 26.4
230 ± 118
230 ± 118
0
0
5.0
0
0
0
850
2028
2028
12
6
96.5 ± 71.1
496 ± 315
4.0
250
0
2
850
2028
12
12
9
96.5 ± 71.1
19.0 ± 12.1
16.3 ± 12.5
4.0
3.0
2.0
12
8
-77.5 ± 74.0
-8.6 ± 6.0
-1.0
-4.5
0
0
0
Maximum
Reduction
-850
-44
850
146
114
Maximum
Increase
144
9
63
TABLE 2.6 DRILLING TIMES IN COPAIBA TREES AT TEKOHAW
DRILLING VARIABLE
Time to Reach Heartwood (min./hole)
Time Drilling Heartwood (min./hole)
Time to Reach Hollow (min./hole)
Time to Reach Oleoresin (min./hole)
Time of Oleoresin Flow (min./hole)
Drill Rate in Bark/Sapwood (cm/min.)
Drill Rate in Heartwood (cm/min.)
N (#TREES)
20
11
4
4
5
17
11
MEAN ± S.E.
4.70 ± 0.53
20.05 ± 3.47
11.38 ± 2.45
16.12 ± 4.14
90.80 ± 34.3
2.75 ± 0.47
1.17 ± 0.27
MINIMUM
1.0
5.0
5.0
10.0
11.5
0.0
0.24
MAXIMUM
10.0
36.0
16.5
28.0
205.0
6.9
3.5
64
TABLE 2.7 OLEORESIN PRODUCTION FROM C. MULTIJUGA NEAR
MANAUS, BRAZIL
HARVEST NUMBER
HARVEST PERIOD
SEASON
AVERAGE RAINFALL
OLEORESIN HARVEST
CLASS N(%)
No Oil
Small Oil (<25 ml.)
Large Oil (>25 ml.)
Hollow
OLEORESIN HARVEST
MEAN (ml./tree)
All Trees (n=82)
Non Hollow Trees (n=73)
1
Mar.-Jun.
‘77
mid rainy
288 ml./mo.
2
Dec.-Jan. ‘78
early rainy
188 ml./mo.
3
Sep.-Nov. ‘78
late dry
75 ml./mo.
4
Sep.-Nov. ‘79
late dry
75 ml./mo.
5
Dec. ‘80
early rainy
188 ml./mo.
28 (34.2)
12 (14.6)
28 (34.2)
14 (17.1)
49 (58.8)
*
19 (23.2)
14 (17.1)
55 (67.1)
*
13 (15.9)
14 (17.1)
59 (72.0)
*
9 (11.0)
14 (17.1)
58 (70.7)
*
10 (12.2)
14 (17.1)
212.7
239.0
119.3
134.0
42.0
47.2
34.3
48.7
34.3
38.5
Notes: Data from Alencar (1982) Tables 2 & 3, Figure 11)(N=82 Trees)
* - Small oleoresin harvest not noted beyond first harvest.
TABLE 2.8 COPAIBA OLEORESIN HARVESTING MODEL BY AREA FOR LONGTERM AND SHORT-TERM PROJECTIONS
VARIABLE
Min. Estimate
Med. Estimate
Max. Estimate
Density (trees/ha)
0.3
0.7
1.5
Total Trees/Area (1000 ha)
300
700
1500
Productive/Total trees
0.15
0.30
0.50
Productive Trees/Area
45
210
750
Resin/Harvest/Tree (l.)
0.2
1
2.5
40 Year Projection
(Productive Life of Tree)
Number of Harvests in 40 Years
1
2
20
Total Resin (l.)/Tree
0.20
2.00
50.00
Resin (l.)/Tree/Year
0.005
0.05
1.25
$/Tree
$ 1.40
$
14.00
$
350.00
$/Tree/Year
$ 0.04
$
0.35
$
8.75
Resin (l.)/ha
0.009
0.420
37.5
Resin (l.)/ha/Year
0.000
0.011
0.938
$/ha
$
0.06
$
2.94
$
262.50
$/ha/Year
$ 0.002
$
0.074
$
6.563
Resin (l.)/Area (1000 ha)
9
420
37500
Resin (l.)/Area (1000 ha)/Year
0.2
10.5
937.5
$/Area (1000 ha)
$ 63.00
$ 2940.00
$ 262500.00
$/Area (1000 ha)/Year
$
1.58
$
73.50
$
6562.00
6 Year Projection
Number of Harvests in 6 Years
1
2
3
Total Resin (l.)/Tree
0.20
2.00
7.50
Resin (l.)/Tree/Year
0.03
0.33
1.25
$/Tree
$ 1.40
$
14.00
$
52.50
$/Tree/Year
$ 0.23
$
2.33
$
8.75
Resin (l.)/ha
0.009
0.420
5.625
Resin (l.)/ha/Year
0.000
0.011
0.938
$/ha
$ 0.06
$
2.94
$
39.38
$/ha/Year
$ 0.01
$
0.49
$
6.56
Resin (l.)/Area (1000 ha)
9
420
5625
Resin (l.)/Area (1000 ha)/Year
1.5
70.0
937.5
$/Area (1000 ha)
$ 63.00
$ 2940.00
$ 39375.00
$/Area (1000 ha)/Year
$ 10.50
$ 490.00
$
6562.50
Notes: All variables in bold are estimates of values based on data from this study and the literature. Other variables are
calculations based on those estimates. Medium estimates are based either on typical value in the literature or an average value
between minimum and maximum values. Area estimates are based on 1000 hectares (ha). Resin estimates are in liters. Value
estimates are based on Brazilian reais ($R) with $7/liter as base price paid to harvesters in 1998. Equivalent value in $US at the
time was approximately $R 1 = $US 1. Multi-year projections of values are not calculated with a discount rate or adjusted for
inflation.
66
TABLE 2.9 COPAIBA OLEORESIN HARVESTING MODEL FOR TWO
COLLECTION METHODS AT FIRST-TIME HARVEST
VARIABLE
Min. Estimate
Med. Estimate
Max. Estimate
VARIABLES FOR BOTH METHODS
Density (trees/ha)
0.3
0.5
1.5
Productive tree/total tree
0.15
0.30
0.50
Productive trees/ha
0.045
0.21
0.75
Square meters searched/min
500
500
500
Search Time/Productive Tree (min)
444
95
27
Resin/Harvest/Tree (l)
0.2
1.0
2.5
Drill Time/Tree (min)
40
40
40
Site Transit Time/Day (min)
90
90
90
METHOD A (Wait for Resin Flow)
Resin Flow Time/Tree (min)
32
110
256
Drill & Flow Time/Tree (min)
72
150
296
Search, Drill, & Flow Time/Tree (min)
516
340
349
Search, Drill & Flow Time/Day (min)
330
330
330
Trees Harvested/Day
0.64
0.97
0.94
Resin Harvested/Day
0.13
0.97
2.36
Trees Harvested/Week
3.8
5.8
5.7
Resin Harvested/Week
0.8
5.8
14.2
Search & Harvest Days/Week
6.0
6.0
6.0
METHOD B (Return for Resin)
Monitor Resin Flow Time/Tree (min)
20
20
20
Drill & Monitor Flow Time/Tree (min)
60
60
60
Search, Drill & Monitor Time/Tree (min)
504
155
87
Search, Drill & Monitor Time/Day (min)
330
330
330
Trees Drilled & Monitored/Day (min)
0.65
2.13
3.79
Trees Drilled/Week
3.3
10.6
19.0
Collect Resin Time/Tree (min)
10
10
10
Inter-tree Transit Time (min)*
16
7
4
Resin Collection & Transit Time/Tree (min)
26
17
14
Trees Collected/Day
3.3
10.6
19.0
Trees Harvested/Week
3.3
10.6
19.0
Resin Harvested/Week (l)
0.7
10.6
47.6
Search & Harvest Days/Week
5.4
5.7
5.8
Resin Harvested/Day (l)
0.12
1.88
8.15
SUMMARY COMPARISON OF METHODS
$/Day - Method A
$ 0.90
$
6.79
$
16.53
$/Day - Method B
$ 0.85
$
13.17
$
57.03
Days to Harvest 1000 ha - Method A
70
216
794
Days to Harvest 1000 ha - Method B
74
112
230
Notes: All variables in bold are estimates of values based on data from this study and the literature. Other variables are
calculations based on those estimates. One day is defined as 5.5 hours of work in the field plus 1.5 hours for travel
between the forest and the village. One week is defined as six days of work. Both methods invole drilling any potentially
productive trees found during the search process. All “productive” trees are drilled two times. Method A harvesters wait
to capture resin flow before resuming search. Resin flow time is based on the equation: Flow time (min)=12.2369 + 97.5 *
liters oleoresin. Method B harvesters search and drill potentially productive trees for five days. They monitor initial resin
flow and set up containers to capture strong flow. They revisit all productive trees on the sixth day to collect accumulated
oleoresin. Inter-tree transit time based on average density of productive trees and walking speed of 0.5 m/sec in the forest.
Value estimates are based on $R 7.00/liter paid to harvesters in 1998. At the time, $R 1 approximately equaled $US 1.
67
FIGURE 2.1 CROSS-SECTION ILLUSTRATION OF SOLID, OLEORESIN
BEARING AND HOLLOW COPAIBA TREES
PERCENT CONDITION IN SIZE CLASS
68
Hollow
Only
100
75
Hollow and
Fire
Damaged
50
Fire
Damaged
Only
25
0
20 - 40
(n = 17)
40 - 60
(n = 31)
60 - 80
(n = 17)
> 80
(n = 4)
DIAMETER SIZE CLASS (DBH cm.)
(n = total # trees in size class)
FIGURE 2.2 PERCENT OF HOLLOW AND FIRE DAMAGED COPAIBA TREES
AT TEKOHAW BY SIZE CLASS
% TRUNK AREA AT DRILL HEIGHT
69
(Measurements only from Drilled Trees)
40
Hollow Area
(n=13 trees)
30
Heartwood
Area (n=47
trees)
20
10
0
25 - 35
35 - 45
45 - 55
55 - 65
65 - 75
> 75
DIAMETER SIZE CLASS (DBH cm)
FIGURE 2.3 HEARTWOOD AND HOLLOW PROPORTION OF TRUNK AREA IN
COPAIBA TREES AT TEKOHAW SITE
MEAN OLEORESIN HARVEST (ml)
70
Average
Average
Amount
Amountper
per
Harvest
Harvest
1000
Amount
of First
Harvest
750
500
250
0
25 - 35
35 - 45
45 - 55
55 - 65
65 - 75
> 75
DIAMETER SIZE CLASS (DBH cm)
(For Trees with >0 ml. Harvest; n=18)
FIGURE 2.4 OLEORESIN HARVEST FROM COPAIBA TREES AT TEKOHAW
BY SIZE CLASS
71
AMOUNT OF OLEORESIN (ml)
Tree #(Hole #)
1400
15(2)
1000
58(2)
66(1)
66(2)
800
67(1)
67(2)
1200
600
400
200
0
0
25
50
75
100
125
150
175
200
TIME OF OLEORESIN FLOW (min.)
FIGURE 2.5 PATTERNS OF OLEORESIN FLOW FROM COPAIBA TREES AT
TEKOHAW
72
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APPENDIX 2-A COMMON NAMES FOR COPAIFERA TREES IN THE AMAZON
REGION
ARGENTINA: timbó-y-atá
BOLIVIA: copaibo
BRAZIL:
Tapajós: copaiba = C. multijuga, copahyba marimary, copahyba angelim; copahiba preta=C. glycycarpa
Trombetas: copahyba marimari = C. langsdorffii; copahyba jutahy = C. martii
Eastern Pará: copaiba-marimari = C. reticulata, copaiba jutaí
Jutai River Falls: copaiba-branca = C. guianensis
Maues: copaiba-cuiarana = C. glycycarpa
Santarem/Obidos: copaiba-rana, cupa-úba, balsam capivi = C. martii
Non-specified region: balsamo de copaiba, cabreuva, copahibera, copahiba, copahyba, c. angelim, c.
cuiarana, c. jutahy, c. marimary, c. parda, c. roxa, c. vermelha, copahúva, copaiba, copaibera, copaúva,
jaboti meutámeutá, jacaré copahiba, oleo, oleo folha, oleo pardo, olho, pau d’oleo, yébaro
CHILE: bálsamo de copaiba, copaiba del Brasil
COLUMBIA: trompo puerco = C. guianensis; aceite, arbol de aceite, canime, copaiba, copaibi, kurruma,
pata de gallo, queiyane (Kurripaco Indian) (possible species = C. officinalis, C. canime); canime vailuno =
C. pubiflora
FRENCH GUIANA: panchimouti (possible species = C. officinalis)
GUAYANA: balsam, maram (possible species = C. officinalis)
PARAGUAY: cupay, kupa’y
PERU: copaiba, copaiva, copaiba blanca = C. reticulata
SURINAM: apaoewa, hoepelboom, hoepelhout, hoepfroe-hoedoe, hoeproe, koepajoewa, koepawa,
pasoemoeti, passiemoetie (possible species = C. officinalis)
VENEZUELA: aceite, cabima, cabimbo, cabimo, calenibo, calimbo, copaiba, currucaí, curracay, kurukay,
maramo, palo de aceite, palo de aceitillo (possible species = C. officinalis; C. venezuelana
NON-SPECIFIED COUNTRY: balsam copaiba, copaiba, copaiva, copahu, Jesuit’s balsam
Sources: Anonymous, 1915; Cortés, N.S.; Record and Hess, 1943; Youngken, 1943; Le Cointe, 1947; Dwyer,
1951; Mors and Rizzini, 1966; Spruce, 1970; Leung, 1980; Chudnoff, 1984; Vega et al., 1984; Correa and
Bernal, 1989; Chichignoud et al, 1990; Sampaio, 1993; Phillips et al., 1994; Uibarri, 1997.
CHAPTER 3
THE HARVEST OF BREU RESIN FROM BURSERACEAE TREES IN THE
EASTERN BRAZILIAN AMAZON AND THE ROLE OF WEEVILS AND BEES IN
ITS FORMATION AND MARKETING
82
ABSTRACT
Many Brazilian Amazon forest dwellers collect and sell resins from various species of the genus
Protium (family Burseraceae). The resin that is locally called breu is used as a fire starter, incense, medicine
and wooden boat caulking material. A study conducted with Tembé Indians in the Alto Rio Guamá Indigenous
Reserve in eastern Pará state revealed that trees yielded an average of 0.8 kg and up to 11 kg of resin during an
initial harvest. Resin flow is stimulated by a previously undescribed species of weevil Sternocoelus new
species #1 (Coleoptera: Curculionidae). Weevil larvae bore into the inner bark and develop inside resin lumps
that form on the bark exterior. This is a novel adaptation by an insect to a terpene-rich compound that is
usually a potent defensive agent against pest species. At the Tembé study area and another site in the region,
resin was found more often and in larger amounts as tree size increased and tree health deteriorated. Trees in
forests that were occasionally flooded had larger resin harvests than ones in upland areas. Resin lump size was
closely correlated with the number and size of bore holes made by weevil larvae. Experimental drilling
demonstrated that large resin lumps only accumulate when trees are repeatedly bored by beetle larvae. Followup harvests at the Tembé site showed that trees that have already been infected by the beetles are more likely to
be colonized by additional beetles than uninfected trees. Weevil larvae may take one or two years to reach the
pupation stage. A regression model that includes tree diameter, the number of resin lumps harvested during the
first harvest, the weight of resin lumps harvested in the first harvest and the time since the first harvest predicts
it will take about four years between successive harvests for resin levels to rebuild to the initial harvest level.
Inventories in the Tembé area show that the types of Burseraceae trees likely to yield harvestable
resin have a density of 9 to 10 trees/ha. This places the site on the low end of one hectare plots surveyed in
various parts of the Brazilian Amazon where densities of such trees can exceed 50 trees/ha. Inventories of 100
ha plots, however, indicate that these high densities may not exist at a larger scale. It is estimated that as
density increases from 1 to 50 trees/ha, one experienced collector can harvest an average of 1 to 19 kg of resin
per day. Since resin is laid out in the sun to dry, these yields drop by 15-20% before the material is sold for
$US 0.25 to .50 per kg. This weight loss is probably due to the loss of volatile components in the resin, but an
83
experiment comparing drying rates of small resin samples that either permitted or excluded access to stingless
bees showed that these stingless bees can remove substantial amounts of resin.
Indigenous communities in the Gurupi River area probably harvest several tons of resin per year.
Since harvesters typically search away from villages, harvesting may have reduced the most accessible
supplies of resin. At a landscape scale, however, current harvest levels can probably be maintained because
the selling price for the resin and the market demand for it are both relatively low. Phytochemists are now
seeking to identify essential oils from breu resin that are marketable to the perfume industry. If such efforts
succeed and increase the market demand and price for breu resin, harvesters could earn more but pressures to
overharvest the resource would also increase. This study suggests that other researchers and communities
involved in resin harvest and medicinal plants in general should further explore possible connections between
plant defensive compounds and specific pest agents that may provoke their production. Such agents might
better be thought of and potentially managed as a new type of beneficial insect instead of a pest.
INTRODUCTION
Neotropical forest dwellers from the Yucatan to the Amazon have collected resins from trees in the
Protium Burman, Tetragastris Gaertner, and Trattinnickia Willd. genera (family Burseraceae) for subsistence
and commercial purposes for hundreds if not thousands of years (Alcorn, 1984a, 1984b; Turner and Miksicek,
1984; Rodrigues, 1989; Neels, 2000). In Brazil, the resin was formerly known as “hard elemi” (Howes, 1949;
Mantell, 1950); the most common current name for both the resin and the trees that produce it is breu
(pronounced “bray-oo”) (Rodrigues, 1989)(Appendix 3-A). The aromatic resin is burned to provide light, start
charcoal fires and make ceremonial smoke (Levi-Strauss, 1952; Daly, 1987; Rodrigues, 1989; Balée, 1994).
Various resins from select species are also used as a flavor, scent and a medicine to treat ailments including
skin infections and parasites, wound maggots, nasal congestion, rheumatism, toothache, hernias, poor vision,
conjunctivitis, respiratory, stomach and menstrual problems (Howes, 1949; Mantell, 1950; Mors and Rizzini,
1966; Reitz, 1981; Secoy and Smith, 1983; Alcorn, 1984(1); Plowman, 1984; van den Berg, 1984; Schultes
and Raffauf, 1990; Balée and Daly, 1990; Kainer and Duryea, 1992; Lima et al., 1992; Milliken et al., 1992;
Balée, 1994; Grimes et al., 1994; Boom, 1996; Comerford, 1996; Johnston and Colquhoun, 1996; Milliken and
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Albert, 1996; Siani et al., 1999(1); Neels, 2000). Sticky forms are used as glues in indigenous handicrafts,
glazing for ceramics, and caulking for canoes (Levi-Strauss, 1952; Whitten and Whitten, 1988; Balée and
Daly, 1990; Daly, 1992; Balée, 1994; Grimes et al., 1994; Johnston and Colquhoun, 1996).
Breu resin is found in various qualities. The most common type is a sticky balsam that comes from
species in the Protium and Tetragastris genera. It exudes from the tree in a clear to milky white liquid (Daly,
1987; Gentry, 1993). The resin lump that coagulates on the trunk may remain white or turn black depending
on the extent of oxidation and polymerization. “Breu branco verdadeiro” (true white breu) is prominently used
for caulking wooden boats (Rodrigues, 1989; Balée, 1994). “Breu jauaricica” or “breu preto” (black breu) is
darker with greenish blotches and is used more for its aromatic properties (Rodrigues, 1989). While there are
at least 35 species in the Protium genus alone in the eastern Amazon, only some of these produce harvestable
quantifies of resin (Daly, 1987; Balée, 1987, 1994).
In the eastern Brazilian Amazon, the Tembé and Ka’apor Indians (both Tupi-Guaraní groups) rely on
the harvest of breu resin (principally from Protium trees) as one of their main economic products (CEDI, 1985;
Balée, 1987, 1994). The quantity of resin harvested appears to be growing in line with their desire to generate
more income. As improvements to local infrastructure make villages more permanent, the local depletion of
natural resources such as breu resin becomes more likely. I conducted this study to quantify the amount of
breu being removed from the forest, understand some of the underlying mechanisms that influence resin
production, estimate the amount of time required for resin to regenerate post-harvest, and make
recommendations to guide development of a sustainable harvest mode. It was part of a larger project that
evaluated the production ecology of several plants yielding marketable non-timber forest products (NTFPs) in
the Alto Rio Guamá Indigenous Reserve (Pará state) and other parts of the region.
The Tembé technique for harvesting breu resin is straightforward. When someone encounters a breu
tree with a resin lump on it and wishes to collect it, he pries the lump off with his hand or machete. If the lump
is higher up on the trunk, he may knock it loose with a long stick. Lumps of dried resin that have fallen off the
trunk are also collected from the ground. Resin collected for personal use is done opportunistically during a
hunting trip or searching for other forest products. When collecting breu for sale, the harvester will dedicate
one or more days to that task with the goal of filling several large sacks with resin. Upon return to the village,
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the fresh resin is laid out on bags in the sun for several days to dry and prepare the material for sale. Collectors
all reported finding larvae (“tapuru”) in the resin lumps, but none claimed to know what these immature insects
turned into or what role if any they played in resin formation (Plowden et al., 2002).
The main objectives of this study were to answer the questions: 1) how much resin be harvested from
a population of breu trees and what factors influence variation in this amount, 2) how often can resin be
harvested from breu trees and what factors influence this frequency, and 3) how much income can forest
people make by harvesting breu resin. Under question one, the role of tree type, tree size, tree condition,
population density, size class distribution, forest type, and harvesting variables (distance from the village;
interval between successive harvests) were specifically investigated. As the involvement of weevils
(Curculionidae), bees (Apidae) and other invertebrates in this resin system became more apparent, other
questions were addressed as subsets of the main questions: 1) what role do weevils play in the formation of
harvestable resin, 2) what is the importance of breu trees and resin to these weevils, 3) how much fresh resin is
removed by stingless bees. A summary of specific research objectives, surveys and methods, analysis and
location of results in the chapter is listed in Table 3.1.
STUDY AREAS
The main study area was in the Alto Rio Guamá Indigenous Reserve located in eastern Pará state of
the Brazilian Amazon. It is the principal homeland of the Tembé Indians. Research was carried out in an area
of about 500 ha of closed tropical rainforest near the village of Tekohaw (020 37.7’ S; 460 33.1’ W) located on
the Gurupi River. Underlying soils are on the border of two major soil types classified by the Brazilian system
as yellow latossols and red-yellow podzols - the latter called spodosols in the American system (Projeto
Radam, 1973; Kalpagé, 1974). The site has three main forest types: upland (“terra firme”) forest, lowland
forest that is occasionally flooded (“baixo”), and forest near the main river and major streams that is usually
flooded during the rainy season. Near this and other villages, forest patches are cleared and burned to plant
manioc and other crops (Sales, 1993, 1994). This land use pattern creates a mosaic of forest in various stages
of current farm use, fallowed secondary forest and old forest. Other important aspects of the landscape are
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swaths of secondary forest that resulted from unintentional forest fires that passed through the area in the early
1980s.
Additional observations were made in a 100 ha terra firme forest plot on state road PA150 near the
town of Mojú, Pará (020 11.5’ S; 480 49.1’ W). The forests in both study areas are dominated by trees in the
Lecythidaceae, Burseraceae, Leguminosae, and Sapotaceae families (Baleé, 1987; Lockman, personal
communication 2000).
METHODS
RESIN HARVEST AMOUNTS PER TREE AND AREA OF FOREST
Breu Tree Density Inventories
Breu tree density at the Tekohaw study site was estimated in two successive belt-transect style
inventories. The first one carried out in January, 1998 surveyed trees and site characteristics in two 1000 ha
blocks of forest near Tekohaw village. Each block had twelve 500 m long north-south oriented transects
originating from the block mid-line that were located through a stratified random selection process. Transects
were sub-divided into 25 m long sections. Six out of the twenty sections in each transect were chosen by
stratified random sampling for 100% inventory of all trees ≥ 10 cm DBH (at 1.5 m) within 5 meters of the
transect. Forest type, common name, and DBH were noted for all inventoried trees. Breu densities were
analyzed in 2.55 ha of 102 forest plots (10x25 meters each) that were not recently burned or subject to
prolonged annual flooding. The second inventory was carried out in the same blocks in November, 1999 to
increase the size of the area surveyed specifically for breu trees. In this study, all breu trees ≥ 10 cm DBH
were inventoried in ten 500 m long transect lines in areas not recently burned or subject to prolonged annual
flooding. In addition to recording the same type of information gathered in the first inventory, the presence or
absence of resin on the trunk was also noted. Inventory results were analyzed to estimate the overall density of
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resin yielding breu trees, describe the size-class distribution of these trees and estimate the percentage of trees
in each size class that had harvestable resin.
Resin Harvest Studies and the Role of Weevils
In June and July 1998 and April 1999, a team of Tembé Indian men searched within 5 km of Tekohaw
village on the Pará side of the river for the kinds of breu trees that typically yield resin. The tree’s common
name, DBH, general health (live, injured, dead), GPS position, and forest type (“terra firme”, “baixo”) were
recorded. Each tree was marked with a numbered aluminum tag, and leaves were collected from some trees
for identification purposes. In the first month of the study, all breu trees were marked whether or not they had
resin. If resin was present, these lumps were harvested and weighed individually. The number and depth of
weevil bore holes found behind each resin lump was then recorded. In the second month, only trees that
actually had resin were tagged. All resin lumps were again harvested, but only the number of resin lumps and
their aggregate weight were recorded. The length and body width of larvae found in resin lumps were
measured. Other arthropods found in resin lumps were classified by general type. Weevil pupae and adults
were collected from host trees on the few occasions they were encountered. Results were analyzed to compare
resin amounts to tree type, tree size, general tree health, forest type, distance from the village (as a potential
indicator of previous harvesting intensity), and extent of weevil activity (number and size of weevils and
weevil bore holes). Data on weevils were also analyzed to describe natural history aspects such as the size
classes that correspond to larval instars and the relationship between weevil larval and bore hole number and
size.
Experimental Drilling
To experimentally compare resin buildup with mechanical wounds similar in shape and size to those
made by weevil larvae, 18 breu trees were drilled at the Tekohaw site in November, 1999. The objectives of
this test were to measure resin production from one time mechanical boring and explore the relationship
between resin production and hole diameter and depth. A hand drill was used to make a series of four holes on
the north side of each tree with a 4.2 mm drill bit; a similar series was made on the south side with a 3.4 mm
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drill bit. Each series consisted of holes drilled to depths of 5, 10, 15, and 20 mm with 40 mm separating
adjacent holes. These hole diameters and depths covered the upper range of hole sizes made by beetle larvae.
Each hole’s position was marked underneath with a small circle of red spray paint. The trees were revisited
four weeks after drilling to measure resin accumulation.
THE RATE OF BREU RESIN ACCUMULATION
Resin Yields in Follow-up Harvests
Two methods were used to measure the rate that resin accumulates on breu trees. These same
methods helped describe host tree colonization preferences of Sternocoelus weevils and the development rate
of their larvae in breu resin lumps. In the first method, I conducted follow-up visits to subsets of the trees that
had been first harvested or monitored in the summer of 1998. These visits occurred in November 1998, March
and July 1999 - approximately 5, 8, and 12 months after the initial harvest. A subset of the trees harvested 8
months after the initial harvest was harvested a third time 4 months later. During these visits, an effort was
made to individually remove and separately weigh resin lumps occupied by no more than one weevil larva.
The length, head width, body width, and weight were measured for larvae found in each resin lump.
Resin Lump Growth Rate and Weevil Development Time
In the second method of measuring the rate of resin accumulation, all resin lumps on some trees first
harvested in the summer of 1998 were individually marked with a numbered aluminum tag in March 1999.
During this and two other visits to these trees in April and July 1999 a thin gauge electrical wire was wrapped
around each resin lump. The wire was then transferred to a piece of paper where a tracing of its perimeter was
used to estimate the lump’s basal area. A comparison of resin lump weight to basal area obtained from other
harvested lumps was used to estimate the growth rate of these individually marked resin lumps by weight.
The results of both studies were analyzed to compare resin regeneration to factors such as breu tree
type, tree size, forest type, time since previous harvest, and amount of resin present at the previous harvest.
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The most significant factors were combined in a regression model to estimate the amount of time that would be
required for resin amounts to rebuild to the average found during the initial harvest. Data from the second
study were used to estimate the time required for one weevil to complete its larval (resin forming) stage.
BREU RESIN HARVESTING ECONOMICS AND MARKET
Breu Tree Search and Harvesting Time
During the initial resin harvest at Tekohaw in July, 1998, the time of arrival and departure at each tree
with resin was recorded. These data were used to estimate the amount of time needed to find a tree with resin.
Resin Drying Studies and the Role of Stingless Bees
.
Resin must be dried prior to selling, but since this process leads to weight loss from volatilization,
evaporation and harvest by stingless bees, it has economic implications for commercial harvesters. Resin
weight loss during drying was quantified at Tekohaw in July, 1998 by laying out batches of freshly collected
resin on plastic sacks for five to eight days outside in the sun. The total weight of each batch and the
proportion of different resin types (“branco” and “sarara”) was recorded before drying. At the end of each day,
the resin was reweighed. The weighing of individual batches concluded when weight stabilized for two
successive days. Results were analyzed to calculate the amount of weight lost during the typical procedure.
A companion experiment was carried out in April, 1999 to estimate the specific effect of stingless bee
resin removal on weight loss during the drying process. In this procedure, enclosures that were 25 cm on a
side were covered with plastic mesh and placed outside four days in a seven day period. Four enclosures were
covered on all sides; four had one side open so they were accessible to bees. One 36g sample of resin taken
from the same original breu “sarara” resin lump was placed inside each enclosure. Resin samples were
weighed at the beginning and end of every day samples were placed outside. On the second, third and fourth
days the samples were outside, the type and number of bees present in each enclosure at one time was recorded
in a spot check about every 30 minutes. A few specimens of the two types of bees that visited the enclosures
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were collected for measurement and identification. Results were analyzed to compare drying rates between
resin samples exposed to bee removal and samples that dried without such exposure.
Breu Resin Sales in the Village and Urban Markets
Information on the amounts and values of breu resin collected and sold by Tembé and Ka’apor
Indians was obtained primarily through conversations with one Tembé man who bought resin from Tembé
living in or near Tekohaw village. He further cooperated with the research by keeping a log book of resin
purchases and sales between July, 1999 and Feb. 2000. I obtained additional information about amounts,
values, and end uses of breu sold in urban markets by interviewing the owner of a forest products business in
Belém and managers of several boat repair businesses in the Belém and Icoaraçi that use resin to caulk wooden
boats.
Breu Resin Harvest Models
Information on average resin yield by size class of breu trees were combined with estimates of drying
rates and values to predict the amount and value of resin that could be harvested from a forest with different
densities of breu trees. A second model uses these variables as well as harvesting efficiency and the rate of
resin accumulation to predict the daily yield and earnings from breu resin harvesting and the amount of forest
that would be needed to employ a breu harvester full-time at different densities of breu trees.
RESULTS
RESIN HARVEST AMOUNTS PER TREE AND AREA OF FOREST
Breu Tree Density Inventories
The Tembé have numerous names in both their language and Portuguese for the wide array of breu
trees in their forest (Balée, 1987) that are based on bark, wood, and leaf characteristics. While most
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Burseraceae species in the area are probably resin producers, there are two major types (that both include
several species) that yield harvestable resin on the trunk. These are called “branco” (white) and “sarara.”
According to the Tembé, red “vermelho” breu trees are not a source of such resin. The belt transect inventories
conducted at Tekohaw showed that “branco” and “sarara” breu trees had a combined average density of 9.6
trees/ha (Table 3.2). The “branco” and “sarara” types contributed equally to this overall average, but the
densities of these two types varied considerably between the two main forest types. “Branco” type trees were
more common in the “terra firme” type forest, while “sarara” type trees had a higher density in the wetter
“baixo” type forests.
In the second inventory with the larger sample size the size class distribution of resin yielding breu
trees was heavily weighted in the 10-30 cm DBH size classes. Trees that were 10-20 cm DBH accounted for
76% of the trees; 20-30 cm DBH trees garnered 20% of the total, and the 30-40 and 40-50 cm DBH classes
each had only 2% of the total trees located.
Resin Harvest by Tree Type
Resin harvest studies at Tekohaw showed that at least seven species of Protium including P. pallidum
Cuatrec., P. trifoliolatum Engl., P. decandrum (Aubl.) Marchand, P. morii Daly, P. giganteum Engl. var.
giganteum, P. polybotryum (Turcz.)Engl., and P. glabrescens Swart and Tetragastris panamensis (Engl.)Kuntze
had resin lumps on them. P. pallidum and P. trifoliolatum were generally classified as “breu branco.” These
trees were found almost exclusively in well-drained “terra firme” type forest. Their resin accumulated in
rounded white lumps that turned grayish with age. The other six identified species were considered different
varieties of breu “sarara.” “Sarara” trees were found evenly distributed in both “terra firme” and wetter
“baixo” (low) areas of the forest. Their resin lumps tended to be flatter than the “breu branco” lumps. These
lumps also blackened to a greater extent and were often covered with dirt and plant-like debris. Resin from
“sarara” type trees is commonly marketed as “breu preto” (black breu).
Results of the initial harvest at Tekohaw showed that the average amount of resin harvested from live
and dead trees that had at least one resin lump was 856 ± 100g per tree (Table 3.3). A few trees yielded more
than 9 kg of resin, but the median harvest amount of 420g per tree better reflects the fact that many trees
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yielded relatively small amounts. In the first month of the harvest when data were recorded for trees with and
without resin, this average was 562 ± 65g per tree. Sample sizes of known scientific species were too small to
compare. A comparison of the main types of resin yielding breu trees showed that “sarara” trees had larger
amounts of resin than “branco” trees with a p-value of 0.09 when their means were compared with a One-way
AOV. The number of resin lumps per tree for the two types was almost identical (“branco”: 2.9 ± .2
lumps/tree; “sarara”: 3.2 ± .2 lumps/tree). Several other types of breu trees, principally the “vermelho” (red)
type, were relatively common, but since they never had resin lumps they were not included in the harvest
survey or inventories.
Resin Harvest by Tree Condition, Size, and Location
The initial resin harvest at Tekohaw showed that tree condition, size and site each explain some
variation in resin yield (Table 3.3). The clearest finding was that while only 12 dead trees were found among
183 trees examined, they had an average of twice as much resin on them as live trees (One-way AOV: p=0.01).
Resin yields in general increased with tree size, but the relationship was not a simple one. In the first
month of the initial resin harvest when all potential resin yielding trees were examined, the percentage of live
breu trees with some resin significantly increased with tree size class (χ2 = 31.91; df=4, p=.010). Only 9% of
live trees in the 5 – 9 cm and 20% of live trees in the 10 – 19 DBH size classes had any resin on them. The
percentage of live trees that were ≥ 20 cm DBH with resin was 68%. Regression analysis comparing resin
amounts found on all live trees in the initial harvest to tree DBH was significant (R2=3.4%; p=0.015; n=171).
Increases in resin amount with size were gradual, though, so means were not significantly different between
different size classes (Table 3.3). Non-parametric comparisons, however, showed that the median amounts of
resin harvested were significantly greater from trees that were ≥ 20 cm DBH compared to trees in the two
smaller size classes.
Resin harvest amounts also varied with the location of the trees at the Tekohaw site. The area
searched most intensively had both “terra firme” and “baixo” type forest. Trees in the wetter “baixo” type
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forest areas that were closest to the streams had almost twice the resin harvest as ones in the drier “terra firme”
forest (Table 3.3)(One-way AOV: F=6.39; p=.012).
The other factor that logically influences resin collection success is harvest intensity. Apart from this
study, there were no records of harvesting activity. Bark discoloration and bore holes on a trunk indicated
where a resin lump was once attached, but it was difficult to tell whether a lump was removed by a harvester or
fell off naturally and how long ago it came off. It was, therefore, hypothesized that harvesting intensity was
proportional to distance from the village. Using GPS positions, the distance of each tree with resin from
Tekohaw village was calculated and plotted against the amount of resin found on that tree. The average
amounts of resin harvested per tree increased with each kilometer (Figure 3.1), but overall such differences
were not significant (Table 3.3). It is noteworthy that all finds of 3.5 kg or more of resin per tree were found at
least 3.4 km from the village.
The Role of Weevils in the Formation of Breu Resin Lumps
During the resin harvests at Tekohaw, larvae were found in almost all fresh resin lumps. When larvae
were present in the second round of resin harvest, there was an average of 1.3 ± 0.1 larva per resin lump
(n=137 lumps). In June of the initial 1998 resin harvest, 93% of resin lumps had weevil bore holes behind
them. In a few cases, resin had apparently formed due to a burn or some other non-weevil related wound, but
weevil larvae were found in a few lumps not associated with bore holes. The average weight of resin lumps
with weevil bore holes behind them was 197 g compared to the 61 g average of resin lumps not associated with
bore holes.
An examination of one pupa and seven adults collected from breu trees and resin lumps at Tekohaw
showed these larvae were immature beetles that belong primarily or exclusively to a previously undescribed
species of bark-boring weevil (Sternocoelus Kuschel new species #1; Curculionidae)(O’Brien, personal
communication 2001). Other species of this weevil genus have been identified as stalk borers on manioc
(Manihot spp.) plants (da Costa Lima, 1956; Bastos et al., 1980), but this is apparently the first documented
association with a Burseraceae forest tree. One adult specimen of a closely related weevil (Eubulus Kirsch
species #1)(O’Brien, personal communication 2001) was also found on one breu tree.
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Weevil larvae removed from resin lumps in the second round of harvesting had an average head width
of 1.8 mm, average body width of 3.6 mm, and average body length of 8.2 mm (Table 3.4). The distribution of
head widths (possibly indicating discrete instars) showed two peaks around 1.0 mm and 2.1 mm with a
possible third peak at 1.4 mm (Figure 3.2). The weight of larvae measured in the field was 0.12 ± .02 g
(n=16). Measurements of a few pupae and young adults showed that mature weevils generally reach 10 mm
in length (including the head) and weighed just over 0.2 g.
Resin associated with weevil attacks on breu trees was usually found in lumps affixed to the lower
two meters of tree trunks although some lumps were occasionally seen higher up. Lower lumps were often
wedged in a crook between the trunk and a buttress or stilt root. Weevil related lumps were quite distinct from
the brittle streaks of resin that formed in machete cuts or small fissures of damaged trees. Lumps were
typically 2 to 6 cm thick and averaged 339 ± 59g in the initial Tekohaw harvest. Most distinct lumps had only
one developing weevil, but it was sometimes difficult to distinguish separate lumps in “sarara” type trees
where resin lumps abutted each other. The weevil larva formed a cavity in the fresh resin that was 3 to 8 mm
deep and extended almost to the edge of the resin lump (Figure 3.3). Yellow to greenish spots were sometimes
found in fresh white resin that may have been larval fecal material. A few weevil pupae and pre-emergent
adults were found in a rounded body-sized tunnel with smooth walls. Adult weevils that were not associated
with resin lumps were usually found in the bark of rotting portions of fire-damaged or dead trees.
Weevil larvae apparently feed on the inner bark of breu trees by chewing a simple cylinder shaped
hole toward the center of the tree. The mean number of bore holes was 10.1 ± 0.7 per resin lump in live trees
(n=171) in the initial resin harvest. While there was not a significant difference in the number of bore holes
between “branco” and “sarara” trees, the average depth of bore holes was significantly greater in “sarara” tree
lumps (7.1 ± 0.3 mm; n=81 lumps (776 holes on 19 trees) than in “branco” tree lumps (4.8 ± 0.2mm; n=27
lumps (226 holes on 13 trees) (One-way AOV; F=13.91; p=0.00). In the second resin harvest, there were 3.7 ±
0.2 bore holes per resin lump. A Fully Nested AOV examination of bore hole numbers and diameters
measured in those harvests showed that differences among resin lumps accounted for a higher percent of the
variation than differences among trees.
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The best evidence of the link between weevil activity and the formation of harvestable breu resin at
Tekohaw was the relationship between the number and dimensions of weevils, their bore holes and the weight
of resin lumps. The number of weevil larvae, sum of weevil head widths, sum of weevil body widths, number
of bore holes, sum of bore hole diameters, and sum of bore hole depths all had highly significant correlation (pvalues ≤ 0.01) with resin lump weight when compared with Pearson’s Correlation Coefficient and Regression
Analysis during the initial resin harvest (Table 3.5). The number and size of bore holes were somewhat better
predictors of resin lump weight than larvae number and dimensions. As seen from the weevil larvae point of
view in the initial resin harvest, the number and size of bore holes they chewed were generally better predictors
of larvae number and size than the amount of resin generated in the process (Table 3.6).
Experimental Drilling
The drilling experiment was conducted to measure the amount of resin flow generated by mechanical
action compared to the resin formed when the same amount of wood is chewed and removed by weevil larvae
making typical bore holes. The trees in the drilling experiment included three “branco” type (DBH ave. 23.8
cm) and fifteen “sarara” type trees (DBH ave. 28.7 cm). During the initial resin harvest the “branco” trees in
this experiment had an average of 660g per tree collected from them; eleven of the “sarara” trees yielded an
average of 1200g per tree. The other trees had no resin previously collected from them. When the trees were
revisited four weeks after drilling, little or no resin was found around these drilled holes. These results sharply
contrast with resin lumps generated by weevil bore holes where an average of 20 g of resin per hole was found
during the initial resin harvest. In 26% of the holes drilled, a tiny amount of resin (estimated to be less than
1g) had dripped out of the hole and dried on the trunk or was still leaking out of the hole. Drill bit size had no
effect on the percentage of holes that had any resin coming from the drill hole. The percentage of holes with
some resin on the trunk declined with increasing bore hole depth, but the difference was not significant.
Regression analysis showed that tree DBH (R2 = 5.4%; p=0.35) and the weight of resin harvested from the tree
during the initial harvest (R2 = 1.8%, p=0.60) were also both poor predictors of the number of holes that had
some resin on the trunk in this experiment.
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THE RATE OF BREU RESIN ACCUMULATION
Resin Yields in Follow-up Harvests
The expectation of follow-up visits to breu trees 4 to 12 months after the initial harvest at Tekohaw
was that resin harvests per tree would steadily increase over time. Due to the small number of trees harvested
each time in this second round, this trend was not found. (Table 3.7). On trees with any resin taken during the
second harvest, the average resin per tree was greatest in the group of trees revisited less than five months after
the initial harvest. The mean amount of resin per tree and number of resin lumps per tree harvested eight
months after the first harvest were greater than trees harvested more than a year after the initial harvest. A
subset of trees harvested for a third time four months after their previous harvest produced the fewest number
of resin lumps and the smallest amount of resin.
The time after the initial resin harvest was not by itself a good predictor of resin accumulation in these
second round harvests, but tree size was again highly correlated with the number of resin lumps (a measure of
weevil infection) and amount of resin taken off the trunk (Table 3.8). Harvest history as measured by the
number of resin lumps and resin weight present at the initial harvest were even stronger predictors of these
variables for the second harvest than tree size. The strongest correlation was found between the number of
resin lumps during the first and second harvests with a correlation coefficient of 0.8 and p-value of 0.000.
While forest type was significantly correlated with resin weight during the initial harvest of breu trees, this
factor was not significant in the second harvest round.
Resin Lump Growth Rate and Weevil Development Time
Monitoring the growth rates of individual resin lumps provided additional information on the rate of
resin accumulation and the development time of weevil larvae. The wire perimeter technique used to measure
resin lump basal area showed that this area and resin lump weight in the study trees were highly correlated
(Regression analysis R2 = 94.6%; p=0.00; n=22 lumps in 5 trees). Resin lump growth per day was estimated
by dividing the difference in estimated weight of the lumps in March and July, 1999 by the number of days that
lapsed between these two measurements. Regression analysis comparing these growth rates with the final
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estimated weight (about 12 months after initial harvest) shows that resin growth rate steadily increased with
resin lump weight until it reached a maximum around 0.56g/day at a resin lump weight of about 295g (Figure
3.4). Extrapolating the quadratic regression model to the point where the growth rate returns to zero (the point
when larval feeding is assumed to stop with the onset of pupation) indicates that maximum resin lump size is
around 589 g. An early suspension of the study prevented additional measurements to test this projection. The
mean amount of resin per larvae in the initial harvest, however, was 263.6 ± 53.9g so it is likely that the
average rate of resin formation sharply declines between 300 and 500 g. This pattern of resin formation
indicates that Sternocoelus weevil larvae may spend one, two or possibly three years in a breu resin lump
before pupating. The study provided no information on how long the pupa stage lasts before a young adult
bores its way out of a resin lump.
Breu Resin Accumulation over Time
Data from the second round of resin harvests and monitoring growth rates of individual resin lumps
were used to generate estimates of the time it would take for the average yield of resin per tree from a second
round of harvesting breu trees to return to the average of the initial harvest. Tree type, tree size, forest type,
initial harvest resin weight, initial harvest resin lump number, and time between first and second harvests were
tested in regression models individually and in combination to assess their ability to predict the amount of resin
harvested (or estimated to be present) during the second harvest. The model judged to be most useful because
of the certainty of its variables and high predictability (p-value = 0.000) included tree size (DBH), weight of
resin at first harvest, number of resin lumps at first harvest, and time between first and second harvests. The
regression equation was applied to all live trees monitored during the initial harvest to estimate resin amounts
that could be harvested from one to eight years after the first harvest (Table 3.9). The model predicts that
following a first harvest it would take about four and a half years before a second resin harvest could obtain the
same amount. The model assumes that the rate of resin accumulation remains constant regardless of the weevil
development time and that no resin yielding trees will die during this period.
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Other Animals Associated with Resin Lumps
Fly larvae identified as an undescribed species of Alipumilio Shannon (Diptera: Syrphidae)(Rotheray,
personal communication 2001) were the principal other insects found associated with breu resin. They were
found only in fresh “sarara” breu resin lumps in a section of the lump that was segregated from a weevil larva
cavity (Figure 3.3). One distinctive feature of resin containing fly larvae is that maggot movement (and
presumably feeding activity) in the resin transformed it from a white sticky cake-like material that breaks apart
when pulled to a dark taffy-like substance that stretched. While it seems possible that these fly larvae may
occasionally provoke some resin flow in breu trees, it was assumed that this makes a negligible contribution to
the amount of resin that is harvested for commercial purposes. The relationship between this fly and the breu
resin system are discussed in detail in Chapter 4 of this dissertation.
The most common visitors to the outside of resin lumps were several species of stingless bees
(Apidae: Meliponinae) of the genus Trigona (Venturieri, personal communication 2000). Bees were observed
collecting resin from breu trees in the forest at Tekohaw (Figure 3.3) and from piles of resin laid out to dry in
Tekohaw village. They first hovered over the resin, landed on it, and then applied bits of fresh resin to the
corbicula hairs on their hind legs. No attempt was made to quantify the amount of resin removed by bees from
resin lumps on trees in the forest. The scope of bee collection of resin from piles of human harvested resin
being dried in the village is discussed in the section on harvesting economics.
Assassin bugs (Reduviidae: Apiomerinae) were also commonly seen around breu resin. In the forest
these bugs were sometimes observed waiting motionless on breu tree trunks near resin lumps (Figure 3.3).
These bugs appeared to have some resin on their forelegs. The presence of these bugs has no major
implication for human resin harvesters. Their predation on stingless bees in the forest may reduce the amount
of resin that these bees remove by a minor amount.
The other invertebrates associated with breu resin are those that use the chambers of hardened resin
lumps left behind by weevils. One type of unidentified ant was the most frequent post-weevil occupant of
resin lumps. When dirty old resin collected in the initial harvest was broken apart, these slim ants were
sometimes found tending to pupae. Other invertebrates encountered in this process were flatworms, and
numerous arthropods including spiders, scorpions, centipedes, millipedes, and earwigs (Overal, personal
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communication 1998). The presence of these arthropods in dried resin has no major impact on human
harvesting, although, harvesters breaking apart dried lumps need to exercise some caution to avoid painful
bites and stings from some of these creatures.
BREU RESIN HARVESTING ECONOMICS AND MARKET
Breu Resin Harvesting Rates
Data from arrival and departure at breu trees during the initial harvest at Tekohaw showed that the
average time to find any breu tree was about 5 minutes per tree (about 9 minutes to find one with resin). The
density of breu trees that might have resin in this forest was estimated to be about 10 trees per ha (4 trees with
resin) so an estimated 12 trees could be found in an hour of searching. This means that harvesters search about
1.2 ha of forest per hour. Once a breu tree is found, it takes about four minutes to look for and harvest any
available resin. Harvesting was very quick when there are only a few readily accessible resin lumps, but the
time was substantially greater when trees had numerous lumps that sometimes required more effort to dislodge.
The combination of search and harvest time meant that about 6.7 trees in 0.7 ha of forest could be harvested
per hour.
Resin Drying Studies and the Effect of Stingless Bees
Sixteen batches of resin with initial weights varying from 2 to 9 kg lost the most weight in the first
three days they were placed outside and tapered off to near zero from the fourth day on (Figure 3.5). The
average mean percent weight loss after five days was 17.0 ± 1.2%. The ultimate weight loss was higher
(although not statistically significant) in batches with the largest percentages of “sarara” type resin in the mix.
Weight loss was assumed to be primarily due to the loss of volatile components of the resin. When resin was
still sticky in the first few days of drying, however, stingless bees were frequently seen collecting resin.
In the experiment testing the affect of stingless bees on resin drying rates, resin samples in the
partially screened enclosures lost a significantly greater mean percent of their original weight than the fully
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screened samples (bee treatment: 14.1 ± .3% vs. screen treatment: 3.5 ± .4%; One-way AOV: F=489.13;
p=.000)(Figure 3.6). The partially screened enclosures were visited by two varieties of stingless bees. During
the first day most or all visits were made by a small yellow bee (mean weight 14.9 ± .7 mg; n=4) called “ipawa
ruwer” by the Tembé (Tetragona sp. - now considered another variety of Trigona sp.)(Venturieri, personal
communication 2001). This bee was the only type that visited the resin samples until the second hour of the
second day of exposure. Larger black bees (mean weight 26.0 ± .6 mg; n=8) that the Tembé call “zawar
iruwera” (jaguar bee)(Trigona sp.) then began to enter the open enclosures, and the small bees were never seen
again for the remainder of the experiment.
Bees burrowed into moist resin and applied bits of it to the corbicula on their hind legs. They
sometimes flew away from one piece and then alighted again to continue gathering resin. One black bee spent
six minutes at the enclosure before flying away. The highest number of bees during spot checks was recorded
during the second day of exposure when an average of 7 bees per enclosure were seen at one time (maximum
of 46 bees during one spot-check in all enclosures combined). The average number of bees present during a
spot check declined to 2 bees per enclosure during the third day of exposure (when heavy afternoon rain
interrupted exposure time) and rebounded slightly to almost 4 bees per enclosure on the fourth and final day of
exposure. The decline over the days of monitoring was, nonetheless, statistically highly significant (One-way
AOV: F=22.31; p=.000). During three days of bee monitoring, the number of bees seen at the open enclosures
steadily increased from morning to afternoon observation times.
Breu Resin Sales
Up until 1996 Tembé men who harvested breu resin usually held on to their product until they or
someone from the government Indian agency (FUNAI) could bring their sacks to a dealer in the town of
Gurupi or the city of Belém to sell. These opportunities were unpredictable and often involved personal costs
for transportation and food. In 1997, two Tembé men in Tekohaw began to buy dried breu resin and titica
vines from Tembé and other Indians in the Alto Rio Guamá and neighboring Alto Turiaçu reserves. Harvesters
were usually paid $R 0.50 (about $US 0.50) per kg of dried resin. Most chose to receive payment in the form
of merchandise acquired by these Indian brokers during their selling trips.
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A sales logbook kept by the larger buyer in Tekohaw showed that 16 people (or small groups of
friends or relatives) sold a total of 1519 kg of breu resin to him between July, 1999 and February, 2000. The
mean amount sold by each person or family group was 95 ± 33 kg (median 29 kg; maximum 441 kg). There
were 39 transactions during this time with each customer accounting for an average of 2.4 sales (median 1.5;
maximum 8) with 39 kg (median 6 kg; maximum 200 kg) per sale.
This one Tembé broker sold one lot of 1000 kg of breu resin in 1998. He sold additional lots of 180,
300, and 770 kg of resin in 1999. When he sold to the dealer in Gurupi, he received 25% of the value of the
sale in cash up front and the rest in merchandise. The dealer in Gurupi usually paid $R 1.00/kg for pure
batches of “branco” resin or $R 0.70 for pure or mixed batches of “sarara”/”preto”(black) type resin. Dealers
in Belém generally offered $R 1.00/kg regardless of resin type and paid 100% of the sales in cash. These
buyers, however, usually deferred payment for several weeks or months until they sold the product to a dealer
in another city.
The main dealer in Belém had a retail store where he sold small quantities of breu resin and dozens of
other medicinal forest products. He sells most of his resin as a wholesaler at $R 3/kg to other dealers who
supplied shipyards in larger cities on the northeast coast of Brazil. My conversations with the owners of
several boat repair yards in Belém and the nearby port town of Icoaraçi confirmed that the type of fresh resin
obtained from Protium and closely related Burseraceae trees in the region is primarily used by commercial
boatyards for caulking the sides of wooden boats that sail in salt water. Boatyards in the Belém area that
service boats sailing in the Amazon River or brackish estuary used either a petroleum based product
(“betume”) or the amber like “breu pes” to caulk decks and sides. This may come from a hardened
Burseraceae resin, but the source of the material has not been identified. Even though these materials were a
little more expensive, the Belém repair people preferred to use them over the Protium derived resins because
the “branco” and “preto” types usually needed to be heated and filtered before they could be used due to their
abundant gritty impurities. One man also said that heating these resins over fire was more dangerous since
overflow could ignite and quickly flare up (Tembé did not mention such problems in their use of breu for
caulking canoes and wooden launches).
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Breu Resin Harvest Models
Data on the density and size structure of breu trees, resin harvest amounts and rates, and marketing
factors were used in three models to estimate the amount and value of resin that could be harvested in a given
area and time depending on the average density of trees.
The first model (Resin Harvest Model – Part 1:Table 3.10) estimates the amount of resin that would
be found on breu trees in a particular size class by multiplying the probability of finding resin on a tree of a
given size class by the average amount of resin found on trees of that size class. As was seen in the initial resin
harvest, the probability of finding resin on a tree and the amount of resin found on trees that had at least one
lump both increased with the size of the tree. These estimates of size class specific averages of resin per tree
range from just under 100 g per tree for trees in the 10-19 cm DBH size class to just over 1 kg per tree for trees
that are greater than 40 cm DBH.
The second model (Resin Harvest Model - Part 2: Table 3.11) predicts that as breu tree density
increases from 1 to 50 trees per ha, the amount of marketable resin that can be collected will vary from 0.3 to
16.5 kg per ha and the value to a harvester will range from $0.14 to $6.85 per ha. While breu tree density was
about 10 trees per ha at Tekohaw, this model covers the wider range of densities of potential resin yielding
Burseraceae trees in other parts of the Brazilian Amazon (Table 3.12). These area estimates of yield and value
are generated by first multiplying the percentage of each size class in a group of breu trees by the total density
of trees to estimate the number of trees of a particular size class that would be present at a specific density.
These estimates of the number of trees of each size class per ha would then be multiplied by the estimated
amount of resin per tree at that size class (derived from the first model) to predict the amount of resin that
would be found in a hectare of forest from the trees in each size class (Figure 3.7). The model predicts that the
two middle size classes (20 – 29 and 30 – 39 cm DBH) would yield the most resin because they are relatively
common and have relatively high probabilities of offering large amounts of resin. Although they are
numerous, smaller trees would contribute less because they rarely have much resin. Larger trees are almost
certain to offer substantial amounts of resin, but they would contribute relatively little to overall harvest
because they are relatively rare.
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The expected amounts of resin from each size class are then totaled to present the expected amount of
resin per ha at breu tree densities ranging from 1 to 50 trees per ha. Freshly harvested resin amounts were then
multiplied by 0.83 to account for the lost weight that occurs during resin drying (Table 3.11) to predict the
amount of marketable resin that could be obtained per ha of forest. Values of resin per ha were obtained by
multiplying these dried resin amounts by the $ 0.50 per kg price harvesters are typically paid for this product.
Values would be proportionally higher if they were based on the higher prices involved in retail level
transactions.
The third model (Resin Harvest Model - Part 3 (Table 3.13; Figure 3.8) predicts that a harvester could
gather 1.6 to 16.8 kg of marketable (dried) resin per day worth $0.78 to $8.37 per day when the density of breu
trees ranges from 1 to 50 trees per ha. These estimates are first based on a constant search rate of 200 m2 per
minute and a four minute per tree harvesting time. At high densities a harvester would be able to collect resin
from a large number of trees each day because they will not need to spend as much time searching for trees as
when the density is low. Resin harvest amounts are based on the predicted average resin yield per ha at
different densities presented in Table 3.11.
The second part of this model (Table 3.13) projects resin yields (fresh and dried), the number of days
it would take one person to completely harvest a 1000 ha block of forest and the potential revenue available in
this area at different densities of breu trees. This is an area of forest that is easily accessible within an hour’s
walk of a village. At the lowest density (1 tree/ha), an area this size could provide 279 kg of marketable resin
worth $140, but all of the available resin could be harvested in six months. At the highest density (50 trees/ha),
harvesters could procure almost 14,000 kg of marketable resin worth almost $7000, and it would take one
person about 30 months to complete a first round harvest of all resin bearing trees.
The studies of resin rates of accumulation show that breu trees will probably need at least four years
between harvests for resin to rebuild to first harvest levels. The final part of the third Resin Harvesting Model
predicts that with a four year resin regeneration cycle, some 1752 ha of forest would be needed to sustain one
person engaged in a full-time pursuit of breu resin collection even at the 50 breu tree per ha density. This
amount would be over 8000 ha if the density was as low as 1 tree per ha. These areas provide baseline
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estimates that could help assess the viability of ongoing resin harvesting in an area or development of
management guidelines.
DISCUSSION
RESIN HARVEST AMOUNTS PER TREE AND AREA OF FOREST
Breu Tree Density Inventories
Inventories in the Brazilian Amazon region show that trees from the Burseraceae family are often the
most species rich and numerous with importance values that frequently rank it among the top four families.
While Burseraceae trees are well represented in the forests around Tekohaw, the estimated density of 10 breu
trees per hectare seems low in comparison to inventories in other parts of the region where Burseraceae tree
density often exceeded 50 trees per ha (see references for Table 3.12). The percentage of those species and
trees that would possibly yield harvestable resin is about 50%. Forests in the eastern Brazilian Amazon
seemed to have a higher percentage of such trees than forests in the western part of the region where generally
non-harvestable species such as Tetragastris altissima were prevalent.
These inventories could ideally be used as a basis for predicting amounts of harvestable resin on a
regional level, but it may be problematic to assume that high densities of particular species found in 1 ha
inventories would be maintained over a landscape scale. Inventories of 100 ha plots in Guyana showed that
Protium trees were present in 18 out of 23 forest types spread throughout the country (ter Steege and
Zondervan, 2000). When present, the density of this genus varied from 3 to 217 trees per 100 ha with an
average density of 0.53 trees per ha.
Resin Harvest by Tree Type
Most species of Burseraceae produce resin in the twigs and bark, (Gentry, 1993), but even among the
35 species of Protium genus trees in the eastern Amazon, only some of these produce harvestable quantifies of
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resin (Daly, 1987; Balée and Daly, 1990; Balée, 1994). This study confirmed the importance of P. giganteum
and P. pallidum as major resin producing species that Balée (1994) found were key commercial sources of
resin for the Ka’apor Indians who live in the neighboring Alto Turiaçu Indigenous Reserve. The Ka’apor also
collect resins from other species of Protium and Trattinnickia for use as an incense, medicine and flammable
material, but it is not known what quantities of these materials are harvested or sold.
At the forest site near the town of Mojú, I found resin accumulation on P. giganteum (classified by
local residents as “breu branco”), P. paniculatum Engl., P. guianense (Aubl.) Marchand, P. pilosum (Cuatrec.)
Daly (locally called “breu preto”), Tetragastris altissima (Aubl.) Swart, and Trattinnickia rhoifolia Willd.
(locally called “breu siquereireua” or “sucuruba”). None of the trees identified as P. krukoffii Swart, P.
pernavatum Cuatrec., Trattinnickia lawrencei Swart var. bolivianum, or Trattinnickia burserifolia Mart. at
Tekohaw or Mojú had any resin on the trunk.
According to literature about Burseraceae species and resin production, the taxa most likely to yield
harvestable amounts of resin are Protium species in the Section Icica Aublet, Section Icicopsis (Engl.)Swart,
Section Sarcoprotium Daly, and Protium trifoliolatum group (Daly, 1987, 1989, 1992; Harley and Daly, 1995)
and select other species in Protium and other genera.
While many Burseraceae species have been considered a source of harvested resin (Appendix 3-A),
the only estimate of the amount of resin that can be harvested from Burseraceae trees without human
intervention is given in an economic valuation of nontimber forest products in Ecuador (Grimes et al., 1994).
This study indicated that 32 P. fibriatum Swart, P. nodulosum Swart, and P. sagotianum Marchand trees in two
upland and one alluvial plot could provide an average of 730 g of resin per tree per year. The authors
cautioned that this amount was based on ethnobotanical surveys with Indian collectors, not on actual yield
studies, and that even experienced collectors did not agree on which species provided harvestable resin.
Resin Harvest by Tree Condition, Size and Location
Tree characteristics associated with resin production in these tropical angiosperm Burseraceae trees
correspond well to findings in temperate conifer research. The starkest difference in resin amounts between
two groups of trees in this study was between live and dead trees. This is consistent with a lodgepole pine
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(Pinus contorta Dougl.) study where diseased trees had higher resin flow than healthy ones (Nebeker et al.,
1995). While Sternocoelus weevils’ preferential attacks of weaker trees may further weaken these trees over
time, they do not appear to quickly kill their host tree as some temperate wood-boring beetles do. Among live
trees at Tekohaw, larger (presumably older) trees were more likely to be attacked by weevils than smaller
(younger) ones, and when they were, they produced more resin per tree (Table 3.3). The positive correlation
between tree size and resin flow paralleled results in studies with lodgepole and Scots pine (Pinus sylvestris L.)
(Lieutier et al., 1993; Nebeker et al., 1995). Sternocoelus preference for attacking a specific height section of
the host tree’s bole was also found in two studies involving the southern pine beetle (Dendroctonus frontalis
Zimm.)(Coleoptera: Scolytidae) and its loblolly pine (Pinus taeda L.) host (Coulson et al., 1976; Fargo et al.,
1978). Differences in resin composition and flow rate have been documented at different heights in loblolly
pine, but these differences have not yet been linked to height preferences in beetle attack distribution (Schmitt
et al., 1988;Tisdale and Nebeker, 1992). No observations were made of the Sternocoelus weevils in flight, but
their flying capabilities are presumed to be very modest (O’Brien, personal communication 2000).
Accordingly their tendency to colonize lower sections of the trunk may be due to this limitation and an
attraction for ovipositing in protected crevices around tree buttress and stilt roots.
Resin harvest amounts were higher at Tekohaw in “baixo” (sometimes seasonally flooded) forest than
in more upland “terra firme” type forest. Average tree size in the “baixo” forest type, however, was larger than
in the drier areas. It may be that weevil attacks and resin flow were greater in these areas simply because
larger trees were found there.
The Role of Weevils in the Formation of Breu Resin Lumps
Bore Hole Patterns and their Implications about the Breu Tree Resin Defense System
Living close to and possibly ingesting resin may give Sternocoelus larvae certain benefits such as
physical protection from outside predators, chemical protection against microbes that often attack larvae
(Evans, 1975), and a source of special materials for pheromones (Slansky, 1992). In spite of these potential
benefits, though, it is unlikely that the weevils would want to maximize resin flow while feeding since coping
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with this viscous and terpene rich material would impose unavoidable energy costs. A weevil would,
therefore, logically try to maximize nutritive tissue intake while minimizing energy expenditures chewing
wood and dealing with fresh resin. The number and size of bore holes and larval development rate, therefore,
reflect the interaction between the weevil feeding strategy and the breu resin defense system. One question
that arises is whether or not resin production varies more with the type and condition of the tree or with the
intensity of the attack. The larger average size of trees might explain why some trees would be attacked more
readily than others, but it does not sufficiently explain why resin lumps are bigger on some trees than others.
One interpretation of the greater resin harvests from larger trees is that they are more vulnerable to
weevil attacks than smaller trees. The contrary may be true if larger trees produce more resin because weevils
need to feed more extensively on them to gain a comparable amount of nutrition. Cross-sections of resin
lumps show distinct layers so they must have been formed by numerous feeding and resin flow events. The
advantage of chewing the same hole wider and deeper is that chewing energy is presumably only expended in
nutrient rich phloem (Coulson and Witter, 1984). One study with Norway (Picea abies (L.)Kar.) and Sitka
spruce (P. sitchensis (Bong.) Carr.) trees even showed that the nutritive value of inner bark tissue actually
increased in response to wounding (Wainhouse et al., 1998). The downside of staying in the same hole is that
resin flow also grows with increases in bore hole size and depth (Table 3.5). It seems that weevils must be able
to gauge the benefit-cost ratio for chewing deeper into a given hole. Once this ratio drops below some
minimum value it seems better to start a new hole even though this means incurring “start-up” costs chewing
through 5 mm of a new section of non-nutritive outer bark that may have its own set of potent structural and
chemical defenses.
In addition to variation in bore hole number and size between trees, these differences also exist for
resin lumps on the same tree. This may be due to microsite differences in resin structures on the trunk. Breu
resin system anatomy has not been closely examined, but studies of a few coniferous trees provide possible
explanations. Some conifers have both radial (horizontal) resin ducts in the phloem or cambium and vertical
resin ducts in the xylem. The density and size of these ducts can vary seasonally and from place to place on
individual trees (Schroeder, 1990; Blanche et al., 1992; Lieutier et al., 1992; Tisdale and Nebeker, 1992; Baier,
1996). Resin viscosity, exudation pressure, or crystallization rate may also vary within and between trees
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(Baier, 1996). While existing resin ducts (the constitutive defense system) can deliver some preformed resin to
a wound site, the wounding process may induce the formation of new resin ducts or additional resin flow in the
original ones (Schroeder, 1990; Popp et al., 1991; Lieutier et al., 1992, 1995; Baier, 1996; Ruel et al., 1998;
Tomlin et al., 1998; Christiansen et al., 1999; Phillips and Croteau, 1999; Nagy et al., 2000). Boring a hole
into a site with an especially high amount or objectionable quality of resin could, therefore, prompt a
Sternocoelus weevil to start a new hole sooner rather than later.
Several conifer studies have further shown that wounding mostly induces formation of vertical resin
ducts and build-up of resin in a vertical orientation (Blanche et al., 1992; Lieutier et al., 1992; Tisdale and
Nebeker., 1992). While it can take six weeks for resin production to become fully functional in newly formed
vertical resin ducts (Blanche et al., 1992), induced resin may be more potent than the constitutive type since it
can have more numerous and higher concentrations of chemicals that may deter or kill insect attackers and
associated fungi (Lieutier et al., 1995; Phillips and Croteau, 1999; Franceschi et al., 2000; Nagy et al., 2000).
These factors may explain why at least one type of bark beetle chews new galleries in its host tree in a
horizontal direction (Lieutier et al., 1992). By moving laterally rather than vertically it may be able to keep in
front of induced resin defenses. Since Sternocoelus weevil bore holes are also often oriented in a horizontal
direction, breu trees may share some resin system features with temperate conifers. Considering these interspecific, inter-tree, and intra-tree variations in resin production, it is not surprising that resin flow is more
closely related to the number of bore holes and sum of bore hole diameters than to larvae size parameters
(Table 3.5).
The occasional finding of weevil larvae in resin lumps not accompanied by bore holes bears some
scrutiny. Without a bore hole it seems hard to imagine how the larvae could access inner bark tissue. It also
raises the question about how the resin came to be formed on that spot. The first issue suggests that larvae are
able to gain some nutritional value directly from the resin. No analysis has yet been done, however, to indicate
whether the liquid that exudes from wound sites contains some nutritional sap in addition to the documented
oleo-resin constituents. In the case of wounded trees, however, it seems possible larvae could feed on bark
tissues or microbes in the sap without making a bore hole. This feeding habit has been well established for the
sub-family of fly larvae found on these and other tropical trees discussed in Chapter 4. The observation that
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one chain saw wounded T. rhoifolia tree at the Mojú site had fewer bore holes per resin lump than unwounded
trees demonstrated this possibility.
Weevil Attack Pattern and Life-History
This study showed that usually only one weevil larvae occupied a resin lump with a single chamber.
Since most trees visited in the second round of harvesting only had several resin lumps per tree, it appears that
female weevils only oviposited one or at most several eggs at one site. Results also showed that the
distribution of attack on host trees by Sternocoelus weevils was not like several pine beetles that depend on
mass attack to successfully colonize (and ultimately kill) a host tree (Raffa and Berryman, 1987; Phillips and
Croteau, 1999). The observation that fresh resin lumps on “sarara” trees often form near other fresh or
abandoned ones does show that a new weevil may gain some advantage by being near the place where another
weevil is or was active. This idea is reinforced by the finding that even trees that had all resin (and presumably
all larvae) removed during the initial harvest were recolonized in proportion to the extent they had been
colonized before (Table 3.8). The extreme example of this was one tree that had 28 resin lumps removed from
it in the first harvest, but eight months later it had already accumulated another 49 resin lumps.
The distribution of weevil larvae head widths indicated two to three peaks. While a beetle body
grows steadily throughout the larval stage, the sclerotized head capsule is fixed at the beginning of the molt.
Within a population there is often a consistent progression of sizes of these hard parts that corresponds to each
new instar (O’Brien personal communication 2000). If this is true for this species of Sternocoelus, it
undergoes two or three instars during its larval stage. Three instars is a typical number for many weevils
(Evans, 1975).
The finding that adult weevils were only found on rotting portions of damaged or dead breu trees has
potentially important implications for the population dynamics of this species. If adult weevils do need dead
host trees for their stage of the life cycle, this requirement could limit the expansion of the Sternocoelus weevil
population if its dispersal range is restricted by poor flying ability. The major forest fire that swept through
hundreds of hectares around Tekohaw in the early 1980s burned but did not immediately kill several breu trees
found in this study. It seems possible that this fire created more potential adult weevil host trees than would
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exist under normal rates of tree mortality. As the forest recovers, the number of suitable host trees for adults
could decline.
Experimental Drilling
This study’s drilling experiment showed that one time drilling of potential resin producing trees
yielded no more than miniscule amounts of resin. Throughout the overall study, resin was sometimes seen in
machete cuts made on breu trees by passing Indians, but there was never much resin in these wounds and it
was always dry. These observations match those of a tree wounding study with other types of Burseraceae
trees in Panama where machete cuts were quickly clogged with resin (Guariguata and Gilbert, 1996). Since
weevil generated holes at Tekohaw produced an average of 12.5 g resin per hole (n=246 holes) compared to
less than 1 g from the drilled holes, the weevils apparently induce greater resin flows by repeatedly chewing
into the breu tree. This is consistent with the observation of layers in resin lumps. Whether weevil stimulation
of resin flow is due to mechanical wounding alone or is supplemented by another agent such as a fungus
remains to be seen.
Martinez-Habibe (1998) showed that Ticuna Indians in Columbia also harvest resin from various
species of Protium that have apparently been formed by weevil attacks. Accounts that other Amazonian
natives slashed breu trees with a machete and later collected resin (Mors and Rizzini, 1966; Spruce, 1970;
Daly, 1987), however, indicate that weevils may not be necessary to provoke resin flow in some types of breu
trees. Neels (2000) showed that indigenous people in Guatemala successfully harvest resin from Protium copal
by scraping out a series of 2 cm wide circles in the bark and reopening the wounds every three days for several
months. She found that these trees yielded an average total of 246 g per tree after being wounded once a week
for 14 weeks. Resin flow increased every week for the first five weeks and then leveled off producing an
average of 17.5 g per tree per collection. Since resin collection is done on a seasonal basis in the region, it was
hard to predict if these yields could be maintained throughout the year. While resin collection in the region is
generally done on a seasonal basis. There are many other examples of tropical and temperate trees that yield
commercial quantities of resin when they repeatedly wounded (Howes, 1949; Langenheim, in press).
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THE RATE OF BREU RESIN ACCUMULATION
Resin Yields in Follow-up Harvests
The harvest history of breu trees was a stronger predictor of the amount of the next harvest than
knowledge of its type, size, and site characteristics. This was shown by strong associations between the
number of resin lumps and resin amount in the initial and follow-up harvest of the same trees (Table 3.8).
These relationships are good evidence that the extent to which a tree has been colonized by weevils before will
strongly influence the extent to which it will be colonized again. There are three possible explanations for
these connections. One is that a tree that was suitable for infection once is simply more likely to be considered
suitable as a host tree again to adult weevils searching for a tree to colonize. The second possibility is that
infected trees with or without current weevil infestation release more resin volatiles that attract beetles in
search of ovipositing sites than ones that have never been infected. The third possibility is that harvesting did
not remove all weevil larvae and apparent recolonization was in fact merely a continuation of the growth of
larvae that escaped the first harvest.
The low resin per tree and resin per lump values in trees harvested for a third time in this study
present a new challenge to explain. Some characteristics of the trees in this group fit well with an expectation
of high resin, while others would indicate the opposite. Four months is not a long time to expect high levels of
colonization and resin build-up, but other factors may be operating. One possibility is that two rounds of
harvesting these specific trees and intensive harvesting in the area may have removed all larvae from the target
trees and possibly reduced the weevil population size in the area. While insect numbers are sometimes thought
to be almost limitless, it is not inconceivable that an insect highly specialized on a few species of trees in a
species rich forest could become vulnerable to intensive harvesting of its larvae.
The other possible explanation for the third harvest’s low resin yield is that seasonality affects resin
production. Several studies with temperate conifers have shown that resin production and susceptibility to
beetle attacks can vary between seasons and years depending on rainfall levels (Blanche et al., 1992; Lorio et
al., 1995; Baier, 1996). Trees suffering from severe drought or other stresses may become more vulnerable to
insects or pathogens. On the other hand, trees may actually develop more resistance to attacks in regular dry
112
periods because they can devote more carbon to making defensive compounds than in wetter times when
resources may be preferentially devoted to growth and reproductive effort (Dunn and Lorio, 1992; Tisdale and
Nebeker, 1992; Lorio et al., 1995; Nebeker et al., 1995; Baier, 1996; Feeney et al., 1998). This principle may
be a factor here since the strong follow-up resin harvest in November 1998 was preceded by a five month dry
season period after the initial harvest in June and July. The weak third round harvest in July 1999 was
preceded by the last four months of the rainy season following the second harvest in March.
Resin Lump Growth Rate and Weevil Development Time
The resin lump growth study could not precisely measure the amount of time that Sternocoelus
weevils spend in the larval stage, but data suggest a minimum of one year and possibly as many as three years.
These weevils have evolved many mechanisms to cope with a noxious resin, but they are probably not immune
to its deterrent properties. Even highly specialized pine beetles that use resin elements to make pheromones
can succumb to host tree resin defenses (Borden, 1982; Schroeder, 1990; Dunn and Lorio, 1992; Hui and
Zhimo, 1995; Phillips and Croteau, 1999). There is wide variation in the generation time of beetles that may
result from the quality of their food resources or periods of dormancy (diapauses) induced by lapses in food
availability (Evans, 1975). Some bark beetles in the genus Dendroctonus can complete 4 to 7 generations in
one season (Stark, 1982) while other beetles may take five years or more to complete their life cycle (Evans,
1975).
Breu Resin Accumulation over Time and the Impact of Harvesting
Estimating the sustainable harvest rate of a plant-based non-timber forest product generally requires
knowing how long one needs to wait to reharvest a product after the previous round of harvesting and what
limits may need to be imposed to avoid negative impacts on the plant population’s reproductive success
(Peters, 1994). In the case of breu resin, this task is more complex since resin formation is connected to both
the trees’ capacity to produce the material and weevil actions that stimulate it. Harvesting has a potentially
greater effect on Sternocoelus weevils than the breu trees themselves because weevil larvae are usually taken
along with the resin.
113
This study’s finding that weevil development is very slow has clear implications for the management
of breu resin harvest. Systematic harvesting of fresh resin with live larvae can set the clock back on resin
production for a considerable amount of time. Repeated harvesting in one area could reduce the Sternocoelus
weevil population and the amount of resin that could be harvested. While these weevils prefer to colonize
older and weakened breu trees, it is not known if their attacks accelerate their decline. It would be interesting
to know if a moderate level of resin harvesting (as generally practiced by the Tembé) actually helps breu trees
remain vigorous longer than if weevils were allowed to colonize trees to their full natural extent.
Other Animals that Utilize Resin Lumps
The only other animals found in fresh resin breu lumps beside weevils were larvae of a syrphid fly.
Larvae of this family have been previously found in tree exudates in both tropical and temperate regions
(Thompson, 1969; Ferrar, 1987), but there is only one case where fly activity directly contributed to resin flow
(Hellrigl, 1992)(See Chapter 4 for details).
My observation that stingless bees collected resin from various types of breu trees in the forest at
Tekohaw and Mojú was also noticed by Neels (2000) in her study of P. copal resin harvest in Guatemala. She
estimated that these bees (possibly Trigona spp.) removed as much as 25% of the resin that had oozed onto the
trunk one week after the trees were wounded in resin harvest experiments. Stingless bee collection of resin
from a variety of plants has been noted throughout the tropics (Roubik, 1989; Nogueiro-Neto, 1997). Apart
from this study, reduvid assassin bugs have also been seen using resin to facilitate their capture of bees and
other insects in Brazil and other parts of Latin America (Adis, 1984; Hogue, 1993).
Some wild mammals also use resin from Burseraceae trees. In Panama, coatis (Nasua narica L.)
groom themselves with the aromatic resin of Trattinnickia aspera (Swart), possibly for its pharmaceutical
properties (Gompper and Hoylman, 1993). Wild peccaries regularly chewed resin masses on Protium copal
trees in Guatemala (Neels, 2000).
114
Should Sternocoelus Weevils be Considered Beneficial Insects?
Weevils are probably the world’s most diverse taxonomic family of organisms with more than 50,000
identified species (Hogue, 1993; Anderson, 1995), but the habits of most are scarcely known. Some weevils
and other bark beetles (mostly Scolytidae) have achieved notoriety by colonizing and sometimes killing
coniferous and hardwood trees in spite of these trees’ defensive systems of sticky and toxic resins (Storer et al.,
1997; Ruel et al., 1998; Tomlin et al., 1998; Phillips and Croteau, 1999). Various weevils are also considered
pests on tropical crops including bananas, rice, coffee, cotton, tea, sugar cane, yams, and palm trees (Lamb,
1974). With the possible exception of an aquatic weevil tested for controlling Eurasian water milfoil (Creed
and Sheldon, 1995; Sheldon and Creed, 1995), the term beneficial insect has rarely been applied to weevils or
other close herbivorous relatives.
The term beneficial insect is usually reserved for insects that provide pollination services, produce
honey, recycle nutrients, prey on pest insects or serve as a direct source of human food, medicine or dye
material (Metcalf, 1993). An insect or pathogen that causes economic damage to human agricultural or tree
crops is understandably called a pest. The beneficial label, however, should be extended to phytophagous
insects whose attacks stimulate production of plant compounds that are in turn used by people, wildlife and
other insects including key pollinators as long as such attacks do not significantly hasten the mortality of the
host plants..
The Sternocoelus weevil should firstly be considered a beneficial insect because it is responsible for
directly generating a significant economic resource for many forest peoples. Secondly, this weevil is providing
an indirect benefit to the forest community by stimulating resin collected by stingless bees that pollinate a
variety of rainforest trees (Roubik, 1989; Nogueiro-Neto, 1997). Since stingless bees are common pollinators
of Protium trees (Daly, 1987), weevil attacks that provoke resin flow may actually provide some benefit to the
offended trees. Provoking resin flow may also provide an essential material for the larvae of the syrphid flies
that live in the fresh resin of some breu trees and probably also pollinate various rainforest plants (Thompson,
1969; Ferrar, 1987).
115
BREU RESIN HARVESTING ECONOMICS AND MARKET
Breu Resin Harvesting Rates
The amount of time required to find breu trees with resin in a species-rich forest depends on the skill
of the searcher and the density of such trees in the forest. It is difficult to quantify the search efficiency of a
“typical” harvester because some individuals had a much stronger intuitive sense of where to find such trees
than others. The seasoned harvesters seemed to have a finely tuned search image for breu trees that helped
them pick out trees that might have resin at a much greater distance than their less experienced colleagues.
When a harvester found a breu tree of the type that might have resin, he first stopped to examine the
trunk and stilt roots for resin stuck to the bark and look around the ground for chunks of dried resin that have
fallen from the tree. If resin was found, additional time was spent removing resin lumps from the tree by hand
or with a machete. If resin was seen higher up on the trunk, a harvester sometimes cut a long sapling to
dislodge the elevated resin lumps. Since time measurements of searching and harvesting were made with a
research team, the accuracy of the estimates of these average times could be improved by accompanying actual
lone harvesters at work.
Resin Drying Studies and the Effect of Stingless Bees
The loss of weight when harvested breu resin is laid out in the sun is potentially due to three factors:
loss of volatile components, evaporation, and removal by stingless bees. Various essential oils have been
identified in the resin of several Protium and Tetragastris species (Zoghbi et al., 1993; 1998; Siani et al., 1999a,
1999b) so the release of aromatic substances from fresh resin is to be expected. Water soluble carbohydrates
(gums) have been found mixed with resins in other Burseraceae genera (principally Boswellia and
Commiphora)(Khalid, 1983), but studies of the Protium have apparently not investigated its resin for materials
that could evaporate.
Since conventional outdoor drying methods lead to a 15 - 20% weight loss in harvested resin,
determining the reasons for these losses could have financial implications for the harvesters. The greater
weight loss of resin samples in partially screened enclosures exposed to bee removal than samples in fully
116
screened enclosures showed that bee resin removal was apparently responsible for just as much weight loss as
chemical processes. Even if bee removal was the main cause for the difference in weight loss in the resin
drying enclosure experiment, it cannot be assumed that bee removal rates from four 36 g samples of resin
could be extrapolated to hundreds of kilograms of resin laid out to dry in a single village. Further evidence of
large-scale bee removal would be needed before any alterations in drying methods would be justified on
economic grounds. Allowing stingless bees full access to harvester resin is certainly providing an important
material to one group of the forest’s most ubiquitous group of pollinators.
Breu Resin Sales
It appeared that the total breu resin harvest in Tekohaw and other indigenous communities in the
Gurupi region is several metric tons per year. These communities have access to tens of thousands of hectares
of relatively undisturbed forest so the current scale of harvesting is probably sustainable at the landscape level.
Forests near villages and areas close to river access points have probably been harvested most often, so there
may be local areas where Sternocoelus weevil populations have been depleted and profitable harvesting will
not be possible without a hiatus of many years to permit populations to rebuild. The Tembé and Ka’apor
Indians readily admit they need to spend hours in the forest in more remote areas in order to make a
commercial breu resin collecting effort worthwhile since the final product only gives them $0.50/kg. The resin
has a long tradition as a medicine and great utility in caulking canoes, but the market demand for it in these
uses is limited. The resin’s principal area of supply seems to be in the wet forests of the eastern Amazon, but
the fact that boat repair yards in Belém prefer alternatives that are more expensive than Protium derived resin
shows that the prospects of more sales to the boat repair market in that area are not good. It is, thus, the lack of
demand and consequent low price that limits the amount of harvesting effort more than the limit of the
resource.
The price paid for breu resin as a raw material for caulking boats in the eastern Brazilian Amazon is
much lower than the $US 9.11 per kg paid for resin from various Protium species in Ecuador (Grimes et al.,
1994). Several groups of Quichua Indians have long used this resin as a ceramic glaze (Whitten, 1976;
117
Whitten and Whitten, 1988), and the collection and sale of the resin for this purpose has made it the most
important economic product for the Quijos Quichua in the Napo Region (Grimes et al., 1994).
While resins from various species of Burseraceae have long been used by indigenous people and other
forest dwellers as an incense and folk medicine, it seems ironic that the major commercial use of breu resin to
caulk wooden boats requires driving off most of the aromatic essential oils. A phytochemist at the Museu
Goeldi in Belém is now analyzing different types of breu resin and other Amazonian plants for their possible
uses as scents in the perfume industry (Zoghbi, personal communication 1999). If this screening process is
successful, it could increase demand for the resin. This demand might not be sufficient to increase the price as
well, but it could at least allow more people to generate modest income from collecting this resin.
Breu Resin Harvest Models
The first two Resin Harvest Models use data on the size class distribution of breu trees at Tekohaw
and the average amounts of resin that were found on each size class. While the format of these models may
have general value in predicting the amount of resin that could be harvested in other locations, the estimations
would need to be adjusted to reflect differences in size class structure and the probability of finding resin on
trees in a certain size class. Observations made at the Mojú site indicated that rates of infection by weevil
larvae there were considerably lower than found at Tekohaw. Neels (2000) estimated that the annual yield of
resin from P. copal trees in Guatemala with a bark wounding harvest method would be about 2.4 kg per ha
where the density of productive trees averaged 12.8 trees per ha. This is only slightly less than the 3.3 kg per
ha yield of fresh resin that was predicted for an initial harvest at Tekohaw with an estimated density of 10
potentially productive trees per ha.
The third Resin Harvest Model indicates that a harvester can only expect to consistently earn the
Brazilian minimum wage ($R 7.00 per day) by collecting breu resin if he is working in a forest that has at least
25 breu trees per ha that potentially yield harvestable resin. This is based on spending five hours on harvesting
activities per day and probably includes several extra hours walking to and from a village or camping spot to a
prime collecting area. Some Tembé collectors stated they have filled several sacks of breu resin weighing 20
118
kg each in a day’s hard work, but such events seem to be unusual days of good fortune for the most
experienced harvesters and does not reflect the average yield of harvesters in general.
The most sobering result of the third Resin Harvest Model is that incorporating a four to five year
hiatus on harvesting the same tree requires that thousands of hectares of forest would need to be available to
sustain even a few harvesters on a full-time basis. Since the Tembé and many other forest dwellers rely more
on subsistence than commercial activities to support their lifestyle, breu resin harvesting offers a modest but
welcome source of cash income to dozens of people rather than full-time paid work for a few. The forests
could provide more resin than is currently being harvested, but this surplus seems destined to remain intact
unless the demand for the material substantially increased.
CONCLUSIONS
Sustainable harvest of a plant-based non-timber forest product typically depends on managing the
harvest within the limits of an individual plant’s capacity to produce the harvested product, its tolerance of the
harvesting method, and the capacity of the plant’s population to withstand the harvesting intensity (Peters,
1994). In this case where commercial quantities of breu resin are only produced when trees are attacked by a
specialized bark-boring weevil, sustainable harvest of the product will also require maintenance of the weevil
population. Routine resin harvesting removes many weevil larvae so intensive harvesting could eventually
reduce local weevil populations to the point where breu tree colonization and resin formation decline.
Resin harvesting, though, may actually help maintain the health of breu trees and prolong their resin
producing lives. Individual weevil attacks do not kill their host trees, but the accumulation of weevil larvae
and their bore holes may increase the exposure of the trees to bacterial and fungal pathogens. Over time
weevil attacks could hasten the demise of the tree. While this study shows that resin harvesting does not stop
recolonization, it may slow down the deterioration of the tree associated with weevil attacks.
Some important evolutionary and ecological questions remain to be answered in future studies. How
widespread is the association between the Sternocoelus or other weevils and Burseraceae trees in other parts of
the Amazon? Other researchers have noted the relationship between unidentified weevils and resin producing
Burseraceae trees in Amazonas states of both Brazil and Columbia (Zoghbi, personal communication 1998;
119
Martinez-Habibe, 1998). Further inquiries about weevil life history and the specificity of its relationships with
host trees throughout the region could make an important contribution to the evolution of weevil-host plant
associations (Anderson, 1993). There is nothing known yet concerning the role that any fungi or other microbe
might play in stimulating resin flow or weevil adaptation to its resin-rich environment. It is clear that the
weevils’ provocation of resin flow provides a material that is important to many forest dwellers in their every
day lives and in commerce. It would be also be valuable to explore the extent to which these resins are critical
materials for other invertebrates such as syrphid flies and stingless bees that provide pollination services to the
forest. The breu trees might not agree, but from a human perspective this new species of Sternocoelus, the
“resin” weevil, might arguably be considered a new sort of beneficial insect.
Given the numerous factors that restrict breu resin harvest potential within close proximity of the
major villages near the Gurupi River, it is not surprising that most people who collect this resin for sale go
upriver to reach larger areas of intact forest. This opens up a much larger area of forest to exploitation, but
there are still limits to this. Collectors generally do not travel farther than several hours into the forest away
from the river and a large portion of the reserve has been occupied and/or deforested by non-Indian colonists,
ranchers and loggers (Sales, 1993, 1994). In theory, overharvesting has the potential to reduce long-term
harvesting potential by reducing the population of weevils that stimulate resin flow. One procedure that could
avoid this possibility would be to restrict resin harvest to dried lumps that no longer contain living larvae. The
non-systematic manner in which harvesters search the forest, however, probably provides a strong buffer
against severe impacts on the weevils. Harvesters have a strong tendency to seek out medium and large trees
that are most likely to yield large amounts of resin. This practice undoubtedly leaves many smaller trees with
some weevils on them as a de facto beetle population reserve.
While depletion of the weevil population may not be an immediate concern, the increasing demand
for cash income by the Tembé could reduce the amount of breu resin available in the most accessible areas of
forest. Even with healthy beetle populations, this study predicts that it will take at least four years for resin
levels to build up to initial harvest levels. The current relatively low market price and demand for resin may
provide a sufficient barrier to overharvesting. Success in developing the resin as a new source of fragrances
for the perfume industry could expand the market and economic benefits for resin collectors; it could also push
120
local harvesting into the non-sustainable realm. If harvests increase, it would probably be advisable for
communities to institute a more proactive form of managing resin harvests. One scheme would be to make
areas off-limits to harvesting for a certain number of years after an intensive harvest. Another management
tool that could be explored is transplanting small resin lumps from trees with multiple lumps to suitable
uncolonized trees. This technique of “seeding” a tree with an insect that produces a commercial product has
worked well with lac insects responsible for producing natural shellac (Sharma et al., 1999).
In conclusion this study has demonstrated that the harvest potential of this NTFP that is considered to
be a plant defensive compound is directly influenced by the host tree’s relationship to a herbivorous weevil.
There are no doubt many other useful resins and defense compounds produced by plants in response to attacks
by insects or microbial pathogens. Investigating these relationships could address many outstanding
coevolutionary questions and help develop strategies to sustainably manage such resources.
121
TABLE 3.1 SUMMARY OF RESEARCH OBJECTIVES AND METHODS
OBJECTIVE
Density of Breu Trees
& Size Class Structure
Amount of Resin on
Breu Trees during
Initial Harvest
Sternocoelus Weevils
and their Role in Breu
Resin Yield
Resin Yield by
Drilling Breu Trees
Rates of Resin
Accumulation on Breu
Trees
SURVEY/METHOD
Belt transect inventories: 7.3
ha inventoried in 10x500 m
plots in 1/98 and 11/99 in
forest 1 km from Tekohaw
Initial Harvest Survey: Resin
was harvested from all breu
trees in two month (6-7/98)
survey of forest <4 km from
Tekohaw
Initial Harvest Survey
Breu Tree Drilling
Experiment: 18 trees drilled
with bore hole dimensions
similar to weevils (11/99)
Follow-up Harvests: subsets
of trees were reharvested in
11/98, 3/99, and 7/99
Resin Lump Growth Study: In
3/99, new resin lumps were
tagged on 16 breu trees last
harvested in 6-7/98.
Follow-up Harvests & Resin
Lump Growth Study
Resin Weight Loss
during Outside Drying
Resin Weight Loss
due to Stingless Bees
Percentage of Breu
Trees with Resin
Amount and Value of
Resin Sales
Resin Amount and
Value per Hectare
Resin Amount and
Value per Time
Harvested
Resin Drying Study: 16
batches observed (7/98)
Resin Drying in Open and
Closed Enclosures (4/99)
Initial Harvest Survey: In 6/98
all breu trees found in forest
were checked for resin
Interviews with Resin Buyers
Initial Harvest Survey, Resin
Drying Study, Interviews
Initial Harvest Survey, Resin
Drying Study, Interviews
MEASUREMENTS
Tree Type (branco or sarara
only); Tree Size (DBH ≥10
cm); Forest Type (terra firme
or baixo)
Tree Type; Tree DBH; Tree
Status (Dead or Alive); Forest
Type; GPS reading; Number
resin lumps; weight of resin
lumps;
Length, width, weight of
weevil larvae, pupae & adults
ANALYSIS
Trees/ha by Tree type, Tree
DBH, Forest Type
RESULTS
Table 3.2
Comparison of Resin
Amount/tree to Tree Type,
Tree DBH, Tree Status; Forest
Type, and Distance from
Tekohaw
Dimensions of weevils and
head width distribution pattern
Table 3.3;
Fig. 3.1
Table 3.4,
Fig. 3.2
Number and size (width) of
weevils in resin lump; number
and size (width, depth) of
weevil bore holes behind resin
lump; weight of resin lump
Number and size (width) of
weevils in resin lump; number
and size (width, depth) of
bore holes behind resin lump
Tree DBH; Presence or
absence of resin in drill holes
one month after drilling
Correlation of resin lump
weight to weevil number and
dimensions and to bore hole
number and dimensions
Table 3.5
Correlation of weevil number
and dimensions to bore hole
number and dimensions
Table 3.6
Percentage of holes with resin;
comparison of Tree DBH to
percentage with resin
Results in
text
Number and weight of resin
lumps (all trees)
Comparison of number of
resin lumps and resin weight
to time since previous harvest
Resin lump area to weight
ratio measured from harvested
lumps. Daily resin growth
rate (g/day) estimated for
tagged resin lumps
Correlation between resin
lump number & weight at first
& second harvest and DBH
Regression Model Estimates
for Resin Accumulation
Percentage resin weight loss
during outside drying
Resin weight loss with and
without bee removal
Probability of resin by Tree
Size
Table 3.7
Average price per kg of dried
resin
Amount and value of resin
harvest per hectare of forest
Amount and value of resin
harvest per day and renewable
harvesting cycle
Results in
text
Table 3.11
Area of individual resin lumps
on tagged trees measured in
3/99 & 7/99. Area and weight
of resin lumps from 5 trees in
Follow-up Harvests
Number of resin lumps and
weight per tree; Tree DBH;
Time since previous harvest
Resin batches weighed daily
until stabilized
Resin sample weights at end
of each day
Tree Size (DBH); Presence or
absence of resin on trunk
Amount of resin bought and
price paid
Same as previously described
Same as previously described
Fig. 3.4
Table 3.8
Table 3.9
Fig. 3.5
Fig. 3.6
Table 3.10
Table 3.12
122
TABLE 3.2 DENSITY OF BREU TREES IN DIFFERENT FOREST TYPES AT
TEKOHAW
Area
Branco Tree
Sarara Tree
Total Tree
Surveyed
Number and
Number and
Number and
FOREST
(ha)
Density
Density
Density
TYPE
No. (%)
No. (%)
Tr./ha
No. (%)
Tr./ha
No. (%)
Tr./ha
6.25
33
5.3
24
3.8
57
9.1
Terra Firme
(86%)
(94%)
(69%)
(81%)
1.05
2
1.9
11
10.5
13
12.4
Baixo
(14%)
(6%)
(31%)
(19%)
7.30
35
4.8
35
4.8
70
9.6
Total
Combined Results of Two Inventories of Resin Yielding Breu Tree Types (≥ 10 cm DBH) in Tekohaw
Survey Blocks
123
TABLE 3.3 BREU RESIN HARVEST PER TREE BY STATUS, TREE TYPE, LIVE
TREE SIZE (DBH), FOREST TYPE, AND DISTANCE FROM TEKOHAW
CATEGORY
OVERALL
STATUS
Live
Dead
TREE TYPE
Branco
Sarara
LIVE TREE DBH
5 - 9 cm
10-19 cm
20-29 cm
30-39 cm
40-49 cm
50-69 cm
FOREST TYPE
Terra Firme
Baixo
DISTANCE
1.0-1.9 km
2.0-2.9 km
3.0-4.2 km
Number of
Trees
183
Mean Tree
DBH(cm)
28.4
Resin Harvest (g)
Mean ± S.E.
856 ± 100
Resin Harvest (g)
Median
420
171
12
28.3
29.2
789 ± 85a
1808 ± 923b
380
930
72
111
24.5
30.8
646 ± 135 a
992 ± 139 a
325 a
508 a
2
33
71
46
14
5
7.2
16.3
25.1
34.9
44.5
54.9
38 ± 28 a
437 ± 133 a
766 ± 145 a
1009 ± 176 a
905 ± 228 a
1394 ± 507 a
38 a
200 a
340 a
480 a
640 a
1120 a
110
71
25.6
32.8
650 ± 98 a
1167 ± 204b
315 a
540b
53
29
99
25.8
27.0
30.2
669 ± 109 a
756 ± 127 a
992 ± 171 a
320 a
540 a
422 a
All categories include live and dead trees with resin except DBH size classes. Sample sizes in some categories are different
due to missing data from some trees in that category. Distance is the distance between the tree and Tekohaw village.
Different letters in mean column indicate that pairwise p-values in AOV comparisons of means were ≥ 0.05. Different
letters in Median column indicate that p-values of pairwise comparisons were ≤ 0.05 with Mann-Whitney Test and Mood’s
Median Test had a p-value ≤ 0.05 in comparing all medians for that category.
124
TABLE 3.4 DIMENSIONS OF WEEVILS FOUND IN BREU RESIN LUMPS
Larvae
Head Width (mm)
Body Width (mm)
Body Length (mm)
Pupae1
Head Width (mm)
Body Width (mm)
Body Length (mm)
Body Weight (g)
New Adult2
Pronotum Len. (mm)
Pronotum Wid. (mm)
Elytra Length (mm)
Elytra Width (mm)
Body Length (mm)
Body Weight (g)
N
Mean ± S.E.
160
158
158
1.84±.04
3.58±.11
8.15±.27
2
2
2
2
1.80±.25
4.85±.69
9.55±2.35
0.12±.06
3
3
3
3
3
3
3.67±.19
4.93±.12
8.47±.38
6.68±.22
11.82±.23
0.22±.01
Measures done on specimens stored in alcohol. Linear dimensions done with calipers examining specimens under
dissecting microscope and measured to nearest 0.01 mm. Weight measurements done with electronic scale to nearest
0.0001 g.
1. Includes one pupa and one animal in transition between pupa and adult condition.
2. Includes two males and one female. Female was larger in all dimensions.
125
TABLE 3.5 BREU RESIN LUMPS WEIGHT CORRELATION TO WEEVIL
LARVAE AND BORE HOLE DIMENSIONS
WEEVIL LARVAE
(per resin lump with larvae)
Number of Larvae
Head Width Sum (mm)
Head Width Maximum (mm)
Body Width Sum (mm)
Body Width Maximum (mm)
BORE HOLES
(per lump with bore holes)
Number of Bore Holes
Diameter Maximum (mm)
Diameter Sum (mm)
Depth Sum (mm)
Volume Sum (mm3)
Correlations with Resin Lump
Weight
No.
Corr.
Lumps
Coef.a
R2 (% )b
137
.456**
20.8**
136
.352**
12.4**
136
.261**
6.8**
129
.456**
20.8**
129
.346**
12.0**
245
244
244
33
33
.540**
.355**
.488**
.632**
.671**
a - Pearson’s Correlation Coefficient ρ (rho)
b - Regression Analysis R2 Value expressed as percentage
**- Correlation and Regression Values significant at p ≤ 0.01
29.2**
12.6**
23.8**
40.0**
45.0**
126
TABLE 3.6 WEEVIL LARVAE DIMENSION CORRELATION TO BORE HOLE
NUMBER AND DIMENSIONS
Bore-Hole Number
Bore-Hole Dia. Sum
Corr.Cf.a
R2 (%)b
Corr.Cf. a
R2 (%)b
WEEVIL LARVAE
Number of Larvae
.311**
9.7**
.336**
11.3**
n=129 resin lumps
Sum of Head Widths
.444**
19.7**
.512**
26.2**
(mm) n=127 resin lumps
Maximum Head Width
.425**
18.1**
.500**
25.0**
(mm) n=127 resin lumps
Sum of Body Widths
.583**
33.9**
.634**
40.2**
(mm) n=120 resin lumps
Maximum Body Width
.535**
28.6**
.616**
37.9**
(mm) n=120 resin lumps
a - Pearson’s Correlation Coefficient
b - Regression Analysis R2 Value expressed as percentage.
**- Correlation and Regression Values significant at p ≤ 0.01.
Bore-Hole Dia. Max.
Corr.Cf. a
R2 (%)b
.133
1.8
.330**
10.9**
.544**
29.6**
.401**
16.0**
.589**
34.7**
127
TABLE 3.7 RESIN LUMP NUMBER AND WEIGHT FOUND ON BREU TREES
DURING FOLLOW-UP HARVESTS
Second Harvest Month
Time Since Last Harvest
N trees
Ave. Resin/tree (g)
Ave. Lump/tree
Ave. Resin/lump (g)
Nov. 1998
139 - 147 Days
11
134.0 ± 50.6
3.3 ± 0.9
40.9 ± 6.9
Mar. 1999
241 - 272 Days
26
164.9 ± 49.5
5.2 ± 1.9
31.8 ± 2.7
July 1999
364 - 388 Days
12
57.5 ± 32.4
2.3 ± 0.6
25.3 ± 6.9
128
TABLE 3.8 CORRELATION OF TREE SIZE AND HARVEST HISTORY WITH
FOLLOW-UP BREU RESIN HARVEST
Tree DBH (cm)
Resin (g)/Tree in
Second Harvest
Resin Lumps/Tree
in Second Harvest
Corr. Cf.a
.407**
R2 (%)b
16.6**
.440**
19.4**
Resin (g)/Tree in
First Harvest
Corr. Cf.a
R2 (%)b
.548**
30.0**
.476**
22.7**
Resin Lumps/Tree in
First Harvest
Corr. Cf.a
R2 (%)b
.716**
51.3**
.802**
a - Pearson’s Correlation Coefficient; b - Regression Analysis R2 Value expressed as percentage
**Correlation and Regression Values significant at p ≤ 0.01
Analysis includes 49 trees with resin and 17 trees without resin during follow-up harvest.
64.3**
129
TABLE 3.9 PROJECTION OF BREU RESIN YIELD AFTER INITIAL HARVEST
All Trees
Trees with Resin
First
Harvest
Resin (g)
per Tree1
562±65
789±85
Year 1
Time of Second Harvest
and Estimated Resin Yield (g) per Tree (Mean ± S.E.)
Year 2
Year 3
Year 4
Year 5
Year 6
153±12
215±14
278±23
390±28
403±35
566±42
528±46
741±56
653±58
916±71
777±69
1091±85
Year 7
902±81
1266±99
Regression Model:
Resin Wt/tree in Second Harvest = 28.4 + 0.000381*WtH1*Days + 0.000017*DBH2 *Days + 0.0443*Lumps*Days
R2=32.6; p=0.000
WtH1=Resin weight collected from tree during initial harvest; Lumps=Number of resin lumps collected from tree during
initial harvest; DBH2 = square of DBH (cm) at 1.5 m.; Days=Number of days after initial harvest
Based on Regression Model from Second Round Resin Harvests and Resin Lump Growth Study (n=82 trees) applied to All
Live Trees (n=240) and Live Trees with Resin (n=171) Monitored in Initial Harvest
Note: 1) Mean ± S.E.
130
TABLE 3.10 BREU RESIN HARVEST MODEL - PART 1. ESTIMATED
AVERAGE RESIN PER TREE BY SIZE CLASS
BREU TREE
SIZE CLASS
10-19 cm DBH
20-29 cm DBH
30-39 cm DBH
>40 cm DBH
PROBABILITY OF
RESIN
0.20
0.62
0.80
1.00
AVE. RESIN PER
TREE FOR TREES
WITH RESIN (Kg)
0.44
0.77
1.01
1.03
EST. RESIN PER
TREE FOR ALL
TREES (Kg)
0.09
0.47
0.81
1.03
131
TABLE 3.11 BREU RESIN HARVEST MODEL - PART 2. DENSITY OF RESIN
YIELDING TREES, RESIN HARVEST AND HARVEST PER VALUE PER
HECTARE
ESTIMATED BREU TREES PER SIZE CLASS AT
DIFFERENT DENSITIES (Trees/ha)
BREU TREE
SIZE CLASS
10-19 cm DBH
SIZE CLASS
PERCENT
0.55
1 Tree/ha
5 Trees/ha
10 Trees/ha
25 Trees/ha
50 Trees/ha
.55
2.75
5.5
13.75
27.50
20-29 cm DBH
0.26
.26
1.30
2.6
6.50
13.00
30-39 cm DBH
0.14
.14
0.70
1.4
3.50
7.00
>40 cm DBH
0.05
.05
0.25
0.5
1.25
2.50
10-19 cm DBH
AVE. RESIN
(Kg/tree)
0.09
20-29 cm DBH
0.47
0.12
0.59
1.18
2.94
5.88
30-39 cm DBH
0.81
0.11
0.57
1.13
2.83
5.65
>40 cm DBH
1.03
0.05
0.26
0.52
1.29
2.59
0.34
1.65
3.30
8.26
16.51
0.28
1.37
2.74
6.85
13.71
0.14
0.69
1.37
3.43
6.85
AREA FRESH
RESIN (Kg /ha)
AREA DRIED
RESIN (Kg/ha)
AREA VALUE
($R/ha)
ESTIMATED BREU RESIN AMOUNT AND VALUE PER HA
AT DIFFERENT DENSITIES (Kg Resin/ha)
0.05
0.24
0.48
1.20
2.40
Equations for Variables
1) Kg Resin per Size Class at Given Density = Resin per Tree at Size Class (Table 3.9) x Number of Trees in Size Class at
Given Density = Kg Resin per Size Class at given Density
2) Area Fresh Resin = Total Kg Fresh Resin at Given Density for All Size Classes Combined = Kg Fresh Resin/ha
3) Area Dried Resin = Kg Fresh Resin/ha x 83% = Kg Dried Resin/ha
4) Area Value (for harvester) = Kg Dried Resin/ha x $R 0.50/Kg = $R/ha
132
TABLE 3.12 DENSITY OF BURSERACEAE RESIN YIELDING SPECIES IN
BRAZILIAN AMAZON INVENTORIES
Ha
SURVEYED
REGION
EAST
(Maranhão,
4
Pará, and Amapa
1
1
4
1
42
1
1
1
6
(3)
(3)
CENTRAL
(Manaus, AM) 70
WEST
(Rondônia)
1
1
0.52
2
6
1
NUMBER OF SPECIES
DENSITY(trees/ha)
Ref
All Species
Resin Species1 All Species
Resin Species1
20
12
80.5
57.8
1
10
7
17
6
9
2
6
4
7
4
3
7
5
10
3
7
1
2
1
3
1
2
72.0
68.0
41.5
56.0
12.3
11.0
35.0
7.0
64.3
52.0
46.0
30.8
30.0
10.8
10.0
5.0
1.0
0.0
1.1
2
3
4
5
6
7
8
9
10
11
48
21
6
1
1
5
8
1
4
1
1
1
2
0
12
70.0
5.0
4.0
25.0
20.8
60.0
9.0
5.0
4.0
1.5
0.7
0.0
13
14
15
16
17
18
Site References: Balée, 1994 (1,5); Balée, 1986(1.1); da Silva and Rosa, 1989(2, 6); Almeida et al., 1993(3); Dantas et al.,
1980(4); da Silva et al., 1987(7); Salomão et al., 1988(8); Salomão, 1991(9); Lopes, 1993 (10); Mori et al., 1989(11);
Rankin-de-Mérona et al., 1992(12); Salomão and Lisboa, 1988(13); Maciel and Lisboa, 1989(14); Lisboa, 1989 (15,18);
Absy et al., 1987(16,17).
Other Notes:
1. Includes species known to be a source of harvestable resin or close relatives of these (Daly, 1987; Harley and Daly,
1995): Protium altsonii, P. carnosum, P. crassipetalum, P. decandrum, P. giganteum, P. guianense, P. heptaphyllum, P.
morii, P. opacum, P. pallidum, P. paniculatum, P. pilosum, P. polybotryum, P. robustum, P. rubrum, P. sagotianum, P.
tenuifolium, P. trifoliolatum, Tetragastris panamensis, Crepidospermum goudotianum.
2. Secondary forest site; others are assumed to be old-growth forest
3. Non-plot survey
133
TABLE 3.13 BREU RESIN HARVEST - PART 3. RESIN HARVESTING RATES
AND VALUE BY AREA AND TIME
DENSITY (Trees/ha)
10
25
9
6
SEARCH AND HARVEST (Min./Tree)
1
54
5
14
TREE HARVEST RATE (Trees/Day)
5.6
21.4
33.3
50.0
60.0
AREA HARVEST RATE (Ha/Day)
5.6
4.3
3.3
2.0
1.2
RESIN HARVEST RATE (Kg/Day)
1.9
7.2
11.2
16.8
20.2
DRIED RESIN YIELD (Kg/Day)
1.6
6.0
9.3
14.0
16.8
VALUE OF EFFORT ($/Day)
0.78
3.00
4.65
7.00
8.37
AMOUNT, VALUE, TIME PER
HARVEST IN 1000 ha
Fresh Resin (Kg)
336
1681
3363
8406
16813
Dried Resin (Kg)
279
1395
2791
6977
13955
Collecting Days (5 hours per day)
180
233
300
500
833
Value($R)
140
698
1395
3489
6977
8111
6257
4867
2920
1752
10139
7821
6083
3650
2190
AREA NEEDED FOR ONE PERSON
DAILY HARVEST (ha)
4 Year Harvest Cycle
5 Year Harvest Cycle
50
5
Equations for Variables
Area Search Rate = 1 min./ 200 m2 x 10,000 m2/ha = 50 Min./ha
Tree Search Rate = 50 Min./ha x 1 ha/Number of Trees = y Min./Tree
Trees Harvest Time = 4 min./tree
Tree Harvest Rate = 5 hrs. (300 min.)/y min./tree = Trees Harvested/Day
Area Harvest Rate = Trees Harvested/Day x 1 ha/Number of Trees = Area Harvested/Day
Resin Harvest Rate = Resin Harvested/ha (Table 3.10) x Area Harvested/Day = Kg Resin Harvested/Day
Dried Resin Yield = Kg Resin Harvested/Day x 83% = Kg Dry Resin Harvested/Day
Value of Effort = Kg Dry Resin Harvested/Day x $R 0.50/Kg = $ Earned/Day ($R ≈ $US in 1998)
Area Needed for Continuous Daily Harvest = Area Harvest Rate x No. Years between Harvests from Same
Trees
RESIN HARVEST PER TREE (g)
134
10000
FITTED LINE REGRESSION PLOT
R-Sq = 1.3%; p=0.124;
N=181 Trees with Resin
5000
2-3 Km.
756 g./tree
1-2 Km.
669 g./tree
3-4 Km.
992 g./tree
0
1000
2000
3000
4000
DISTANCE FROM TEKOHAW VILLAGE (m)
FIGURE 3.1 RELATIONSHIP OF DISTANCE FROM VILLAGE TO BREU RESIN
HARVEST
PERCENT IN EACH WIDTH CLASS
135
14
12
10
8
6
4
2
0
1
2
3
HEAD CAPSULE WIDTH (mm.)
(n = 160 larvae from Tekohaw)
FIGURE 3.2 DISTRIBUTION OF WEEVIL LARVAE HEAD CAPSULE WIDTHS
FROM BREU RESIN LUMPS
136
a. Sternocoelus weevil larva feeding on breu tree in bore hole adjacent to chamber in resin lump
b. Alipumilio fly larvae in resin lump
c. Trigona stingless bee collecting fresh resin
d. Reduvid assassin bug stalking bees and other insects attracted to fresh resin
FIGURE 3.3 ILLUSTRATION OF BREU RESIN LUMP AND ASSOCIATED
INSECTS
137
Growth rates based on estimated weight differences
in resin lumps measured at 9 and 12 months after
initial resin harvest. N=53 Resin Lumps on 16 Trees.
0.9
Estimated Resin Lump
Growth Rate (g/day)
0.8
0.7
Fitted Line Regression Plot
(Quadratic Model)
R-Sq = 59.7 %
0.6
0.5
0.4
0.3
0.2
0.1
0.0
0
100
200
300
Estimated Resin Lump Weight (g) at
12 Months after Initial Resin Harvest
FIGURE 3.4 ESTIMATED GROWTH RATE OF INDIVIDUAL BREU RESIN
LUMPS AT TEKOHAW
CUMULATIVE WEIGHT LOSS (%)
138
Interval Plot: Mean plus 95% Confidence Interval
18
13
8
1
2
3
4
5
DAYS EXPOSED OUTSIDE
(N = 16 Batches of Resin)
FIGURE 3.5 BREU RESIN WEIGHT LOSS IN OPEN OUTSIDE DRYING AT
TEKOHAW
139
CUMULATIVE WEIGHT LOSS (%)
Interval Plot: Mean plus 95% Confidence Interval
Closed
to Bees
14
Open
to Bees
9
4
Day 1
Day 3
Day 4
DAYS IN SCREENED ENCLOSURES
N= 4 Samples Breu "Sarara" per Treatment
FIGURE 3.6 BREU RESIN WEIGHT LOSS WITH AND WITHOUT STINGLESS
BEES
140
BREU RESIN PER
HECTARE (kg)
15
DBH SIZE
CLASS (cm)
Mean Contribution of Resin per
Hectare by each DBH Size Class Based on Resin Harvest and Size
Class Population Data at Tekohaw
> 39
30-39
20-29
10
10-19
5
0
1
5
10
25
50
BREU TREES PER HECTARE
FIGURE 3.7 ESTIMATED FIRST TIME RESIN HARVEST AT DIFFERENT
DENSITIES OF BREU TREES
141
DRIED RESIN (Kg) /// REVENUE ($R)
Based on Resin Harvest Amounts and Collection Rates at Tekohaw
KG DRIED BREU RESIN
HARVESTED PER DAY
15.00
12.50
10.00
7.50
$R EARNED PER DAY
HARVESTING RESIN
5.00
2.50
0
0
10
20
30
40
50
BREU TREES PER HECTARE
FIGURE 3.8 ESTIMATED DAILY HARVEST AND SALE VALUE OF DRIED
RESIN AT DIFFERENT DENSITIES OF BREU TREES
142
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150
APPENDIX 3-A SCIENTIFIC SPECIES AND COMMON NAMES OF RESIN
YIELDING SPECIES OF NEOTROPICAL BURSERACEAE
TAXA
REGION/GROUP
COMMON NAMES
PROTIUM
P. altsonii Sandw. (includes syn. P.
paraense)
Venezuela
catamajaca, chipoi-yek
Guyana
Surinam
Brazil (Amapa)
Brazil (Amazonas)
Brazil (Ka’apor)
tsepur
tiengi-monnie
breu mescla
breu branco
ara-kanei’y, kandeiape’y,
kanei’y pitag
hykata’ywching
breu grande
icica
hykata’ywpihun
icica
caraña
caraña
icica
ma-mee-ree’-ma
caraño
breu, breu vermelho
hikeuteu ‘y pihun,
kanei’y pitag, yawar-’a’y
hykata’ywra
elemi, tacamahaca
shillquillo
kirreri
encens blanc, encens gris
breu branco, breu manga,
breu preto, breu vermelho
breu branco, kandeiaka’y
hikeuteu ‘yw chig,
kandei-y tuwyr
haiwa
icica
caraña, copal, elemi,
sasafras
hee-ta-ma-ka
o-mo-ta’
anime, anime blanco, brea,
caraña, guacamayo,
guacharaco, incienso,
pergamin, tacamahaco
currucai, tacamahaco
haiowa, sipipio, shipu,
almecega, icica
breu branco do campo
ara-kanei’y, hikeuteu’y,
breu branco verdadeiro,
kanei’y pitag, yawar-’a-’y
P. apiculatum Swart
P. aracouchini (Aubl.)March
P. brasiliense (Spreng.)Engl.
P. carana Marchand*
P. crassipetalum Cuatrec.
P. decandrum (Aubl.)Marchand
P. elemifera*
P. fibriatum Swart
P. giganteum Engl.
P. giganteum Engl. var. crassifolium
P. giganteum var. giganteum
Brazil (Tembé)
Brazil
Brazil
Brazil (Tembé)
Brazil
Columbia
Venezuela
Brazil
Amazon (Kuripako)
Columbia (Amazon)
Brazil
Brazil (Ka’apor)
Brazil (Tembé)
Columbia
Ecuador (Quijos Quichua)
Columbia (Amaz/Piapoco)
Fr. Guiana
Brazil (Pará)
Brazil (Ka’apor)
P. heptaphyllum (Aubl.) Marchand ssp.
heptaphyllum (includes P. tacamahaca)
Guyana (Arawak)
Brazil
Panama
Amazon (Tanimuka)
Amazon (Kubeo)
Columbia
Venezuela
Guyana (Arawak,Carib,Warrau)
Brazil
Brazil (Ka’apor)
REF.
8
8
8
8
8
1,8,3
2
13
12
2
12
7
18
12
18
19
13
3, 1
2
7
5
19
8
8
3,8,1
10
12
6
18
18
6,19,
18,7
18, 6
12
16, 9,
13,12
3
151
Appendix 3-A (Continued)
TAXA
P. icicariba (DC.) Marchand
P. llanorum Cuatrec.
P. macrophyllum H.B.K.
P. nervosum Cuatrec.
P. niloi Pires
P. nodulosum Swart
P. opacum Swart ssp.opacum
P. pallidum Cuatrec.
P. paniculatum Engl. var. paniculatum
P. pilosum (Cuatrec.)Daly
P. polybotryum (Turcz.)Engl. ssp.
polybotryum
REGION/GROUP
Venezuela
Brazil
Venezuela
Columbia (Amazon)
Columbia
Columbia
Brazil (Tembé)
Ecuador (Quijos Quichua)
Brazil
Peru
Brazil (Ka’apor)
Brazil (Tembé)
Peru
Brazil
Fr. Guiana
Brazil (Pará)
Columbia (Amazon)
Brazil (Ka’apor)
Brazil (Tembé)
Fr.Guiana
Fr.Guiana (Paramaka)
Surinam
P. puncticulatum J.F. Macbr.
P. robustum (Swart) D. Porter
P. sagotianum Marchand (includes syn.
P. insigne)
P. spruceanum (Benth.)Engl.
P. tenuifolium Engl.
P. trifoliolatum Engl.
P. unifoliolatum Spruce ex Engl.
Surinam (Carib)
Brazil (Pará)
Brazil (Ka’apor)
Peru
Peru
Brazil
Brazil (Ka’apor)
Brazil (Tembé)
Columbia
Ecuador (Quijos Quichua)
Venezuela
Columbia (Amazon)
Brazil (Ka’apor)
Brazil (Tembé)
Brazil
Brazil (Ka’apor)
Brazil (Tembé)
Brazil (Tembé)
Brazil (Yanomami)
COMMON NAMES
guacamayo
Brazilian elemi,
almecega, jauaricica,
brea branca, breu branco
tacamahaco
hugucal, incienso
anime, cunday,
guacharaco hediondo
anime
kiriwa’yw
shillquillo
breu preto
breu branco,
hikeuteu ‘yw chig,
kanei-’y tuwyr,
yawara-’a’cu’y
hykata’ywching
copal, shininga arana
almecega
encens
breu branco
caraño
hikeuteu-iran-’yw
kanei-ape’y, yawar-’a-’y
kiriwa’yw
encens blanc, wet-sali
moni
rode bast tingimoni,
ajawabali, joelliballi
beleroe
peraka, tamoene poeleka
breu branco
akakandei’y, kandeiape’y
breu branco
kanei-aka’y,
yawara’a’cu’y
hykata’ywching
currucay, urrucay, caraño
shillquillo
cabimbo
incienso
kandei’y tuwyr,
kanei-’y tuwyr
hykata’ywching
breu, breu preto
kanei-ape’y, yawar-’a-’y
hykata’ywching
hykata’ywrang
warapé kohi
REF.
18
15, 9
18
19
19, 7
19
2
5
13
17
3,1
2
8
16
8
8
19
3, 1
2
8
8
8
8
8
8
17
17
13
1,3
2
7,21,19
5
18
19
8,1
2
13
1,3
2
2
14
152
Appendix 3-A (Continued)
TAXA
Protium spp.
REGION/GROUP
Central America
Ecuador
Columbia
Columbia (Amazon)
Columbia (Amaz/Kubeo)
Columbia (Amaz/Kurripaco, Yeral
Columbia (Andina)
Columbia (Caribe)
Venezuela
Guyana
Fr. Guiana
Brazil
Bolivia
TETRAGASTRIS
T. altissima (Aubl.)Swart
T. mucronata (Rusby) Swart
T. panamensis (Engl.)Kuntze
TRATTINNICKIA
T. aspera (Standl.)Swart
Peru
Columbia
Brazil
Brazil (Ka’apor)
Brazil (Tembé)
Columbia (Amazon)
Columbia (Amazon)
Columbia (Amaz/Piapoco)
Brazil
Brazil (Ka’apor)
Columbia (Amazon)
Columbia (Andina, Pacific/Cholo
Brazil
COMMON NAMES
caraño, copal, fontole,
fosforito, pom, tontol
anime blanco
anime, caraño, currucay
breo, copal, de-ep, tovake
tamamuri, tatamoco
taloake
kakaine, keterre, tabaiba
carano, incienso
indio desnudo
anime, azucarito, caraño
haiawa, kurokay, porokay
encens blanco, tinguimoni,
encens gris, encens rojo
almecega, aruru
caraño
guacamayo, tacamahaca
breu manga, sali
breu manga, waruwa’y
iwapepirangyw
copal
caraño
kirrei
barrotinho, breu preto,sali
breu preto, papara-’yg,
waruwa-ywa-pitag-’y
REF.
4
4
4
19
19
19
19
19
4
4
4
4
4
17
7
13
3
2
19
19
19
13
3
caraño, echicorai-guna
19
caraño, ampó
19
T. burserifolia Mart.
amescla,
13
breu sucuruba branco
Brazil (Ka’apor)
kyryhu-’y, kyryh’y-’y
1,3
T. rhoifolia Willd.
Brazil
breu sucuruba
13
Brazil (Ka’apor)
breu mescla, kyryhu’-y,
3
kyryh’y-’y
Brazil (Tembé)
kiriwa’yw
2
Trattinnickia spp.
Columbia (Andina)
caraño
19
Columbia (East Llanos)
caraño
19
Brazil
amesclao
4
BURSERACEAE (non-specified)
Peru
caraña, caraño, copal
8
Surinam
salie, tiengi-monnie
8
Brazil (west Amaz)
breu sicantá, sicantá
8
Brazil (central/south)
almecegueira, almesca,
8
almisca, almiscar,
amescla, icicariba
Bolivia
isígale, isigo
8
References: 1)Balée and Daly,1990; 2)Balée,1987; 3)Balée,1994; 4)Chichignoud et al.,1990; 5)Grimes et al., 1994;
6)Correa and Bernal,1989; 7)Cortes, n.s., and Renner et al.,1990; 8)Daly,1987; 9)Howes,1949; 10)Johnston and
Colquhoun,1996; 11) Levi-Strauss,1952; 12)Martini et al.,1993; 13)Milliken and Albert,1996; 14)Mors and Rizzini,1966;
15)Paula and Alves, 1997; 16)Pinedo-Vasquez et al.,1990; 17)Pittier, 1926; 18)Shultes and Raffauf, 1990; 19)Vega et al.,
1984. Notes: * - questionable species
CHAPTER 4
THE ASSOCIATION OF AN ALIPUMILIO FLY (DIPTERA: SYRPHIDAE) WITH
BURSERACEAE TREE RESINS IN THE EASTERN BRAZILIAN AMAZON
154
ABSTRACT
There are numerous species of flies (Diptera) whose larvae develop in plant saps or rotting exudates.
There are relatively few examples of immature flies that have adapted to resin flows of coniferous or
angiosperm trees. Most of these cases have been members of primitive syrphid genera (Syrphidae) including
Cheilosia in the temperate region and Alipumilio in the neotropics. A recent study of resin harvested by
Tembé Indians in the eastern Brazilian Amazon has revealed a potentially new species of Alipumilio that
develops in resin masses on various species of Burseraceae trees. These resin flows are primarily stimulated
by the actions of a previously undescribed species of bark-boring weevil in the genus Sternocoelus
(Coleoptera: Curculionidae). Resin lumps are sometimes also associated with severe physical tree wounds.
This fly larva’s morphological features, presence and movement in the resin mass, and unsuccessful
rearing apart from fresh resin indicate that this Alipumilio species may be consuming bacterial and fungal
spores as well as sap materials coming out of tree wounds. While Sternocoelus weevils are found in resin
lumps in a range of Protium and other Burseraceae tree species in the region, Alipumilio larvae were only
found in the resin of some of these species. The study speculates that its absence on some species may be due
to chemical properties of resin types that inhibit build-up of micro-organisms that nourish this kind of larvae.
If this distinction is borne out, the presence or absence of mycophagous syrphid larvae could serve as an initial
indicator of certain medicinal properties of resins in this diverse group of trees found throughout the Amazon.
INTRODUCTION
Almost 6,000 species of syrphid flies (Diptera: Syrphidae) commonly called flower or hover flies
have been described so far (Vockeroth and Thompson, 1987; Sommaggio, 1999). Adult syrphids are often
mimics of stinging Hymenoptera, and many are important pollinators (Vockeroth and Thompson, 1987;
Teskey, 1976; Speight, 1978; Ervik and Feil, 1997). Larvae of the family have diverse feeding patterns that
include mycophagous, phytophagous, saprophagous and entomophagous habits (Vockeroth and Thompson,
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1987; Ferrar, 1987; Rotheray and Gilbert, 1999). They are typically found under the bark, tree exudates,
decaying sapwood of living trees, dead rotting trees, or under fallen leaves. Their diet may include bacterial or
fungal spores, sap, cambial tissue, decaying wood, aphids or immature ants (Hartley, 1961; Teskey, 1976;
Perry and Stubbs, 1978; Gilbert, 1986; Ferrar, 1987; Foote, 1987; Vockeroth and Thompson, 1987; Rotheray
and Gilbert, 1999). There are well documented cases where beetles (Coleoptera) have overcome resin defenses
of coniferous trees (Raffa and Berryman, 1987; Phillips and Croteau, 1999), but apart from Cecidomyiidae gall
midges, few flies have apparently been able to circumvent the potent physical and chemical deterrents posed
by plant resins (Teskey, 1976; Pyenson, 1980; Coulson and Witter, 1984; Thompson, 1985).
Cladistic analysis disputes the division of Syrphidae into three monophyletic sub-families (Rotheray
and Gilbert, 1999), but closely related genera typically grouped in the primitive Eristalinae or Milesinae
provide the best examples of flies that have adapted to resin-based tree defenses (Rotheray and Gilbert, 1999;
Rotheray et al. 2000). Hellrigl (1992) described finding larvae of the spruce resin fly Cheilosia morio Zett. in
resin lumps on the trunks of spruce trees in northern Alps region of Italy. While these resin lumps resembled
ones made by attacks of the wood-boring beetle Dendroctonus micans Kug., observations indicated that the
flies were directly responsible for the resin flow in these trees. In Washington State, U.S.A., Burke (1905 cited
in Teskey, 1976 and Rotheray, 1990) observed that Cheilosia alaskensis Hunter fed on the sap and cambium of
hemlock (Tsuga heterophylla (Rafin.)Sarg.) trees by scraping cavities next to the sapwood. In the neotropics,
some larval specimens from the Alipumilio Shannon genus were reared in the pitch of Araucaria australis trees
in southern Brazil (Thompson, 1972). Larvae of Alipumilio femoratus Shannon were reared in the resinous
sap of a tree in the Ecuadorian Amazon (Rotheray et al., 2000; Whitten and Thompson, personal
communication). Only a dozen specimens of Alipumilio found from Argentina to Mexico have so far
accounted for nine species, so the diversity of this genus and examples of resin resistant lifestyles is probably
much greater than is reflected in its poor representation in collections and field research (Thompson, 1972).
A recent study of resin harvested from the trunk of various Burseraceae trees in the Alto Rio Guamá
Indigenous Reserve in Pará state, Brazil (Plowden et al., 2002) has revealed another and probably new species
of Alipumilio (Rotheray, personal communication 2001) whose larvae shares the unusual habit of developing
in tree resin. Similar larvae were found in another eastern Brazilian Amazon site near the Mojú River. The
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Burseraceae trees and the resin that exudes from bark wounds as a clear to milky white liquid (Daly, 1987;
Gentry, 1993) are most commonly known in the Brazilian Amazon as breu (Rodrigues, 1989). The aromatic
and sticky balsam resins of breu trees have been collected by forest dwellers throughout the Amazon for
subsistence and commercial purposes including incense, flavors, medicinal products, illumination, handicraft
glues, and caulking material for wooden boats (Mors and Rizzini, 1966; Reitz, 1981; Plowman, 1984; van den
Berg, 1984; Balée and Daly, 1990; Schultes and Raffauf, 1990; Daly, 1992; Lima et al., 1992; Balée, 1994;
Boom, 1996; Siani et al., 1999a). Stingless bees and other invertebrates also utilize the resin for various
purposes (Chapter 3).
The larvae of a previously undescribed species of bark-boring weevil Sternocoelus Kuschel (O’Brien,
personal communication 2000) are primarily responsible for the build-up of resin on at least a dozen species of
trees in the Burseraceae genera Protium Burman, Tetragastris Gaertner, and Trattinnickia Willd. at several sites
in the eastern Brazilian Amazon (Chapter 3). Like other phloem boring beetles, this weevil’s larva first uses its
mandibles to chew through the tree’s outer bark to access more nutrient rich inner bark. It makes simple
cylinder shaped bore holes in the cambium that average 0.7 cm deep and sever resin ducts interspersed in this
tissue. As resin drips out of the wound onto the trunk and begins to solidify, the larva burrows into the
malleable resin and shapes a chamber inside the resin lump. Fly larvae were also sometimes found in resin
lumps associated with weevil larvae. This report will present information about the natural history of this
Alipumilio resin fly and its potential use as an indicator of resin anti-microbial potency.
STUDY SITES
The main study area (Tekohaw site) was an area of about 500 ha of closed tropical rainforest near the
Tembé Indian village of Tekohaw (020 37.7’ S; 460 33.1’ W.). This village is located on the Gurupi River in
the Alto Rio Guamá Indigenous Reserve (eastern Pará state, Brazil). A second study area (Mojú site) in
eastern Pará was located in a similar forest on state road PA150 near the town of Mojú (020 11.5’ S; 480 49.1’
W.) Both forests are dominated by trees in the Lecythidaceae, Leguminosae, Sapotaceae, and Burseraceae
families (Baleé, unpublished data; Lockman, personal communication).
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MATERIAL AND METHODS
In June and July 1998, a team of Tembé Indian men located 250 breu trees within 5 km of the village
of Tekohaw. The tree’s common name as well as tree size, tree status (live, injured, dead), forest type (“terra
firme” and “baixo”) were recorded. Each tree was marked with a numbered aluminum tag, and leaves were
collected from a sample of trees to assist with identification. Measurements were made on the number and
weight of resin lumps during this initial and follow-up harvests in November 1998, March and July 1999. The
number of insect larvae found in resin lumps was measured in a sample of trees. Once the larvae were
distinguished as weevil and fly larvae, they were separated into these categories and counted separately.
Lengths, widths and weights of fly larvae stored in alcohol were later measured in the lab since precise
instruments were not available to do so in the field.). Type and paratype specimens will be deposited in Brazil.
A similar protocol was used to measure resin lumps and tree-insect relationships on trees at the Mojú
site in March 2000. Resin lumps were harvested from several species of unwounded Protium and Tetragastris
trees and one Trattinnickia rhoifolia Willd. tree that had a large amount of resin exuding from a chainsaw
wound. Leaf specimens were collected from a sample of Burseraceae trees at the site including all those that
had resin to identify host tree species of the weevil and fly.
Since only a small fraction of immature tropical syrphids have been identified (Thompson, 1987),
several attempts were made to raise the Alipumilio larvae. During the initial attempts, fly larvae in resin lumps
collected at the Tekohaw site were placed in mesh cages and jars with mesh tops. Fly larvae living in resin
collected from the Trattinnickia tree at the Mojú site were placed in glass vials with medium used for rearing
Drosophila fruit flies.
158
RESULTS
RESIN LUMPS AND HOST TREE PREFERENCES
The initial resin harvest at Tekohaw showed that dead trees had significantly larger amounts of resin
on them than live trees. Large trees were also colonized more often by Sternocoelus weevils than small trees
(Chapter 3). Leaf specimens identified by botanists at the Museu Goeldi in Belém revealed that at least eight
species of Protium including P. pallidum Cuatrec, P. trifoliatum Engl., P. decandrum (Aublet) Marchand, P.
morii Daly, P. giganteum Engl. var. giganteum, P. polybotryum (Turcz.) Engl., and P. glabrescens Swart. and
Tetragastris panamensis (Engl.) O Kuntze had resin lumps at the Tekohaw site. P. pallidum and P. trifoliatum
were generally classified as “breu branco” (white breu) by local Tembé while the other six species were
considered different varieties of “breu sarara.” While Sternocoelus weevil larvae were found in resin lumps in
both types of trees, Alipumilio fly larvae were only found in resin lumps on “sarara” type trees.
The Mojú site also had resin accumulation on P. giganteum as well as on P. paniculatum Engl.,
Tetragastris altissima (Aubl.) Swart., and Trattinnickia rhoifolia Willd. The last species was only encountered
with resin on the trunk of a tree wounded with a chainsaw. Weevil larvae were again found in resin lumps on
all of these types of trees. Fly larvae, however, were only found in the resin of the wounded Trattinnickia tree.
None of the trees identified as P. krukoffii Swart., P. guianense (Aublet) Marchand, P. pernavatum Cuatrec, P.
pilosum (Cuatrec) Daly, Trattinnickia lawrencei var. bolivianum Swart., or Trattinnickia burseraefolia Mart. at
either site had any resin on the trunk.
CHARACTER OF RESIN LUMPS
Resin associated with weevil attacks on breu trees was usually found in round lumps affixed to the
lower part of tree trunks although some lumps were occasionally seen much higher up. Lumps were often
wedged in the crook between the trunk and a buttress or stilt root. Lumps were typically 2 to 6 cm thick and
averaged 339 ± 59 g in the initial Tekohaw harvest. The weevil larva forms a cavity in the fresh resin that is 3
to 8 mm deep and extends almost to the inner edge of the resin lump (Figure 3.3). Fly larvae were found in
159
resin lumps both with and without weevil larvae at both sites. When resin lumps contained both kinds,
however, the flies were in a separate section of the lump that usually occupied no more than 25% of the lump.
In cases where flies were found in resin without weevils, some lumps had bore holes behind them indicating
prior weevil activity.
One distinctive feature of a resin mass containing fly larvae was that it had a dark color and taffy-like
consistency that stretched out when pulled apart. This was different from the cake-like quality of resin
associated with weevils that was sticky but readily broke into separate pieces. While weevil larvae were
usually found quiescent in a resin chamber or occasionally in a bore hole, maggots were almost always
observed moving in their section of resin. This activity that presumably accompanies feeding, therefore,
seemed responsible for the transformation of resin consistency. Dried resin lumps further revealed the
different feeding strategies of the immature resin weevil and fly. Resin lumps occupied solely by weevils had
distinct layers indicating different resin flow events. These layers logically resulted from successive boring in
the cambium during the weevil’s slow larval development. Resin sections that were inhabited by fly larvae,
however, dried in a well-mixed matrix interspersed with narrow meandering tunnels.
NUMBER, SIZE, AND IDENTITY OF FLY LARVAE
During the initial harvest at Tekohaw, the identity of the fly larvae was not known, so precise counts
are not available. Finding all fly larvae in resin lumps was also more difficult than locating immature weevils
since the maggots are immersed in a mass of sticky resin. During the second round of resin harvests at
Tekohaw, 9 out of 38 (24%) of resin lumps that had weevil or fly larvae in them had at least one fly larva in
them, and 6 of these (16%) had only fly larvae. This measure overestimates the number of resin lumps
associated only with flies since only 15 lumps out of 910 examined (1.6%) did not have weevil bore holes
behind them. In the chainsaw damaged tree at Mojú, 5 out of 22 (23%) resin lumps with larvae had only fly
larvae in them, but only 3 of these lumps had no weevil bore holes behind them.
At both sites, two “types” of fly larvae were found. Both types had similar mouth hooks, but one
“type” that had three pairs of spiracular slits was considerably smaller than the “type” that had numerous
spiracular slits radially oriented around the ecdysial scar (Table 4.1). Rotheray (personal communication
160
2001) believes that these two “types” of larvae probably represent Stage 2 and 3 instars of the same species of
Alipumilio. The average body width of the larvae changed from 1.2 to 3.0 mm and the average length went
from 5.1 to 7.7 mm in the shift from Stage 2 to Stage 3 larvae. The average wet weight during these stages
was 4.2 and 37.1 mg respectively, but these figures should probably be increased to account for weight loss
that occurs during storage in alcohol. Rotheray further noted that the form of the thorax and the head of this
larva is very similar to A. femoratus Shannon. The third stage larva has a number of interesting and novel
features, such as a very flattened anal segment, different arrangement of spiracular openings to A. femoratus
and has an almost complete band of spicules around the segments (Rotheray, personal communication 2001).
The number of Stage 3 larvae present in a single resin lump was rarely more than one at the Tekohaw
site. When Stage 2 larvae were present, however, there were usually several and as many as fifty in one lump.
This pattern was repeated at Mojú where there was rarely more than one third stage larva present in one resin
lump, but two to twenty-five second stage larvae were present in some lumps in the wounded Trattinnickia
tree.
None of the attempts to rear these breu resin flies was successful. Over a period of months, the resin
lumps collected at Tekohaw that were placed in cages and jars became covered with mold and eventually dried
out before any mature flies emerged. Fly larvae placed in vials with Drosophila medium also did not survive
to maturity (Overal, personal communication 2000).
DISCUSSION
The finding that Sternocoelus weevil attacks and resin amounts were highest in large and dead trees is
consistent with earlier research on temperate coniferous trees (Lieutier et al., 1993, Nebeker et al., 1995).
While the Alipumilio fly is sometimes capable of colonizing trees without these weevils, it seems that in most
cases this fly relies on the weevil to first bore into a tree before it can access a food source.
Figuring out the diet and feeding method of this Alipumilio fly during its immature stages is one of
the principal natural history questions raised by this investigation. The answers are central to determining if
the fly plays some direct role in provoking the flow of resin that is harvested by people. As noted, several
161
species of closely related Cheilosia larvae, feed directly on cambial tissues so they are not limited to feeding on
tree sap that is exuding due to other causes (Rotheray, 1990; Hellrigl, 1992; Rotheray and Gilbert, 1999). An
examination of the mouth parts of A. femoratus, however, did not produce conclusive evidence regarding the
diet of this species. It has well-developed mouth hooks and other characteristics that indicate it could feed on
bacteria, yeasts and other micro-organisms in its host sap or it might be a phytophage that feeds on cambial
tissues where the sap exudes (Rotheray et al. 2000). Its overall morphological characteristics place it in the
most basal position of the Syrphidae in proximity to the Eumerus and Merodon genera that are primarily
considered mycophagous (Rotheray and Gilbert, 1999; Rotheray et al., 2000). This study seems to affirm the
probability that Alipumilio does not feed directly on the cambium since it was exclusively found moving about
in resin masses. Their frequent movement in the resin is consistent with the filter-feeding mode of
mycophagous larvae living in semi-liquid mediums (Rotheray and Gilbert, 1999).
There have been several studies of the volatile components of Burseraceae resins (Zoghbi et al., 1993,
1994, 1995, 1998; Siani et al., 1999a, 1999b), but studies have not yet analyzed these materials for the
presence of microbes or nutrients that might come out of wounds along with the resin. The failure of the initial
attempts to rear these fly larvae on small lumps of resin removed from the tree or the Drosophila medium
suggest the maggots require a fresh source of organic material from the host tree in order to complete their
development. It is possible that bacteria or fungus attracted to the wound site or brought in by the weevils need
to build up to a certain level before these larvae can meet their nutritional needs. In order to successfully rear
several other types of immature Syrphid flies, Rotheray (personal communication 2001) needed to keep them
supplied with an ongoing supply of fresh exudate from their typical host tree.
The Alipumilio larva’s discriminating diet raises an interesting question about the variation in hosttree preferences between the weevil and the fly. Weevils attacked many but not all species of Burseraceae in
the same forest while Alipumilio were apparently more selective in their host preferences. The notable
difference was the presence of fly larvae at Tekohaw in “sarara” type resin lumps and its total absence in
“branco” type resin lumps at Tekohaw and Mojú. While “branco” and “sarara” type breu trees represented
various taxonomic species, “branco” type resin lumps all formed as white masses that slowly turned gray.
Fresh “sarara” type resin also dripped out of the tree as a milky white liquid, but its resin lumps quickly
162
darkened and were almost black when dry. “Sarara” lumps were also covered with unidentified gritty and
small plant-like material.
These visible differences between “branco” and “sarara” type resin are likely due to differences in
their chemical composition. Several studies have found significant variation in the volatile components of
resins from various Protium species (Zoghbi et al., 1993, 1994, 1995, 1998; Siani et al., 1999a, 1999b) that
may impart their resins with specific medicinal properties. P. heptaphyllum (Aublet) Marchand is one type of
“branco” breu tree whose resin is used by native peoples in Northeastern Brazil as a treatment for infectious
skin diseases, although extracted essential oils did not show great effectiveness against several dermatophytes
in one test (Lima et al., 1992). Another pharmacological test did demonstrate its potency as an antiinflammatory agent (Siani et al. 1999a).
A comparison of the chemistry and microbial communities of various types of breu resin could help
explain whether Alipumilio larvae are not found in the resin of certain host trees because it has physical or
chemical properties that are inhospitable to them or because the resin simply lacks sufficient microbes to
nourish them. It is interesting to note that “branco” resin sometimes earns a higher market price than the
“sarara” type so it has some medicinal or other properties that has made it more attractive to human consumers
than flies.
Sommaggio (1999) has suggested that the large diversity and differences in environmental
requirements in syrphid flies make them well suited as bioindicators. As this case demonstrates, the presence
or absence of one probable mycophagous syrphid larvae in the resin of a group of closely related trees could be
a first-level indicator of the anti-microbial potency of each tree species’ defense compounds. Other
mycophagous larvae might serve a similar purpose in analogous situations.
TABLE 4.1 BODY DIMENSIONS OF IMMATURE ALIPUMILIO FLIES FROM
BREU RESIN AT TEKOHAW
Dimension
Body width (mm)
Body length (mm)
Spir.Plate width (mm)1
Weight (mg)2
Stage 2 Larvae (n = 9)
Maximum
Mean ± S.E.
1.53
1.21 ± 0.09
6.80
5.12 ± 0.40
0.47
0.39 ± 0.02
8.10
4.24 ± 0.81
1 - Posterior Spiracular Plate
2 - Wet weight of larvae preserved in 70% alcohol.
Stage 3 Larvae (n=49)
Maximum
Mean ± S.E.
4.10
2.98 ± 0.12
12.74
7.66 ± 0.27
1.40
1.05 ± 0.04
100.70
37.05 ± 3.29
164
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Ervik, F. and J.P. Feil. 1997. Reproductive biology of the monoecious understory palm Prestoea schultzeana in
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Hellrigl, Von K. 1992. Die fichtenharzfliege Cheilosia morio Zett. (Dipt. Syrphidae) als physiologischer
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CHAPTER 5
THE ECOLOGY, HARVEST AND MARKETING OF TITICA VINE ROOTS
(HETEROPSIS SPP.: ARACEAE) IN THE EASTERN BRAZILIAN AMAZON
ABSTRACT
The aerial roots from various species of the hemiepiphytic vine Heteropsis (Araceae) locally known
as “titica” have long been used in the Amazon region as a strong flexible material in construction and making
handicrafts. In recent decades commercial demand for this product has increased for use in making wicker
furniture. I conducted studies in the Alto Rio Guamá Indigenous Reserve and other sites in the eastern
Brazilian Amazon to describe and quantify several aspects of titica ecology and root biology (density, host tree
size and type, root abundance, root growth and maturation rate), root harvesting, and transformation of the
roots into a commercial product. I collected information on these topics by doing inventories of titica vines,
accompanying harvesters, experimentally cutting roots, and interviewing titica dealers and furniture makers in
urban areas.
The average density of host trees with titica roots in relatively undisturbed upland “terra firme” forest
areas with moderate to light titica harvesting was 371 trees per ha with 115 of these having commercially
harvestable roots. In these areas there was an average of 0.9 commercially harvestable roots and 2.4 nonharvestable roots per host tree generating a titica root density of 1332 roots per ha with 26% of these being
commercially harvestable. Titica plants did not show a positive preference for particular types of host trees,
but the number of titica roots per host tree did increase with host tree size. The vines were virtually absent
from forest areas subject to periodic flooding. Titica was also rarely found on pioneer tree species in patches
of secondary forest recovering from a fire that occurred almost twenty years ago.
Commercial quality titica roots need to be relatively thick (usually ≥ 3.5 mm) and have long (≥ 2 m)
sections without nodes (bulges). In order to prepare roots for sale, nodes, bad pieces, and the root cortex were
removed. Once the processed roots dried, their weight was reduced to 19% of the initial amount harvested in
the forest. Since harvesters remove some nodes from roots immediately after pulling them from host trees, the
weight of roots they end up selling is about 30% of the amount they bring out of the forest. Harvesters
typically collected 10 to 40 kg of roots per day in the forest and depending on the amount and extent of help
from other family members, they spent an extra day processing the roots from each day of harvesting. Since
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dealers in the region usually pay about $1.00 per kg of processed roots, a harvester collecting 35 kg of roots
per day will earn about $US 5.25 per day for the 7 days he spent traveling and harvesting roots and the 3 days
he and his family spent harvesting and processing the roots. The revenue of first time harvests of titica roots
was about $US 10 per ha.
My experimental cutting of mature titica roots showed that harvesting may have a severe impact on
root survival. In two treatments where at least half of mature titica roots per host tree were cut about 4 m
above the ground, an average of 63% cut roots died and only 16% showed some regrowth after seven months.
The roots that did start to grow back had an average of 1.7 new roots arising from them, but each cut root had
only one commercial quality root to replace it. These commercial quality roots were growing at an average of
220 cm per year. The high mortality and slow regrowth following root harvest indicate that many decades may
be needed in between intensive titica harvests in one area. Options for managing titica harvests include
reducing the percentage of roots removed per plant, setting up blocks of forest that would be harvested in
succession, and in some cases integrating titica harvests into long-term timber cutting cycles.
INTRODUCTION
Indigenous people and other forest dwellers in the Amazon region have long harvested Heteropsis
(Araceae) vines for use as a lashing material in construction and making handicrafts (Boom and Moestl, 1990;
Oliveira et al., 1991; Milliken et al., 1992; Balée, 1994; Paz y Miño C. et al., 1995). The strong flexible aerial
roots of these hemiepiphytic plants have also been sold to make wicker (“rattan”) furniture, baskets and other
woven articles in Brazil, Guyana, Venezuela, and Peru (Pio Corrêa, 1931; Madison, 1979a; Rodrigues, 1989;
Whitehead and Godoy, 1991; Balée, 1994; Berry et al., 1995; Hoffman, 1997; Troy and Harte, 1998; Bown,
2000). As deforestation and over-harvesting reduce supplies of rattan from Asian palm species, the demand
for the roots of Heteropsis and other suitable vines from the Amazon seems to be increasing in both South
American domestic and international markets (Whitehead and Godoy, 1991; Hoffman, 1997; Troy and Harte,
1998). Like many non-timber forest products (NTFPs) from the region there is relatively little information
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about the amounts and impacts of current harvesting and how much could be available if the harvest was
managed on a sustainable basis.
There are about 13 species of Heteropsis plants (Croat, 1988; Gentry, 1993), mostly found in the
Amazon region. The species that have been most utilized are H. flexuosa (Kunth) Bunting, H. longispathacea
Engl., H. spruceana Schott, and H. oblongifolia Kunth (Pio Corrêa, 1931; Boom and Moestl, 1990; Oliveira et
al., 1991; Balée, 1994; Berry et al., 1995; Paz y Miño C. et al., 1995, Troy and Harte, 1998). While H.
jenmanii Oliver has also been widely referenced in the literature, this name is now properly considered a
synonym for H. flexuosa (Bunting, 1979). The vernacular name most often used for the commonly harvested
species in the Brazilian Amazon is “cipó titica” (titica vine)(Pio Corrêa, 1931; Rodrigues, 1989; Oliveira et al.,
1991; Balée, 1994, Troy and Harte, 1998).
Titica plants are one type of Araceae vines with a hemiepiphytic habit (sensu Putz and Holbrook,
1986; Appendix 5-A) found in primary tropical moist forests (Hoffman, 1987; Croat, 1988; Bown, 2000).
Unlike “true” epiphytes, the seeds of this secondary type of hemiepiphyte germinate on the forest floor and
reach reproductive status in a host tree. Young seedlings appear to use a skototropic mechanism to grow along
the forest floor toward shady spots in search of a support structure (Hoffman, 1975) as described for a related
Monstera vine (Strong and Ray, 1975). This support is usually provided by a host tree, but rocks have also
served as a climbing substrate for Heteropsis (Croat, 1988). When a seedling reaches a potential host, it
switches to a light seeking mode and grows up the trunk with the aid of fine adventitious anchoring (climber)
roots that adhere to the bark (Wilder, 1992; Appendix 5-A; Figure 5.1). The juvenile stage is reached when the
stem reaches the lower part of the host tree canopy (typically where the first main branch is located) and
produces its first lateral branches with larger leaves (Ray, 1992; Hoffman, 1997; Figure 5.1). As the titica
plant approaches adulthood, the plant’s original connection with the soil is broken when the lower stem
senesces. Around this time, a second type of adventitious root called an absorbing root (Appendix 5-A)
sprouts from the main titica stem or a lower branch and grows downward until it enters the ground and
branches out to give the plant a renewed supply of water and nutrients from the soil (Kelly, 1985; Putz and
Holbrook, 1986; Croat, 1988; Benzing, 1990; Wilder, 1992; Hoffman, 1997, Bown, 2000; Figure 5.1). For the
remainder of this chapter, a “titica root” will refer to the aerial portion of an absorbing root unless otherwise
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specified. Titica plants can also reproduce vegetatively via a flagellar shoot that can arise from a main stem on
a standing or fallen host tree. It may reclimb the same host tree or grow scandently on the forest floor in search
of a new one (Hoffman, 1997).
A titica root usually descends in close contact with the host tree where it wraps around the trunk or is
intertwined with other titica roots or roots of other vines. Occasionally a root that arises from a lateral branch
hanging out from the host tree descends directly to the ground. Hoffman (1997) calls these two types of roots
“trunk roots” and “drop roots” (Appendix 5-A; Figure 5.1). In the process of growing down, a root apical
meristem will often abort or be injured and form a node that is a pronounced bulge about 1 cm long (Wilder,
1989; Hoffman, 1997; Figure 5.2). In many cases, a replacement root (Appendix 5-A) of similar width will
sprout above this node and continue the downward growth (Wilder, 1989, Wilder and Johansen, 1992).
While Heteropsis is an herbaceous monocot that is technically not woody (Putz, 1984a), its roots and
stem are tough. Heteropsis roots are distinguished from other equally strong vines and are particularly
attractive for commercial purposes because the dark epidermis and cortex can be easily peeled away to leave a
smooth, blond, and rot-resistant inner stele packed with long strong fibers. This core can be further split along
the vine’s length into slender pieces that retain strength and flexibility (Rodrigues, 1989; Whitehead and
Godoy, 1991; Potiguara and Nascimento, 1994; Hoffman, 1997).
The plant’s presumed abundance in primary forests and resilient growth has led to optimistic
projections about titica harvest revenues for forest-based communities (Godoy and Whitehead, 1991; Paz y
Miño C. et al., 1995; Troy and Harte, 1998). Two recent studies, however, indicate that severed roots often die
and harvesting all roots often kills the whole plant (Hoffman, 1997; Durigan, 1998). In Guyana, harvesting
impacts are reduced by limiting harvest to the relatively small number of long drop roots that hang free from
the host tree trunk (Hoffman, 1997). Most harvesters in Brazil, though, apparently pull down most mature
roots (Troy and Harte, 1998; Durigan, 2000). There is, therefore, a need for more information about the basic
biology of titica vines, harvesting intensity and economics, and regeneration potential under different
harvesting regimes.
As part of my research to evaluate the ecological and economic aspects of several NTFPs in the Alto
Rio Guamá Indigenous Reserve, I conducted observations and field experiments to describe the harvest and
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commercialization of titica vines in this reserve and regional markets. The main objectives of these titica
studies were to: 1) quantify the density of titica plants and roots on a per area basis, 2) quantify the number and
size of titica roots per host tree, 3) describe the relationship between titica plants and host tree diameter and
type, 4) estimate the maturation time and natural mortality of titica roots, 5) quantify the impact of harvesting
on root mortality and regrowth, 6) quantify the length and weight of roots harvested and transformation during
commercial processing, 7) document the daily yield of commercial titica harvesting and the time involved in
collecting and processing roots, and 8) describe the structure and economics of titica harvesting and marketing.
I will finally discuss how different management scenarios could address some of the problems apparently
caused by current harvesting practices in the eastern Brazilian Amazon.
STUDY AREA
The main study area for observing titica ecology and harvest was the Alto Rio Guamá Indigenous
Reserve in eastern Pará state, Brazil. This 278,000 ha area of closed tropical rainforest is the principal
homeland of the Tembé Indians. Studies were carried out near the village of Tekohaw on the Gurupi River and
the village of Cajueira on the Uraím River in the southern part of the reserve. Additional observations on
plants were made in similar forests on Fazenda 7, a private property near the city of Paragominas, Pará.
Dealers and furniture makers that handled titica roots were interviewed in the town of Gurupi on the Gurupi
River in Maranhão, and Belém, the capital city of Pará.
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METHODS
TITICA BIOLOGY AND POTENTIAL HARVEST OF AERIAL ROOTS
Density of Titica Host Trees and Roots and Number of Roots per Host Tree
The density of host trees with titica plants, the number of roots per unit area, and the number of roots
per host tree was estimated during November and December 1999 in three inventories at Tekohaw and
Cajueira. These sites represented a gradient of forest use and disturbance. The first inventory at Tekohaw was
conducted in an area of forest about 1 km from the village that was heavily used by many of the village’s 25
household members in trips to farm fields, hunt, or collect forest products for personal use. The area was a
mosaic of mostly intact “terra firme” type forest and secondary forest that had burned about 20 years ago.
Titica plants were surveyed in 1.7 ha divided into 48 unburned and 19 burned 10x25 m (250 m2) plots that
were located through stratifed random sampling along 500 m long belt transects. The second Tekohaw
inventory was conducted in an area of forest about 3 km from the village. This area was next to a commonly
used trail, but use of forest resources was less extensive there than the site closer to the village. The third
inventory site was located about 2 km from the village of Cajueira. This intact forest was lightly used by
people from the small village of Cajueira, but it was becoming a major site for commercial titica collection by
people from other villages in the reserve. The second and third inventories were each conducted in 0.6 ha of
forest divided into unburned 10x25 m plots located by stratified random sampling of six plots each in four 250
m long belt transects.
The information collected during these surveys was the number of host trees in every plot that had
either a titica root or climbing stem. Climbing stems generally represented titica seedlings or juveniles
(Appendix 5-A) while roots represented adult titica plants. The number and type (described below) of roots
that were present at 1.5 m above the ground were recorded for each host tree. Since roots generally did not
branch out until they got close to the ground, this number approximated the number of roots attached to the
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main plant stem in the lower canopy of the host tree. All live roots were classified as immature or mature
(Appendix 5-A). Immature roots had soft outer layers and may or may not have entered the ground. All
mature roots had entered the ground and had fully hardened outer layers. At the medium-use site (3 km from
Tekohaw) and light-use site (at Cajueira), experienced collectors and I further classified mature roots as
commercial or non-commercial. Commercial roots were generally at least 3.5 mm wide and had relatively few
nodes visible at ground level. Dead roots were detected by their degraded state and were easily broken when
pulled. These were not counted in surveys.
The average number of host trees with titica plants and the average number of titica roots of different
types per 250 m2 plot at each site was multiplied by 40 to generate estimates of titica host tree, titica root
density and titica climbing stem density per ha. Data on host trees with titica roots were combined from the
medium-use (Tekohaw) and light-use (Cajueira) sites to analyze the average number and percentage of
commercial roots per host tree.
Comparative Dimensions of Commercial and Non-commercial Mature Roots
The differences in the width and abundance of nodes on mature commercial and non-commercial
roots were quantified by harvesting the mature roots from all 73 host trees that had commercial roots in 0.5 ha
(20 plots) at the Cajueira inventory site. Prior to measurements, experienced harvesters classified each root as
commercial (n=112 roots), non-commercial due to thinness (n=113 roots), or non-commercial due to too many
nodes (n=11 roots). The width of each root was measured at one meter intervals. When the measuring point
would have fallen on a node, the measurement was taken 4 cm below it. The number of nodes was counted on
every root and divided by the root length to calculate the average number of nodes per meter.
The Relationship Between Titica Plants and their Host Tree Type and Diameter
Host Tree Type and Diversity
The common Tembé or Portuguese name (morphospecies) of the host tree bearing titica plants was
recorded in the three inventories described above to gauge the diversity of these hosts. The number of host
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trees belonging to each morphospecies was divided by the total number of morphospecies at the heavily used
site at Tekohaw to calculate the percentage that each morphospecies contributed to the total. These
percentages were ranked from most common to least common and compared to a similar abundance ranking of
morphospecies of all trees (≥10 cm DBH) that were inventoried in 0.9 ha of the same belt transects (36 plots x
250 m2 per plot). Botanical specimens were not collected from host trees, so this comparison could not be
made on the basis of scientific species.
Host Tree Diameter
The DBH (diameter at 1.5 m) of host trees bearing titica plants was recorded in 0.25 ha (10 plots) in
the heavily used site at Tekohaw. The DBH size class of host trees ≥10 cm was compared with the size class
distribution of all trees ≥10 cm DBH inventoried in 0.2 ha (8 plots) of the same belt transects. Data were
analyzed to compare the size class distribution of titica host trees to all trees. The average density of host trees
(≥10 cm DBH) with titica roots was divided by the average density of all trees with ≥10 cm to estimate the
percentage of trees in the area that served as hosts for adult titica plants.
The DBH of host trees with at least one mature titica root was recorded in all 0.6 ha (24 plots)
surveyed in the medium-use site at Tekohaw. The number of climbing stems, number and type of roots was
compared with the host tree DBH to examine the relationship between host tree size and titica plant
development.
Rate of Root Maturation and Natural Mortality
The amount of time that roots spend in various phases was investigated by individually marking 56
immature roots (already in the ground) on 31 host trees and 159 mature roots on 39 host trees in the mediumuse site at Tekohaw. Each root type was spray painted a different color in December, 1998. The status of
these roots was recorded as immature, mature or dead in follow-up observations in March, April and July,
1999 – 76, 134, and 210 days respectively after the initial observations. Results were analyzed to determine
the maturation and mortality rate of immature roots and the mortality rate of mature roots.
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TITICA HARVEST IMPACT, YIELD AND SHORT-TERM ECONOMICS
Impact of Harvesting on Root Mortality and Regrowth
To measure the effect of harvesting on titica roots and test the hypothesis that the amount of roots cut
per plant could affect the mortality and regrowth of these roots, mature aerial roots were cut from two
treatment groups of host trees at the medium-use site at Tekohaw in December, 1998. In the first treatment, 37
host trees had half (or half plus one) of all commercial quality titica roots present at 4 m above the ground
(accessed by ladder) cut with pruning shears approximately 4 cm below the nearest node. In the second
treatment with 56 host trees, all of the commercial roots were cut in the same way. Host trees were assigned to
treatment groups so there was approximately an equal average number of roots per tree in both groups before
any roots were cut. Nodes of the severed roots were spray painted to facilitate future observations. Roots were
cut rather than pulled to facilitate follow-up observations of regrowth.
Host trees were revisited 3 and 7 months later to record the status of these painted cut roots as dead or
alive. If living roots had any regrowth within a meter of the point where the root was cut, the number, lengths,
and widths of newly sprouted roots were also recorded. The percentage of roots that were dead, alive, and
alive with growth per host tree were calculated and compared for both treatment groups. The mortality rate of
uncut mature roots observed at ground level (described in the previous section) provided information on the
natural rate of mature root mortality. Where regrowth was observed on cut roots, replacement roots ≥3.5 mm
wide were classified as commercial quality. Regrowth data at 7 months was multiplied by 1.7 to estimate the
regrowth length of all roots and commercial quality roots per year.
Root Transformation During Commercial Preparation
The first stage of preparing commercial quality titica roots for sale was breaking and cutting harvested
roots into stems that are free of nodes and other pieces that are split or damaged. I measured the effect that this
process had on root length, weight and the number of root stems per host tree by processing the commercial
roots harvested from 37 host trees in 0.25 ha (10 plots) at the Cajueira site (half of the host trees measuring
dimensions of commercial and non-commercial roots described above).
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Following the removal of nodes and bad pieces, the outer layers of titica roots need to be stripped off
to prepare them for sale. Since roots in eastern Pará are sold by weight, I measured the loss in weight that
accompanies each transformation between harvested and saleable roots. I conducted this study by harvesting
and processing all the commercial roots from 20 host trees with at least two commercial roots found near the
medium-use inventory site at Tekohaw. The roots from each host tree were weighed immediately after
harvest, after node and bad piece removal, and after cortex removal. Total length of roots was also measured
before and after node removal. The stripped root stems were then stored in the shade and reweighed every day
for nine days until there was no further weight loss from evaporation. The average percentage weight loss of
roots per host was calculated for each stage of commercial preparation.
Harvester Daily Yield and Time Involved in Collecting and Processing Titica Roots
Data on harvesting yield and rates were gathered by accompanying one man collecting titica vine
roots at Cajueira for five days. After pulling down a root, this commercial collector broke roots into pieces that
had only one node at the end so root stems could be easily tied in a bundle at the top where the nodes were
bunched together. The number of host trees and root stems harvested per host tree were recorded each day.
The number of commercial roots collected per host tree and the GPS position of each host tree were noted for
four days. GPS data was analyzed to describe the harvester’s search pattern and to estimate the average
distance between host trees and area searched. The time of arrival and departure at each host tree was recorded
for the first two days and analyzed to estimate the average search and harvest time per host tree. All
observations were made by two researchers so the normal working pace of the harvester was not interrupted.
The amount of time required to process roots was measured for a sample of roots harvested at
Cajueira. Observations included the starting number of stems, and starting and finishing weight of each bundle
of root stems. Since titica root processing is often a family endeavor, the age of the person stripping the roots
was recorded along with the number for each bundle of root stems. Results were analyzed to determine the
average number and grams of root stems processed per minute and the relationship between age of processor
and the processing rate.
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Titica Vine Market Structure and Economics
In November 1998, I interviewed two dealers in the town of Gurupi and the city of Belém who buy
large quantities of titica vine roots and other forest products and the managers of four businesses in Belém that
make artesenal style furniture with titica roots and other plant materials. These managers were asked to
comment on sources for titica and the amount of titica roots they bought or consumed per year and per finished
product and amounts of money related to these purchases and sales.
To assess commercial activity in the reserve, I requested a Tembé Indian forest product buyer in
Tekohaw to maintain a log book that recorded the name of the collector and amounts of titica roots he bought
from Tembé and Ka’apor Indians in the area. He provided information on transactions that occurred between
July, 1999 and January, 2000.
RESULTS
TITICA BIOLOGY AND POTENTIAL HARVEST OF AERIAL ROOTS
Density of Titica Host Trees and Roots and Number of Roots per Host Tree
Inventories at Tekohaw and Cajueira showed that as the intensity of site use increased, the density of
host trees with titica plants and the number of titica roots per ha decreased. The most heavily used forest plots
close to Tekohaw had the lowest density of host trees in these three inventories of unburned forest sites with
any titica roots (mean 143 ± 22 per ha). Comparing this number to the density of all trees ≥ 10 cm DBH in the
same plots indicated that as many as 48% of these trees had a titica plant with at least one root. The lightly
used site at Cajueira had the highest titica host tree density (mean 453 ± 32)(Figure 5.3). The density of host
trees with commercial quality roots was 67 ± 15 per ha at the medium-use site at Tekohaw (24% of host tree
density with any roots) and 163 ± 14 per ha at Cajueira (36% of host tree density with any roots). The effects
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of the 20-year-old fire were still evident at the Tekohaw site since there was an average of only 2 trees per ha
in the burned plots in the heavy-use area that hosted any form of titica plant.
The number of titica roots per ha in the unburned plots at the heavy, medium and light uses sites
reflected the same pattern as the host tree density. The lightly used Cajueira site had the highest density with
an estimated 1748 ± 142 titica roots per ha while the heavily used site at Tekohaw had the lowest with 544 ±
84 roots per ha (Figure 5.3). The percentage of these roots that was commercial quality at Cajueira was 32%
while only 15% of the roots at the medium-use Tekohaw site could be harvested for sale.
The estimated density of climbing stems per hectare was almost identical at the medium-use site at
Tekohaw and light use site at Cajueira and Tekohaw with a combined average of 267 climbing stems per ha.
Evidence of seedlings at the heavy-use site at Tekohaw was very low since there was only an average of 19
climbing stems per ha found there.
Analysis of roots per host tree showed minimal differences between the medium and lightly used sites
at Tekohaw and Cajueira so these data were combined to measure the average number and type of titica roots
per host tree in the reserve. There was an average of 3.6 ± 0.2 roots in host trees with any type of root with an
average 1.0 ± 0.1 commercial quality roots per host tree (Figure 5.4). On host trees with at least one
commercial quality root, the average number of roots was 5.8 ± 0.4 with an average of 3.1 ± 0.3 commercial
quality roots per host tree.
Comparative Dimensions of Commercial and Non-commercial Mature Roots
The average width of commercial quality roots harvested from the Cajueira inventory site was 5.4 ±
0.1 mm. This was significantly greater than the roots considered non-commercial because they were too thin
that averaged 2.5 ± 0.1 mm (One-way AOV: F=586.4; p=0.000) and not different from roots considered unfit
for sale because they had too many nodes. There was an average of 1.2 ± 0.1 nodes per meter on commercial
quality roots. This was significantly less than the combined average of both categories of roots considered
non-commercial that had 2.1 ± 0.1 nodes per meter (One-way AOV: F=50.8; p=0.000).
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The Relationship Between Titica Plants and their Host Tree Type and Diameter
Host Tree Type and Diversity
The number of morphospecies that served as host trees for titica plants was the least at the heavily
used site at Tekohaw (29 morphospecies from 183 trees in 1.2 ha inventoried), intermediate at the heavy-use
site at Cajueira (44 morphospecies from 364 trees in 0.6 ha inventoried) and greatest at the medium-use site at
Cajueira (52 morphospecies from 224 trees in 0.6 ha inventoried). The imprecision of combining common tree
names into morphospecies groups means that strict numerical comparisons of these numbers are not
appropriate. The results in general showed that titica plants do not show any strong preferences for the type of
host tree they colonize.
The approximate match between the relative abundance of all trees and the relatively abundance of
titica host trees was demonstrated in the comparison of host tree morphospecies at the heavily used Tekohaw
site and the inventory of all trees (≥10 cm DBH) in the same transects. These results revealed that five of the
six most common host trees for titica (caçador (Lecythis spp.), tiriba (Eschweilera spp.), macucu (Licania
spp.), breu (Protium spp.), and faveira (Parkia spp.)) were also among the eight most common trees in the
general tree survey. Trees that were in the ten most common types found in the general survey that were
almost never recorded as titica host trees were embauba (Cecropia spp.), açai (Euterpe spp.), inga (Inga spp.),
lacre (Vismia spp.) and marupá (Simaruba spp.).
Host Tree Diameter
The survey of titica host trees at the heavily used site at Tekohaw showed that 21.2% of host trees
with titica roots are ≤10 cm DBH (min. 4.2 cm). Most host trees were in the 10 to 19 cm DBH range (40.4%)
with 17.3%, 13.5%, and 7.7% found respectively in the 20-29, 30-39, and ≥40 cm DBH size classes. No data
were available on the proportion of all trees in the smallest size class, but a comparison of the percentages in
each of the size classes ≥10 cm DBH between titica host trees (n=43 host trees) and all trees (n=85 trees) in the
same area was highly correlated (Pearson’s Correlation Coefficient: ρ=.976; p=.024) so in general adult titica
plants had no special association with the diameter of their host trees.
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The titica inventory at the medium-use site at Tekohaw showed that the average number of all titica
roots and commercial roots increased with the diameter of host trees with any type of titica root (Figure 5.5).
The mean number of roots on host trees with 4-9 cm DBH was significantly less than the average in all larger
size classes (One-way AOV; p ≤ 0.05). The greatest value was found in 30-39 cm DBH host trees (5.3 ± 1.0
roots per host). The association between large host trees and commercial titica roots was even stronger than
with roots of all sorts since there was a mean of 2.5 ± 0.7 commercial roots per host that was significantly
greater than found in all three of the smaller size classes of trees <30 cm DBH (One-way AOV: p ≤
0.05)(Figure 5.5).
The relationship between host tree DBH and titica plants with climbing stems is the reverse of the one
with adult plants with roots. The inventory showed that 84% of the 122 host trees with titica climbing stems
were <10 cm DBH. Less than 6% of all host trees with a titica climbing stem were ≥20 cm DBH.
Rate of Root Maturation and Natural Mortality
Immature aerial roots that had penetrated the ground in December 1998, steadily reached maturity
during the three follow-up observations of titica roots at Tekohaw (Figure 4.6). Seven months after the initial
observations, 86% of the roots that were immature at the start of the observations had developed the hard
exterior and strength characteristic of mature feeder roots. About 9% of them died before they reached
maturity. Roots that reached maturity had a higher life expectancy since only 2 out of 159 (1.3%) of mature
roots died during the seven months of these observations.
TITICA HARVEST IMPACT, YIELD AND SHORT-TERM ECONOMICS
Impact of Harvesting on Root Mortality and Regrowth
Three months after titica roots were cut at the Tekohaw medium-use site, 33 % of cut roots had died
on host trees where half the commercial roots were cut; the mortality of cut roots was 45% in host trees where
all commercial roots were cut, although this difference was not statistically significant. Seven months after
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cutting, the mortality of cut roots was close to 63% for both groups (Figure 5.7). It was not possible to observe
effects of cutting on the whole plant from the 4 meter elevation. No mortality of unsevered roots was observed
in the first treatment group where only half of the commercial quality vines had been cut.
Three months after root cutting, only 2 out of 224 roots had a new root growing. Seven months after
cutting, regrowth was observed more often in the 100% root cut treatment (Figure 5.7), but the difference was
not significant. Overall regrowth was seen in both treatment groups combined on 23 cut roots (16.9 ± 3.7% of
cut roots) from 22% of host trees that had roots cut. When this occurred there was an average of 1.7 new roots
per root cut (maximum 4) that usually grew out from the nearest or second to nearest node above the cut.
These new roots sometimes diverged into two roots within centimeters of their origin from the main root.
Judging by their diameter, 59.5 ± 9.4% of these new roots were commercial width. It is not known, however,
what percentage of these new roots would ultimately make it to the ground and mature to commercial quality.
If linear growth of new roots produced after old root cutting is averaged over all host trees that had
roots cut, the average cut root will regrow an estimated 19.7 ± 5.3 cm per root per year. If only host trees with
regrowing cut roots are considered, this same figure is 91.7 ± 17.1 cm per root per year. For host trees as a
whole, the estimated amount of regrowth from cut roots is 76.6 ± 22.3 cm per host tree per year when all host
trees are considered; it is 356.1 ± 77.0 cm per host tree per year for host trees with at least one cut root that
sprouted a new root. For all of these regrowth measurements, the amounts were higher for the treatment where
all commercial roots were cut, but these differences were not statistically significant.
Root Transformation During Commercial Preparation
After nodes and unsellable pieces were removed from commercial roots harvested from the Cajueira
survey plots, the number of stems produced was about twice the number of roots harvested per host tree
(Figure 5.8). The total length of commercial roots harvested per host tree was 13 meters at the Cajueira survey
site; it was 22 meters per host tree from the 20 host trees harvested for the product transformation observations
at the Tekohaw sample harvest. After initial processing in the Cajueira survey and Tekohaw harvest sample,
these total lengths were reduced by 27 to 30%.
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The weight of root stems after node and bad-piece removal was 31% less than the starting weight of
harvested roots from these two samples. After the cortex was removed from root stems derived from the host
trees harvested at Tekohaw, the weight immediately dropped from an average of 420 ± 60g per host tree to 183
± 26 (Figure 5.9), a further weight loss of 56%. In the days that followed, the uncovered stele lost additional
weight, presumably due to evaporation of water. After eight days, the average weight of the cleaned root stems
per host tree had declined to 119 ± 17 g.. A daily charting of this process showed that the biggest losses
occurred in the first five days. After that the percentage weight loss due to drying leveled off at 35.1 ± 1.1%
(Figure 5.10). Comparing the weight of roots harvested from the host trees to that of the saleable root stems
that have had their nodes and cortex removed and dried, only 19.4% of the original weight remained. Since
harvesters typically remove some nodes and bad-pieces immediately after roots are harvested, the ratio of root
weight brought out of the forest to the ultimate weight of roots sold is probably about 30%.
Harvester Daily Yield and Time Involved in Collecting and Processing Titica Roots
Accompanying a harvester collecting titica roots for sale at Cajueira provided a variety of measures of
harvesting yields, rates and effort. During five consecutive days of harvesting, he collected roots from an
average of 44.8 ± 4.1 host trees per day (n=5 days). This led to a daily average of 192.0 ± 12.4 roots collected
per day on the final four days (harvested roots were not counted on the first day). Roots gathered during the
entire five-day period had some nodes removed in the forest and were broken into 2435 root stems that
weighed 167.5 kg producing daily averages of 487 ± 19 stems per day and 33.5 ± 1.4 kg of root stems per day .
The average time he spent between two harvested trees was 1.2 ± 0.2 minutes per tree (n=78 host
trees in the first two days). Using the straight line distance between two harvested trees (calculated by the
difference in GPS readings), this means that 39.1 ± 4.4 meters were searched per minute (n=50 host trees), an
estimated search rate of 26.1 ± 4.9 min. to search one ha of forest if the search path was 10 meters wide. The
average time spent at a tree was 4.9 ± 0.4 minutes per tree (n=84 host trees in the first two days). This time
included cutting roots to be harvested at the base, pulling the descending roots from the tree, breaking them
into stems and tying them into a bundle. As harvesting progressed, multiple bundles were tied together into a
“cabeça” – head. Finally several “cabeças” were united in a large bundle called a “feixa” that weighed 15 to
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20 kg. Depending on the search pattern, a completed “feixa” was sometimes left by a trail for a time and
picked up later so the harvester would not have to carry this load from tree to tree. It was then usually picked
up on the way back to the village. This harvester’s usual goal was to collect two “feixas” per day. Older men
usually had a more modest goal of one small “feixa” (10 to 15 kg) per day while strong men intent on making
as much money as possible said they sometimes harvested three to four “feixas” per day if they were in an area
with lots of titica.
The average distance between successive host trees was 59.8 ± 3.8 meters (134 host trees in 4 days).
A map of the host tree positions visited for four days shows that the harvester searched in one general direction
throughout the day (Figure 5.11), but when he found an area that seemed to have many host trees with titica in
close proximity, he wandered around that area for awhile before returning to the general search pattern
direction. An examination of the order that host trees were visited on the first day (Figure 5.12) shows that his
movements included frequent backtracking to look for trees with large numbers of titica roots. If the
harvester’s average search path was 10 meters wide, the density of host trees with commercial quality titica
roots he was collecting from was 29.2 ± 2.3 host trees per ha. The density of commercial roots he was
harvesting was effectively 122.3 ± 15.2 roots per ha. These are both substantially less than the densities of
commercial roots and host trees with commercial roots estimated from the survey of nearby transects at
Cajueira and the Tekohaw medium-use site.
Roots collected by the harvester at Cajueira were brought back to Tekohaw where his family all
participated in cutting off the top node from each root stem and stripping off the cortex. Processing the
product of the father’s five days of solo collecting activity took about two and a half days for the family with
two to seven members working at any point in time. The average rate for processing these root stems was 0.96
± .04 stems per minute (roots from 147 host trees). By initial weight, stems were stripped at the average rate of
64.8 ± 2.8 g per minute. The rate of root processing was significantly correlated with the age of the person
stripping off the cortex (Pearson’s Correlation Coefficient: ρ=.192; p=0.02). Children that were 10 years and
under were given smaller root bundles to process, and they averaged 0.35 to 0.92 stems per minute at this task
while people who were 13 years and over all stripped just over one stem per minute (Figure 5.13) although
these differences were only statistically significant between groups of children 8 years and under and people
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who were 10 years and older. If one skilled adult had processed all these roots at the rate of 1 stem per minute,
it would have taken them just over 40 hours – about eight hours per day for five days.
Titica Vine Market Structure and Economics
Between 1997 and 1999, the prices that harvesters were paid for cleaned titica roots varied from $R1
to $R2 per kg. This represented an equivalent amount of $US in late 1997 and gradually declined to prices that
were 50 – 60% of these values. Before 1998, titica roots collected in the Tembé reserve were periodically sold
to a small dealer in a river town or the large city of Belém. The advantage of selling to the river dealer was
that he immediately paid for the vine roots in cash, but he usually paid about 25% less than the dealers in the
city. The downside of getting a higher price from these city dealers was that they deferred full payment until
they sold them to other customers. These large city dealers typically only bought batches of hundreds of
kilograms of cleaned roots at one time so collectors had to wait until a large amount had been gathered before a
sale was possible.
From 1998 on, two Tembé men in the village of Tekohaw began buying titica roots and breu resin
(from Protium spp. Chapter 3) from Tembé and Ka’apor Indians in the area. Harvesters received cash or credit
toward merchandise that the Tembé middlemen bought when the roots were sold. Between July 1999 and Jan.
2000, the major Tembé buyer in Tekohaw bought 1754 kg of processed titica roots from 19 different
collectors. Conversations with Tembé Indians indicated they usually devoted several weeks to commercial
harvest of titica roots per year. These efforts usually yielded 10 to 45 kg of roots (with some nodes removed)
per day that yielded an estimated 3 to 15 kg of sellable root stems per day of collecting.
The dealers from Gurupi and Belém sell some titica to other dealers outside of Belém, but most
consumers are local artesenal style furniture makers. These businesses are all small. In the smallest, the owner
along with a few other people do most of the work. The largest one had five regular employees, but up to 60
people would be hired on a temporary basis if the owner secured a large order. All stores had ready-made
items for sale, but most of the business seemed to depend on custom orders..
The mainstay of these businesses was making vine furniture, but to varying extents all of the shops
also made wooden or metal furniture where vines are either a marginal component or absent. Titica vine was
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the principal vine used in these enterprises and was the dominant material in many types of chairs and sofas. It
is sometimes mixed with other vines from the region and cultivated “vime” vines cultivated in southern Brazil.
A small to medium chair consumed three or four kg of titica with a large one using up five to eight kg. A large
basket can consume one kg of titica while leftovers from larger pieces are used to make small baskets. Most of
these items were made only from titica root stems only and are usually sold in their natural color. These stems
were also used to wrap joints in furniture made with other principal materials such as other vines or metal.
Titica can be split and soaked in caustic soda to make it extra pliable. It was also sometimes painted or
stained..
Most of the furniture makers said that they bought titica from three or four regular suppliers who
came by with the material in hand since deals were rarely brokered in advance. These suppliers came from
both Pará and the adjoining state of Amapá. The smallest shop used 50 to 100 kg of titica per month; the larger
ones used 100 to 200 kg per month. A few bought only a month’s supply at a time, while others laid in a stock
of 500 to 1000 kg. The prices paid for cleaned titica roots ranged from $US 1 to 4 per kg; $US 2 per kg was a
rough average. The larger scale dealers paid lower prices for buying larger quantities at one time.
Wicker furniture making is a labor-intensive enterprise. A large chair generally took one experienced
person one and half to two days complete; a complicated piece could take three days. Since workers were paid
about two minimum salaries ($US 12 per day), the cost inputs for a large chair were about $US 10 for material
and $US 24 for labor for a total of $US 34. A regular large chair usually sold for $US 100 to 180. A very
fancy one could fetch from $US 500 to 1000, but such sales are rare. These dealers sold all or most of their
products to customers in Belém. Larger dealers did some business with buyers in São Paulo or outside Brazil
(U.S., Japan, France, and Belgium). One of the two larger dealers said that business was not as good as it had
been in the past. He used to have 20 - 25 regular employees and had reduced his staff to five. It took these
people several years to learn their craft well so he was not training any more.
In sum, the market for titica roots in Belém was fairly strong, but like many commodities the price
paid to harvesters fluctuated by 100% within several months. It seemed that the maximum price, however,
would never exceed the price for a readily available cultivated fiber from southern Brazil.
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DISCUSSION
TITICA BIOLOGY AND POTENTIAL HARVEST OF AERIAL ROOTS
Density of Titica Host Trees and Roots and Number of Roots per Host Tree
Estimates of the density of trees with adult titica plants at Cajueira and Tekohaw varied from 143 to
453 host trees per ha. This range is higher than H. flexuosa host tree densities in Guyana that varied from 61 to
232 host trees per ha (based on surveys in 0.1 ha plots)(Hoffman, 1997). This Guyana population colonized an
average of 9 to 41% of all trees ≥ 10 cm DBH in various habitat types, lower than the 48% colonization
estimate for the heavily used site at Tekohaw. The lower H. flexuosa host tree density in Guyana may be
partially due to the large abundance of greenheart trees (Chlorocardium rodiei) that were rarely colonized by
the vine (Hoffman, 1997). Titica densities from both the Tembé reserve and Guyana study were dramatically
larger than the 1.4 to 5.3 titica host tree per ha figure reported by Durigan (1998) for the Jaú Park in the central
Brazilian Amazon.
From a harvester’s point of view, the density of roots per hectare matters more than the density of host
trees alone. Given the superior densities of titica commercial stems at Cajueira compared to Tekohaw, it is not
surprising that Tembé men from Tekohaw were willing to spend a whole day traveling by boat to Cajueira
when they wished to gather titica vine roots for sale. While both Tembé sites had a higher density of host trees
with titica than in Guyana, the Guyana sites had an estimated 997 to 1175 mature roots per ha that matched
figures for the Tembé sites because titica plants in Guyana had a higher average number of mature roots per
tree (Hoffman, 1997).
The other noteworthy aspect of the density measurements was the near absence of titica host trees in
forest areas near Tekohaw that were burned in the early 1980s. The causes for this probably relate to past
direct effects of fire, the physiology of titica and the characteristics of the trees most commonly found in the
secondary forest of these areas. It is apparent from the relationship between titica root number and host tree
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size and Hoffman’s (1997) observations of H. flexuosa’s slow growth rate that it takes many decades for a
titica plant to progress from the juvenile stage of climbing a host tree to the time that aerial roots descend from
a main stem high up on the trunk and mature in the ground. A fire that happened 20 years ago that was severe
enough to kill most rainforest trees would have also killed their attached vines so the lack of titica plants with
numerous roots in these burn zones is easily understood. While numerous healthy titica plants are close
enough to recolonize the recovering forest, pioneer trees such as Cecropia, Vismia, and Simaruba that are most
common in this area’s secondary forest almost never had titica plants on them. This may be due to specific
vine avoidance traits of these trees or their mere location in light rich environments that are beyond the
temperature and humidity tolerances of titica plants. López-Portillo et al. (2000) have noted that aroid
hemiepiphytes die quickly when suddenly exposed to high light environments possibly because their hydraulic
architecture is not adapted to this condition. Whatever specific reasons lie behind the low density of titica
plants in secondary forests, the progressive degradation of Amazonian forests by logging, agricultural clearing
and fire has stark implications for titica populations. When a forest is degraded, titica density will decline
along with its role as an economic resource for a long time.
While a few host trees with titica plants at Cajueira and Tekohaw had as many as 15 absorbing roots
on them, the mean number of three mature roots per host tree in the Tembé reserve is considerably less than the
7.7 roots per host tree (calculated from data in Durigan, 1998) in the Jaú National Park in Amazonas state,
Brazil and the 6.6 to 10.5 roots per host tree averages found by Hoffman (1997) in various forest types in
Guyana. The difference between the eastern and central Brazilian Amazon sites may be due in part to
Durigan’s (1988) selection of host trees by the nearest-neighbor method (Durigan, 1998) rather than censusing
all titica host trees in a plot since this method was intentionally designed to mimic a harvester selection
process. Accompanying a harvester collecting titica roots in Cajueira showed a mean 6.9 roots per host tree
average in an area of forest immediately adjacent to the survey plots with less than half that number. It was
apparent that this harvester intentionally avoided spending time collecting roots from host trees that offered
few or no commercial quality roots. The comparison between his root per host tree average and the figure
obtained from the nearby inventory shows that caution is needed in estimating the actual abundance of a
resource based solely on data derived from commercial collection. The large difference in the titica roots per
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host tree average found in the inventories conducted in Guyana and the Tembé sites in Brazil indicate that
differences in forest composition, structure, or climate can affect the productivity of titica plants.
Comparative Dimensions of Commercial and Non-commercial Mature Roots
The average length of individual roots harvested at the Cajueira site (4.7 meters) was considerably
shorter than the 10.1 meter average found by Hoffman (1997) in Guyana. This variation is probably due more
to different harvesting practices in Guyana and the Tembé site than to biological factors. At the Tembé sites,
most or all mature titica roots that are considered thick enough to sell are harvested from host trees except
when they clearly have a large number of nodes on them. Most of these are trunk roots (Figure 5.12; Appendix
5-A) that wrap around the host tree trunk while descending and become tangled with other titica roots or stems
of other locally common climbing vines such as “tarakuwa” (tentatively Philodendron grandiflorum (Jacq.)
Schott (Balée, 1994) and timbo-açu (tentatively Evodianthus spp.(Balée, 1994; Troy and Harte, 1998).
In Guyana, harvesters generally only harvest drop roots that grow straight down from a lateral branch
(Hoffman, 1997). These roots consequently have fewer nodes than trunk roots, and yield longer roots when
pulled and longer sections of node free sections. This marketing preference for longer stems presumably
increases the number of host trees that are harvested in Guyana since Hoffman (1997) found that the number of
trunk roots was often ten times greater than the number of commercial (“drop”) roots per host tree and 73% of
all host trees had only trunk roots. The reason there may be relatively few drop roots is that an absorbing root
held on a branch far from the main stem would increase the risk of breaking off the branch whose main
purpose is to hold leaves out to gather sunlight. As Ray (1986) points out, the branch already has to make a
trade off between the length and width of a branch and the weight of leaves it can support. Accordingly
holding an aerial root farther from the trunk would force a reduction in leaf mass with no apparent gain.
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The Relationship Between Titica Plants and their Host Tree Type and Diameter
Host Tree Type and Diversity
The wide diversity of host trees for titica plants at both Cajueira and Tekohaw indicate that this vine
does not preferentially select certain taxa of trees to serve as climbing substrates. Hoffman (1997) had the
same result in Guyana where more than 42 identified tree taxa served as hosts for H. flexuosa. It was the
apparent lack of colonization on several common trees that was more noteworthy in the titica survey of the
heavily used site at Tekohaw.
Kelly (1985) found that most host trees for a variety of vascular epiphytes in Jamaican tropical forest
had single straight trunks for most of their height, somewhat narrow crowns and thin, smooth or narrowly
fissured bark. Other features that may discourage climbing vines on pioneer and other trees are rapid growth,
flexible trunks, large leaves, exfoliating and chemical emitting bark, and the presence of protective insects
(Putz, 1984a, 1984b; Hegarty, 1991; Talley et al., 1996). Three of the common species in the heavily used and
partially burned site at Tekohaw that apparently lacked titica vines were pioneer trees in the Cecropia, Vismia,
and Simaruba genera. Putz (1984a,b) and Janzen (1969) have both noted that Cecropia is often associated with
Azteca ants that vigorously defend their host trees from any plants that contact their host trees. The lack of
Euterpe palms serving as titica hosts at Tekohaw paralleled the apparent absence of palms as a substrate for H.
flexuosa in Guyana (Hoffman, 1997). Other authors indicated that some palms are relatively free of vines due
to their large long compound leaves that are shed as units (Putz, 1984a; Hegarty, 1991). The tendency for
Euterpe palms to be located near small streams in terra firme forests may have also contributed to their
unsuitability as a titica host tree since Heteropsis seems to prefer well-drained soil habitats (Hoffman, 1997).
Host Tree Diameter
Results of the surveys in Tekohaw indicated that for trees ≥10 cm DBH there was no special
relationship between tree DBH and the probability that it was colonized by a titica plant. This is what would
be expected for a vine that ascends its host with adventitious roots (Putz, 1984b; Hegarty, 1991). The finding
that the number of titica roots increases with host tree size reinforces the finding of Durigan (1999) who found
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a significant positive correlation between host tree DBH and the number of mature roots on it. This
relationship may not hold true for the largest trees since the number of titica roots per host tree peaked in the
30 - 39 cm DBH size class and slightly declined (although not significantly so) in the host trees ≥40 cm DBH.
Hoffman (1997) noted that very few emergent trees in the Guyana rainforest had H. flexuosa vines in them.
While titica plants do not apparently have positive preferences for host tree species, they do show
strong preference for locating their main stem and branches in the upper bole and lowest branching parts of
their host trees (Hoffman, 1997). Vertical specialization and stratification of epiphytes in their host trees has
been observed in other neotropical forests and apparently relates to each species’ preferred conditions for light
and moisture (Putz, 1984b; Kelly, 1985; Todzia, 1986; ter Steege and Cornelissen, 1989). The location of
titica’s main stem and foliage below the host tree’s upper canopy provides it with intermediate levels of light
and moisture (Hoffman, 1997). It seems possible that if host trees become emergent, titica plants could arrive
in an environment that exceeds their tolerance for direct sunlight.
The Tekohaw inventory finding that 84% of titica plants with climbing stems (presumably seedlings
or juveniles) were found on trees ≤10 cm DBH shows that only examining host trees with adult titica plants
may omit important observations about this vine’s establishment phase. Putz (1984b) found that certain
twining vines can only climb host trees ≤10 cm DBH, but it seems this structural limitation would not apply to
titica seedlings because they use anchor roots to adhere to host trees. Additional studies are, therefore, needed
to further examine if Heteropsis climbing stems seek out or have better survival on small host trees and what
might explain this facet of their natural history.
Rate of Root Maturation and Natural Mortality
As was done at Tekohaw, Hoffman (1997) tracked the development of immature titica roots. Six
months after initial observations, he found that 52% of these roots had matured, 33% remained the same and
9% had died. This mortality rate was identical to that found at Tekohaw. The maturation rate was less than the
86% figure found at Tekohaw after seven months, but both studies indicate that immature roots probably take
six to nine months to mature once they have taken root in the ground. This maturation process observable
above ground is accompanied below ground by the extension and ramification of the root away from the host
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tree just below the surface (unpublished data from Fazenda 7 observations; Wilder, 1992). While 9% or
slightly more of these immature roots may die during this stage, the low mortality rate of fully established
mature roots found at Tekohaw showed that they live for many years.
TITICA ROOT HARVEST YIELD AND SHORT-TERM ECONOMICS
Impact of Harvesting on Root Mortality and Regrowth
Documenting the effect of conventional harvesting on the affected roots is a logistically challenging
proposition because there is no way to predict where a root will break when pulled from below and they
strongly resist being broken by bending. Hoffman (1997) determined that some roots break off near their
attachment point, but many others did not. Thus to simulate the effect of root breakage, he cut about 150
commercial roots in 61 host tree canopies. Six months after cutting, 17% of the severed roots had died. The
percentage of roots that died in this experiment was considerably less than the 63% average mortality found in
the cutting experiment at Tekohaw seven months after cutting. While the average number of roots cut per host
tree in the Guyana experiment was similar to the average number cut at Tekohaw (2.5 in Guyana vs. 1.8 and
2.9 in the two Tekohaw treatments), the Guyana treatment only cut an average of 30% of the commercial roots
per host tree versus 50% and 100% cut at Tekohaw. It is also possible that the Guyana plants suffered less
because they were cut in the February in the middle of the rainy season while the titica roots were cut at
Tekohaw in December when the rainy season was just beginning.
The Tekohaw cutting experiment did not make observations on the effect of harvesting or root cutting
on the whole titica plant, but Hoffman (1997) noted that when 100% of the roots were cut the whole plant died.
The plant showed signed of serious stress including branch dieback and yellow leaves when just over 80% of
roots were cut, sometimes showed a stressed response when 50-80% of roots were cut, and rarely showed signs
of stress if less than 50% of roots were cut. Durigan (1998) did follow-up observations of titica plants
following conventional root harvesting and found that 15% of the titica plants died within two months if all
commercial roots were pulled down. One year after harvesting, the whole plant mortality rate had risen to an
average of 46% (range 17 to 83 % at four sites).
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In the early phase of their establishment on a host tree, Heteropsis and other closely related secondary
epiphytes secure a supply of water and nutrients from the ground through their original climbing stem. There
may be an intermediate phase when this connection with the ground has deteriorated and the stem is only
receiving modest nourishment via the anchoring roots. In the closely related Monstera genus, the size and
shape of leaves changes with progressive life stages in ways that suggest responses to shifting light conditions
and transpiration demands (Ray, 1990, 1992). Heteropsis maintains a monopodial growth pattern throughout
its development (Hoffman, 1997) and does not go through dramatic heteroblastic leaf development as
Monstera does, but its leaves can go through minor changes in shape as it matures (Croat, 1988). Finally when
absorbing roots succeed in becoming established in the ground, the plant’s access to a more abundant source of
water and nutrients in the soil is firmly renewed. Its leaves can grow larger with the expectation that their
transpiration potential will not be overwhelmed by warmer and drier conditions they are exposed to higher on
the trunk.
The root cutting experiments done with Heteropsis in Guyana and Tekohaw and similar tests
conducted with a Monstera vine (Simmonds, 1950) show that a sudden severing of these conduits can lead to
stress or death of the plant. An anatomical study of the roots of three species of Cyclanthaceae that have
similar life histories to some Araceae vines showed that absorbing roots often have ten times as many
treacheary and sieve elements and nonperiferal xylem and phloem fascicles (bundles) as anchor roots (Wilder
and Johansen, 1992). Absorbing roots may be especially important to secondary hemiepiphytic aroids since
they deliver water and solutes from the ground to the main stem (Wilder, 1992) and may help the plant store
water and reverse embolisms in the xylem (López-Portillo et al., 2000). The sudden loss of the thickest
absorbing roots would, therefore, leave any remaining thin absorbing roots and anchor roots ill-equipped to
fulfill these critical functions for the plant by themselves.
The most obvious potential impact of conventional root harvesting from titica plants would be that the
large amount of force applied by a harvester to break thick aerial roots would dislodge the main stem of the
plant before the root gave way. In practice this rarely happens. I only saw a harvester pull down the main
stem of a titica plant one time in the course of witnessing hundreds of plants being harvested. Likewise
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Hoffman (1997) and Durigan (1998) only saw main stems pulled loose by harvesters a few times during their
repeated observations of titica root collection.
Hoffman’s (1997) cutting experiment with H. flexuosa in Guyana (1997) found that 33% of cut roots
had some regrowth six months later while 44% were still alive but had no regrowth. This percentage of
regrowth was twice as great as found in the Tekohaw cutting study where an average of 17% cut roots had
some new roots growing near the cut portion. The average number of new roots sprouted per cut root with
regrowth in the Guyana study (Hoffman, 1997) was 1.7 – identical to the finding at Tekohaw, but these new
roots were generally smaller than the root that had been cut. My finding that about 60% of the new roots were
still commercial size seven months after being cut conforms well with studies of Cyclanthaceae vines that
showed only one replacement root (Figure 5.2; Appendix 5-A) arises from an aborted apice (forming a node)
or above the excised point of a parental root. This replacement root is generally a little narrower than the
parental root, but it is thicker than other non-replacement first and higher-order roots that may also emerge
from parental roots (Wilder, 1989, 1992).
Hoffman (1997) measured the length of new roots that grew from H. flexuosa roots six months after
they were cut and found that each cut root with regrowth produced an average of 20 cm of new roots per month
or 240 cm per year if this rate held steady. Since an average of 2.5 roots were cut per tree and 33% of those
had regrowth, the average amount of titica root regrowth per host tree would be about 198 cm per year. This
was again a more robust response than found at Tekohaw where the estimated growth of all new titica roots on
all host trees where roots were cut was about 77 cm per host tree per year. Only 60% of this growth (46 cm per
year) would contribute to the production of new commercial quality roots. Hoffman (1997) found that
immature absorbing roots growing down under natural circumstances grew 26 cm per month – 30% faster than
the new roots emerging from cut roots so longer term studies are needed to properly estimate the regrowth of
roots from harvested titica plants.
One tempting conclusion of this comparison between regrowth rates of cut titica roots in Guyana and
Tekohaw is that cutting a smaller percentage of roots in the Guyana experiment left those plants with more
resources to survive the cutting and regrow more often and faster. The fact that the Guyana plants also had a
higher number of roots per host tree to begin with indicates that genetic and/or environmental differences may
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be just as responsible for their greater resilience to root cutting as much as differences in the cutting treatments.
Harvesting impacts need to be further explored with a wider range of treatment conditions at the same site or
multiple sites to better address this issue.
Root Transformation During Commercial Preparation
The Tembé practice of removing the cortex from titica root stems before sale is required by almost all
regional dealers. This is distinct from the practice in Guyana where harvesters generally sell bundles of H.
flexuosa with the cortex intact. Removing the cortex is done to give the product more aesthetic appeal and
flexibility in making furniture and handicrafts. Doing so, however, leads to a significant transformation in the
weight of the roots collected in the forest (Figure 5.9) and may leave the cleaned roots prone to mildew
(Hoffman, 1997). When used as a lashing material in house construction, Tembé leave the titica root cortex on
because removing it takes considerable time and it has greater durability in that form when exposed to the
elements.
Harvester Daily Yield and Time Involved in Collecting and Processing Titica Roots
In examining the data on search times and distance between host trees it should be remembered that
the harvester spotted and sometimes inspected other trees with titica roots that were not harvested.
Consequently the search time between host trees with any titica plant would be less than the value reported.
The actual distance between all host trees would also be less than reported, but the distance walked between
the host trees harvested would be greater than this value since the search pattern between trees was often
circuitous. With their well-honed search images for titica root, some Tembé harvesters can no doubt spot a
vine laden tree from a considerable distance, but the dense forest mandates that most host trees that are
harvested are spotted from close proximity. If 10 meters is a reasonable estimate of the harvester’s reliable
search path width, his perception of the forest is that it has only 29 titica host trees and 122 roots per ha.
Assuming that the total titica host tree and root densities in the forest area being searched by the harvester and
the adjacent survey plots is similar, the lower densities estimated from harvester data are at least partially
explained by his greater selectivity of host trees and roots. It may also mean that the harvester is not locating
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many titica plants that are hidden on the far side of host trees or otherwise obscured from view. While
harvesting most of the commercial quality (i.e. thick) absorbing roots from selected host trees may have
negative consequences for the harvested titica plants (Hoffman, 1997; Durigan, 1998), the probability that
harvesters do not systematically harvest the most robust roots from all titica plants in an area means that a
sizeable number are left unharvested – at least until another harvester searches the area. Accompanying the
commercial harvester at Cajueira showed, however, that he quickly moved through an area that had been
recently harvested (as evidenced by cut and broken roots around a host tree) until he found another area that
seemed to have a high concentration of unharvested plants.
The involvement of family members in removing the nodes and cortex from titica roots is a practice
that has economic and social importance for the indigenous harvesters. Although young Tembé children are
not generally expected to contribute directly to their family’s sustenance, helping out with titica root stripping
is one activity where young children can directly help generate income for their family and save adults time to
do other activities that they are uniquely capable of doing. Measurements of titica processing rates show that
by the time children get to be 10 years old, they have attained almost the same level of root stripping efficiency
as their elders.
The social and economic aspects of titica harvesting are quite different in Guyana where the market
readily accepts titica roots with the cortex on (Hoffman, 1997). Although dealers there seem willing to pay
more for roots that have been stripped, the market (and perhaps social) structure) does not support this.
Titica Vine Market Structure and Economics
While titica harvesting is the most important source of income for many Tembé and other forest
peoples in Brazil (Durigan, 1998), the hard work is poorly remunerated. As an example, the harvester whose
efforts were closely monitored at Cajueira spent two days traveling there and back from Tekohaw and five
days harvesting to bring back about 168 kg of rough root stems. He and his family spent another three days
removing, nodes, bad parts and the cortex. Once these cleaned stems had dried for a week, they were left with
about 33 kg of roots to sell for about $ 1.00 per kg. This means he made about $33.00 for ten days of his time
(not including his family’s labor), about $US 3.30 on average per day – just under the Brazilian minimum
197
wage at the time. These earnings are lower than are generally obtained by harvesters in Guyana who can make
$US 5 to 10 per day in the forest (not including travel time and expenses) by collecting one or two bundles (an
estimated 10 to 20 kg) of prime quality roots and selling them without having to remove the cortex (Hoffman,
1997).
A simple average value of titica harvest by area can be estimated by assuming there are 100 host trees
per ha with commercial vines and that each of these trees will yield 0.5 kg of titica roots. This means that each
hectare will yield 50 kg of harvestable unprocessed roots – or 10 kg of commercial roots. These will sell for
about $ 10. This is a better per hectare value than can be obtained from other NTFPs examined in this study,
but it should be remembered that these harvest levels are derived from harvests from forests that have not been
harvested for a long time and would not be able to be harvested again at this level for many years to come.
Considering the high mark up between the cost of the raw material and the finished product, it is
tempting to contemplate the extra income Tembé or other harvesters could earn if they made and possibly sold
titica based furniture themselves since they possess an aptitude for working with this material. There are
several challenges that could make this a problematic venture. First, it is much easier to transport large batches
of straight vines than bulky furniture on small boats or vehicles. Second, the furniture dealers in Belém only
buy vine roots - not finished product. Troy and Harte (1998) provide additional details about the wicker
furniture businesses established in Belém, which indicate it might be possible for rural artisans to increase sales
of handicrafts made from titica and other vines, but there would be significant marketing challenges. Indians
or other harvesters would need to rent their own space in the city which would incur considerable costs and
marketing expertise. Since much of the local business is based on custom orders, there would also need to be
an effective means to link customers with artisans to discuss customer needs. Current communication systems
do not favor this. The one option for expanding the profitability to harvesters in the short-term would be for
them to explore markets for the vines in more distant cities such as Fortaleza where furniture making is also
apparently prominent. It is possible that the higher prices paid for delivering debarked roots to these cities
would justify the extra cost and logistics of bringing them beyond Gurupi or Belém.
198
CONCLUSIONS: CHALLENGES FOR SUSTAINABLE TITICA HARVEST
Like many NTFPs, harvesting of titica vines presents a challenging dilemma. It is a plant that has
proven useful to Amazonian forest peoples as a vital material in building structures and making other objects
for every day use. The vines are abundant in many primary forests, and minor root harvesting for personal use
would not make an appreciable dent in their populations. The increased demand for titica roots as a prime
material for making wicker furniture has created a very welcome source of cash income for forest peoples, but
this has also placed a much greater pressure on titica resources in the forest. In Guyana, commercial harvest of
Heteropsis is generally limited to drop roots that are only found on 27% of host trees and then only comprise
an average of 31% of the mature roots. By contrast Tembé Indians and other Brazilian titica harvesters
generally take most thick trunk and drop roots they encounter. The fact that individual harvesters do not find
every host tree in an area and spare non-commercially viable roots provides some buffer for titica populations
in areas that are lightly harvested. It is evident, however, that commercial harvesting near the largest villages
in the reserve is no longer viable, so these activities have moved to smaller villages and more remote areas. A
great deal of forest interior will remain unharvested because it is not practical to carry heavy vine bundles more
than a few kilometers to the nearest river so the species itself is not endangered. Without some form of
proactive management of titica harvesting, though, the most accessible areas will probably be exhausted as a
commercial source of vines in relatively few years.
Considering his measurements and analysis of H. flexuosa stem growth, root growth and maturation
rates, Hoffman (1997) estimates it takes 20 to 66 years for one titica plant to go from the climbing stem stage
to the point when the first absorbing root reaches maturation in the ground. This long time horizon suggests
three possible management strategies. Since there are strong indications that current harvesting practices in the
eastern Brazilian Amazon have a deleterious effect on titica plants, one option would be to encourage or
require harvesters to reduce the number or percentage of roots taken from host trees. In order to frame this
recommendation on a scientific basis, longer term research will be needed to establish the rate at which
harvested roots grow back and the rate at which new absorbing roots are formed on branches from the main
stem. This information could help determine the number and frequency at which absorbing roots could be
199
harvested without jeopardizing the whole plant’s well-being. Hoffman (1997) recommends that even the
modest harvesting rate in Guyana should be reduced by half (to about 1.25 drop roots per host tree) to reduce
harsh impacts on individual plants and preserve a future harvest opportunity. Implementing a more
conservative harvesting practice where access to primary forest is not strictly controlled would be challenging.
If one harvester pulled down less than all available commercial roots, the next person to come along would be
inclined to take the rest. It would take understanding and commitment among a group of harvesters in an area
of common property (such as an indigenous or extractive reserve) to abide by such limits.
Traditional beliefs about natural history of the forest can often provide the basis for sound
management of resources without the need to interject an outside scientific interpretation. In the case of titica,
however, some of these beliefs may make adoption of stricter harvesting norms more difficult. Tembé men
who have lived in the forest their whole lives stated that the slender stems of titica that had only ascended
several meters up a host tree were a different type of titica than the plants whose roots they harvested. Given
their secondary hemiepiphytic habit and the possibility that these original climbing stems disintegrate before
aerial roots of an adult plant descend, it is understandable why the two life stages would not be connected.
Another belief of some Tembé and other Amazonian indigenous groups is that the aerial roots of titica and
other common secondary hemiepiphytic vines are generated from the extending legs of different species of
dead ants (Lenko and Papavero, 1979). These stories principally come from rich mythology concerning the
painful stinging tucangueira ant (Paraponera clavata). The link between these and other ants and the
descending roots of some vines may derive from the observation that after ants infected and killed by
Cordyceps and Hirsutella fungal parasites exhibit stringy mycelia that sprout from the ant’s corpse and
resemble young brown roots (Nelson and Papavero, 1979; Luis Diego Gomez, personal communication 2000).
Tembé understand the biological significance of fruits and are intimately familiar with ones that serve as
sources of food for people and terrestrial game animals. The reproductive structures of titica, however, are
generally not familiar even to men who frequently harvest the roots. This may be because the fruits are usually
out of sight and are likely dispersed in the canopy by animals such as bats and primates (Madison, 1979b;
Vieira and Izar, 1999). It may be arrogant to challenge traditional beliefs, but it seems that harvesters who
appreciate the long time span and stages involved in vegetative growth and reproduction of titica plants would
200
be more open to developing alternate harvesting practices that would preserve an important economic and
subsistence resource.
The second management option would be to treat titica harvests more like a logging operation where
the expectation is that most harvested plants will die and future harvests will depend on waiting for the target
species to regenerate to harvestable size. If this system was adopted, harvesting zones could be established
where roots could be collected on a rotating basis – possibly with zone limits established after pre-harvest
inventories. The area would remain closed to further titica root collection until periodic monitoring indicated
that the population and commercial root density had sufficiently recovered. Some of the same challenges
described for management option one would also exist for this system, but it might be easier to enforce within
a community than a tree by tree harvesting standard. Enforcing protection of forest areas from outsiders is
another problem that has already reduced the titica resource base in the Tembé reserve. As forest clearing
progresses in the eastern Brazilian Amazon, the invasion of indigenous and other protected areas by settlers
and loggers has become increasingly acute. The Brazilian Natural Resource Agency (IBAMA) has fined nonIndians for harvesting titica vines (and timber) in the Alto Rio Guamá Reserve, but such enforcement actions
are rare in relation to the extent of such illegal harvests (Potiguara, personal communication 1998).
The third option for managing titica could operate in situations where logging or forest clearance was
a certainty. If it was known that a significant amount of a forest canopy was going to be opened by some
activity, it seems likely that titica plants would not survive well in such open environments. In the case of light
individual tree removal, additional research could show what disturbance thresholds impact titica plants on
remaining trees. Where major canopy removal was anticipated, it would make sense to fully harvest roots of
titica plants in these areas before the area was logged or cleared. Fruits of reproductive titica plants could be
collected when possible and used for germination tests. It would be logical to test planting titica in both intact
forest and advanced secondary forests where non-pioneer species had become established and would be
allowed to grow for the foreseeable future. If Hoffman’s (1997) lower-end estimate of titica maturation rates
proves to be correct, integrating titica vines and root harvest into a long-term timber rotation (50 years or
longer) might be ecologically feasible. It’s hard to say at this point if the demand for titica will get to the point
where developing more sophisticated silvicultural approaches would be economically viable.
201
TABLE 5.1 SUMMARY OF TITICA STUDY OBJECTIVES, METHODS AND
ANALYSIS
TITICA BIOLOGY AND POTENTIAL HARVEST OF AERIAL ROOTS
OBJECTIVE
SITE AND SAMPLE MEASUREMENTS
SIZE
Density of Titica Host
Tekohaw Heavy-use
1) Number of host trees
Trees and Roots
Site (1.2 ha unburned;
with a titica root per
0.5 ha burned)
250m2 plot
Tekohaw Medium-use
Site (0.6 ha)
Number and Type of
Titica Roots per Host
Tree
Dimensions of
Different Root Types
Titica Relation to Host
Tree Type
Relation of Tree DBH
to Titica Colonization
Relation of Host Tree
DBH to Titica
Climbing Stem and
Root Number
Root Maturation and
Natural Mortality
Cajueira Light-use Site
(0.6 ha)
Tekohaw Medium-use
Site (171 host trees)
Cajueira Light-use Site
(306 host trees)
Cajueira Light-use Site
(73 host trees)
Tekohaw Heavy-use
Site (titica survey 1.7
ha; all tree survey 0.9
ha)
Tekohaw Medium &
Cajueira Light-use
Sites
Tekohaw Heavy-use
Site (titica survey 0.3
ha; all tree survey 0.2
ha)
Tekohaw Medium-use
Site (143 host trees)
Tekohaw Medium-use
Site (Immature Roots
on 31 host trees;
Mature Roots on 39
host trees)
2) Number of roots
(immature, mature,
commercial) per host tree
Number of roots
(immature, mature,
commercial) per host tree
Length, width and number
of nodes per root
Host tree common name;
Common names of all
trees ≥10 cm DBH
Common names of titica
host trees in density
inventories
1) Titica host tree DBH
2) All tree DBH
1) Titica host tree DBH
2) Number of climbing
stems and roots per host
tree
Track color coded
immature and mature
roots for 7 months and
note transitions to
maturity or death
ANALYSIS
1) Multiply average
number of host trees
per plot by 40 to
estimate density per ha
2) Multiply average
sum of roots per plot by
40 to estimate density
per ha
Compute combined
mean number of roots
of different types per
host tree
Compute and compare
average dimensions for
commercial and noncommercial roots
Compare relative
abundance of titica host
trees with all trees in
the area
Estimate number of
host tree
morphospecies per site
Compare % of host
trees in each DBH size
class to % in all tree
survey
Compute mean number
of total roots and
commercial roots per
host tree
Calculate % of
immature roots that die
or mature and % of
mature roots that die
RESULTS
PRESENTED
Figure 5.3 and text
Figure 5.4 and text
text
text
text
text
Figure 5.5 and text
Figure 5.6 and text
202
TABLE 5.1 (Continued)
TITICA HARVEST IMPACT, YIELD AND SHORT-TERM ECONOMICS
OBJECTIVE
SITE AND SAMPLE MEASUREMENTS
SIZE
Effect of Two Cutting
Tekohaw Medium-use 1) Monitor condition of
Treatments on Root
Site (37 host trees with roots (dead, live, live with
Survival and Regrowth
50% commercial roots regrowth) 7 months after
cut; 56 host trees with
cutting.
100% commercial
2) Measure number and
roots cut))
width of new roots arising
from cut roots.
Root Length and
Cajueira Light-use Site 1) Number and length of
Weight Loss in
(37 host trees)
roots per host tree before
Commercial Processing
node removal
2) Number and length of
root stems per host tree
after node removal
Tekohaw Medium-use Weight of roots after node
Site (20 host trees)
removal, cortex removal
and each day of drying
Harvester Daily Yield
Accompanied
1) Host trees, whole roots
and Search Pattern
Comercial Harvester
and root stems collected
near Cajueira for 5
per day
days
2) Host tree GPS position
Root Processing Rate
Observed 9 family
members processing
titica roots at Tekohaw
Value of Titica Roots to
Harvester
Inventories at
Tekohaw and Cajueira
Value of Titica Roots to
Forest Product Dealers
and Wicker Furniture
Makers
Interviews with forest
product dealers and
furniture makers in
Gurupi and Belém
1) Starting and stopping
time for processing each
bundle of roots
2) Initial and final weight
and stem number of roots
per processed bundle
3) Age of processor
1) Buying and selling
prices for titica roots
2) Weight of titica roots in
common furniture pieces
ANALYSIS
1) Compare % cut roots
dead, alive, and alive
with regrowth between
treatments.
2) Estimate regrowth
rate of commercial
quality roots.
Calculate average %
loss in length and
increase in root stem
number that
accompanies initial
commercial processing
Percentage weight loss
after each stage of
processing
1) Calculate average
number of host tree
visited and roots
collected per day
2) Calculate average
distance between
harvested host trees
3) Plot host tree search
pattern
1) Calculate overall
average of root stems
and root weight
processed per minute
2) Compare root
processing rate by age
of processor
1) Calculate value of
titica roots per ha
2) Calculate value of
titica harvest per day
collecting and
processing
Compare value of titica
to collector with
dealers and furniture
makers
RESULTS
PRESENTED
Figure 5.7 and text
Figure 5.8 and text
Figures 5.9 & 5.10
and text
Figures 5.11 &
5.12 and text
Figure 5.13 and
text
text
text
203
a. Absorbing Trunk Root b. Absorbing Drop Root c. Absorbing Underground Root
d. Main Climbing Stem with Anchor Roots e. Lateral Branch with Leaves and Fruit
f. Senescing End of Climbing Stem g. Seedling Climbing Stem
FIGURE 5.1 ILLUSTRATION OF TITICA SEEDLING AND ADULT ON HOST
TREE WITH PRINCIPAL PLANT PARTS
204
a. Absorbing Root Node
b. Absorbing Replacement Root
d. Aborted Absorbing Root Apical Meristem
c. Non-replacement Root
FIGURE 5.2 ILLUSTRATION OF TITICA ABSORBING ROOT AND
PRINCIPAL PARTS
205
Interval Plot: M ean plus 95% Confidence Interval
1800
TITICA ROOTS
PER HA
DENSITY
1500
1200
900
Tekohaw
Heavy Use Site
(1.2 ha Surveyed)
Tekohaw
Medium Use Site
(0.6 ha Surveyed)
Cajueira
Light Use Site
HOST TREES PER HA
WITH TITICA ROOTS
(0.6 ha Surveyed)
600
300
0
Heavy
M edium Light
Heavy
M edium Light
INTENSITY OF PREVIOUS SITE USE
FIGURE 5.3 DENSITY OF TITICA HOST TREES AND TITICA ROOTS IN
LIGHTLY TO HEAVILY USED FOREST SITES IN THE ALTO RIO GUAMÁ
INDIGENOUS RESERVE
206
MEAN NUMBER OF
ROOTS PER HOST TREE
7
ROOT TYPE
53% Commercial
6
5
4
M ature
Commercial
M ature NonCommercial
27% Commercial
Immature
3
2
1
0
HOST TREES
WITH ANY
TITICA ROOT
HOST TREES WITH AT
LEAST ONE TITICA
COMMERCIAL ROOT
(N=477 Host Trees)
(N=150 Host Trees)
FIGURE 5.4 NUMBER AND TYPE OF TITICA ROOTS PER HOST TREE AT
MEDIUM AND LIGHT-USE FOREST SITES AT TEKOHAW AND CAJUEIRA
207
TITICA ROOTS PER HOST TREE
Interval Plot: M ean plus 95% Confidence Interval
7
6
All Roots
5
4
3
2
1
Commercial Roots
0
4-9
10 - 19
20 - 29
30 - 39
>39
HOST TREE DBH SIZE CLASS (cm)
(143 Host Trees with Titica Roots)
FIGURE 5.5 RELATIONSHIP BETWEEN HOST TREE DBH AND THE
NUMBER OF TITICA ROOTS PER HOST TREE AT THE TEKOHAW
MEDIUM-USE SITE
208
IM M ATURE TITICA ROOT DEVELOPM ENT PHASE
Live Immature
Dead Immature
New Live M ature
PERCENT OF ROOTS IN
DEVELOPMENT PHASE
100
75
50
25
0
Start
76
134
210
DAYS AFTER START OF OBSERVATION
N = 56 Immature Titica Roots from 31 Host Trees at Tekohaw
FIGURE 5.6 DEVELOPMENT TIME AND MORTALITY OF IMMATURE
TITICA ROOTS AT THE TEKOHAW MEDIUM-USE SITE
209
MEAN PERCENT RESPONSE OF
TITICA ROOTS PER HOST TREE
CUTTING
TREATM ENTS
50% Commercial Roots Cut (37 Host Trees)
100% Commercial Roots Cut (56 Host Trees)
70
Mean + 95% Confidence Interval
60
50
40
30
20
10
0
50%
100%
CUT ROOTS
DEAD
50%
100%
50%
100%
CUT ROOTS
CUT ROOTS
ALIVE
WITH REGROWTH
FIGURE 5.7 CONDITION OF TITICA ROOTS 7 MONTHS AFTER CUTTING
AT TEKOHAW MEDIUM-USE SITE
210
LENGTH PER TITICA ROOT
AND ROOT STEM (m)
ONE HOST TREE AT CAJUEIRA YIELDED
AN AVERAGE OF 2.9 COM M ERCIAL
QUALITY TITICA ROOTS WITH NODES
5
4
(M ean Total length: 13.6 m)
(M ean Total length: 8.9 m)
3
2
1
0
THESE ROOTS LOST A M EAN OF 30%
TOTAL LENGTH WHEN NODES WERE
REM OVED AND YIELDED 5.8 ROOT STEM S
FIGURE 5.8 LOSS OF TITICA ROOT LENGTH AFTER NODE REMOVAL
FROM 37 HOST TREES HARVESTED AT CAJUEIRA
211
Interval Plot: M ean plus
95% Confidence Interval
MEAN WEIGHT OF
ROOTS PER TREE (g)
740
640
Commercial Titica Roots
Harvested from 20 Host
Trees at Tekohaw Medium
Use Site
540
440
340
240
140
Roots pulled Nodes & bad
from tree
pieces removed
Cortex
removed
8 days after
cortex removed
STAGE OF ROOT PROCESSING
FIGURE 5.9 WEIGHT LOSS DURING PROCESSING OF TITICA ROOTS
212
PERCENT WEIGHT LOSS
OF STRIPPED ROOTS
Interval Plot: M ean plus 95% Confidence Interval
30
20
10
1
2
3
4
5
6
7
8
DAYS OF ROOT DRYING WITHOUT CORTEX
Commercial Titica Roots Harvested from 20
Host Trees at Tekohaw M edium Use Site
FIGURE 5.10 WEIGHT LOSS OF TITICA ROOTS FROM DRYING AFTER CORTEX
REMOVAL
NORTH AXIS POSITION (meters)
213
Day
Day
Day
Day
4500
4400
4300
1
2
3
4
4200
4100
4000
3900
3800
3700
3600
100
200
300
400
500
600
700
800
900
EAST AXIS POSITION (meters)
FIGURE 5.11 LOCATION OF TITICA HOST TREES HARVESTED IN 4 DAYS BY A
COMMERCIAL COLLECTOR AT CAJUEIRA
NORTH AXIS POSITION (meters)
214
4000
31
32
27
28
26
29
20
30
22
3900
25
19
24
23
18
17
21
3800
5
12
1
3700
3
2 6
8
15
9
13
16
7
14
10
11
4
400
500
600
700
EAST AXIS POSITION (meters)
FIGURE 5.12 LOCATION AND ORDER OF TITICA HOST TREES HARVESTED BY
A COMMERCIAL COLLECTOR AT CAJUEIRA ON ONE DAY
ROOT STEMS STRIPPED PER MINUTE
215
Interval Plot: M ean plus 95% Confidence Interval
1.5
Titica Roots Collected by O ne
Harvester at Cajueira and Processed
by his Family at Tekohaw
1.0
0.5
4-6
7-8
n=(2)7
n=(2)12
10
13
n=(1)24
n=(2)59
25
n=(1)7
42-44
n=(2)36
AGE OF TITICA PROCESSOR (Years)
N=(Number of people)Number of Root Bunches
FIGURE 5.13 TITICA ROOT STEM STRIPPING RATE BY AGE OF PROCESSOR
216
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APPENDIX 5-A GLOSSARY OF TERMS FOR TITICA VINES AND ROOT HARVEST
PLANT GROWTH HABITS
CLIMBER/VINE – A plant that is rooted in the ground but needs support for a weak stem (Kelly, 1985).
TRUE EPIPHYTE – A plant whose growth is restricted to growth on trees, shrubs, or rocks (2).
HEMIEPIPHYTE – A plant whose natural life cycle includes both an epiphytic and terrestrial phase. It excludes
woody lianas and vines that germinate terrestrially and maintain lifetime contact with the ground (3, 5).
PRIMARY HEMIEPIPHYTE – Plants that start as epiphytes in a tree and later become rooted in the ground (5).
SECONDARY HEMIEPIPHYTE – A plant that is hybrid of a climber and an epiphyte because it germinates in the
ground, ascends a tree and then loses contact with the ground. Some plants including titica produce
adventitious roots that later reenter the soil (2, 5).
GROWTH STAGES
SEEDLING – A plant with one apical shoot meristem on a seedling shoot that grows along the forest floor. It
reaches a more advanced stage when it finds a host tree and a climbing stem begins to ascend it (4, 6).
JUVENILE – A plant whose climbing stem has produced lateral leafy branches without reproductive structures (4).
ADULT – A plant with multiple leafy lateral branches capable of producing a terminal inflorescence. Absorbing
roots have arisen from the main climbing stem or lateral branches and descended into the ground. Flagellar
shoots may also descend from branches and reclimb the same host tree or grow along the ground in search
of a new one (4, 6).
CLASSIFICATION OF ROOTS
ANCHORING ROOT/CLIMBING ROOT – A short and narrow aerial root whose primary function is to secure a
climbing stem, main plant stem or lateral branches to the host tree (6, 7).
ABSORBING ROOT/FEEDER ROOT – A long adventitious root that grows from a climbing stem or lateral branch
toward the ground. Its main function is to provide water and nutrition to the main stem once it has
entered the ground (6, 7).
MATURE ROOT – An absorbing root that has entered the ground and presumably has extensive growth in the soil.
Its aerial portion has a hardened outer layer (1)
IMMATURE ROOT – An absorbing root that is still descending or has not yet grown extensively in the soil. When
a finger nail is scraped over the outer layer of its aerial portion, a greenish color is revealed underneath (1).
REPLACEMENT ROOT – A root that grows back from an anchoring or absorbing root when the original root
apical meristem aborts or is damaged through mechanical injury or herbivory. Replacement roots continue
growing in the direction of the parent root and are about the same thickness (4, 6).
NONREPLACEMENT ROOT – A root that grows off of any portion of an anchoring or absorbing root. They
generally grow in a different direction and are narrower than the parent root (6).
ROOT NODE – A bulge in an absorbing root that occurs just above the point of an aborted apical meristem or
injury. Replacement roots frequently emerge just above a node (4)
ROOT INTERNODE – The portion of an absorbing root between two successive nodes (4).
TRUNK ROOT – An absorbing root that descends in close contact with the host tree trunk and other vines (4).
DROP ROOT – An absorbing roots that grows straight down from its attachment point on the plant stem (4).
COMMERCIAL ROOT – A mature absorbing root that is fit for sale to wicker furniture makers. The criteria varies
regionally, but such roots are judged to be relatively thick (≥ 3.5 mm) and have relatively few nodes (1).
COMMERCIAL STEM – A section of a mature absorbing root that has been separated from nodes and in some
cases stripped of its outer layers (epidermis and cortex). In the eastern Brazilian Amazon, such stems are
generally at least 3.0 mm thick and at least 1 meter long although most are considerably longer (1).
References: 1) Author Description; 2) Croat, 1988; 3) Gentry, 1991; 4) Hoffman, 1997; 5) Putz and Holbrook, 1986;
6) Ray, 1992; 7) Wilder, 1992; 8) Wilder and Johansen, 1992
CHAPTER 6
SUMMARY CONCLUSIONS
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FACTORS THAT AFFECT NON-TIMBER FOREST PRODUCT HARVEST
AMOUNTS AND PROFITABILITY
The first and second objectives of this study were to quantify the amount of marketable non-timber forest
product (NTFP) that could be obtained per plant and per area of forest and identify key factors that influenced
variation in these amounts. The case studies showed that the size of the resource base and available product were
most influenced by the density of the plant, plant size, land use practices (e.g. fire), and sometimes plant
interactions with insects or microbes. Once products were harvested, the amount of saleable product that an area
of forest would yield depended on the conversion efficiency of the raw product to commercial product.
The third objective of the study was to estimate the amount of harvest that would be possible on a
repeated basis and the length of this harvest cycle. The case studies and work done with other NTFPs in the study
area showed that some plants could be harvested several times a year while other plants might only yield
marketable quantities of product once in their life. The amounts varied depending on what material was removed
from the plant and how fast it regenerated.
The fourth main study objective was to estimate the amount of time a harvester needed to spend to find,
harvest and process a unit of product and how much income they would earn for the time invested. The case
studies found that search time was directly related to plant density and the proportion of harvestable plants within
the population. Products that were obtained from the outside of trees (resin lumps on tree trunks, vine roots on
host trees, seeds on the ground) were collected much more quickly than products that required drilling or scraping
trees to procure interior products (latex and oleoresin in heartwood). The time devoted to procuring raw materials
was much greater if collectors needed to travel far from their villages to find concentrated populations of
harvestable plants. The degree of processing raw products into commercial products varied extensively between
products and took as much or more time for some products as the time required to find and harvest the product in
the forest. The amount of income that harvesters could earn from each NTFP of course also depended on the
demand for the product and its market price. Among the products examined, there was a 10 ten fold difference
between the unit price of the least expensive to the most expensive, but the most important sources of revenue for
Tembé NTFP harvesters were from products that were the most abundant – not the most expensive.
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SIZE OF RESOURCE BASE AND AVAILABLE PRODUCT
The most important factors that influenced the size of the resource base were the density of the plant and
the percentage of those plants that offered harvestable product. Among the three case studies in this thesis, titica
vines were by far the most abundant since they were found on several hundred host trees per ha, and at least 50
plants per ha offered harvestable aerial roots (Table 6.1). This number was two orders of magnitude greater than
copaiba trees whose density was close to 0.5 trees per ha. The density of the case study trees varied with habitat
changes in the Tekohaw study area. Titica vines, and to a somewhat lesser extent, copaiba trees were found
almost entirely in upland “terra firme” sites. Breu trees were found almost equally in “terra firme” and wetter
“baixo” type forests, but different species were found more commonly in one habitat or the other.
Larger and presumably older individual plants consistently had a harvestable product more often and
offered larger amounts per plant among the NTFP case studies. The percentage of the size range that offered
harvestable quantities of product from each NFTP, however, varied considerably. Copaiba trees had a fairly
narrow size range of trees from which oleoresin was reliably obtained, so only 15% of the trees could be
considered productive from a harvester’s point of view. In contrast, the probability of finding resin on breu trees
steadily increased with their size, and overall 40% of them offered harvestable product. My studies conducted
with trees that produced “amapá” latex showed that Couma guianensis Aublet (called “black amapá” by the
Tembé) was an exception to this pattern. The amount of latex collected from these trees had no correlation to tree
size throughout the five week period of their harvest (Appendix 6-A).
A marketable product sold by one name may come from a variety of species. This was most apparent
with Burseraceae trees where some types of trees had considerable resin while others had none. Among those
species that were commonly harvested, however, there was no significant average difference in resin yield.
Additional research on specific species rather than loosely defined groups will probably reveal more definitive
differences. The difficulty of trusting common names to reflect distinct taxa was also apparent with amapá latex.
The Tembé used the term amapá to refer to both Parahancornia amapa (Huber) Ducke and C. guianensis trees in
the Apocyanaceae family and used their latex interchangeably as a medicinal tonic. The latex from Brosimum
potabile Ducke(family Moraceae) was also frequently sold in Belém and other parts of the Amazon as “amapá”
(van den Berg, 1984).
223
Land use practices and site history also had a strong influence on the size and productive potential of the
NTFP resource base. A fire that passed through the Gurupi section of the reserve in the early 1980s burned
extensive swaths of forest within several kilometers of the village. Almost 20 years later, though, titica vines are
still almost absent from these areas that are dominated by pioneer tree species. Some copaiba trees that were
burned in the fire survived, but their wounds apparently led to an early hollow condition and consequent loss of
oleoresin storage capacity. The Tembé are also losing part of their resource base for wild populations of NTFPs as
forests are cleared for agriculture and adjacent forest areas are burned by fires escaping from farm plot
preparation. As forest burning related to logging, ranching and agriculture increases throughout the Amazon
(Nepstad et al., 1999), there could be a corresponding direct and indirect depletion of the region’s NTFP resource
base.
Changing settlement patterns among the Tembé could also affect their access to both commercial and
subsistence types of NTFPs. Some villages in the Gurupi area have received additions to their physical
infrastructure in the form of running water systems and construction of health clinics and school buildings. More
people have been attracted to these villages and they are taking on a more permanent character than in the past
(Potiguara, personal communication 1999). The apparent result of this pattern is that more forest close to the
village has been converted to agriculture and people need to go farther to hunt, fish, and collect forest products. It
was already noted in the case studies that breu and titica harvesting trips were conducted hours away from
Tekohaw. People also reported that close to the village it was getting harder to find adequate supplies of the palm
frond “ubim” (probably Geonoma baculifera Kunth. (Balée, 1994)) used as the primary roof thatch material and
long stems of the herbaceous “guarimã” plant (probably Ishnosiphon spp.) used for making baskets. In response to
the shortage and relatively short useful life of “ubim” (two to three years), more people were cutting down trees to
make wooden shingle roofs.
As people in the village buy more electronic goods and gas stoves, the need for more batteries and fuel
increases. This requires more trips to the city necessitating greater gasoline and diesel fuel consumption and more
frequent boat engine repairs. All of these things, of course, cost more money that leads to greater pressure on
natural resources. Such depletions and shifts in resource use that accompany acculturation have been noted with
other indigenous groups (Robinson and Redford, 1991; Baksh, 1995; Stearman, 1995). This issue will no doubt
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pose a mounting challenge for the Tembé and other people who wish to maintain access to the full range of their
traditional wild plant and animal resources while they integrate more aspects of the non-Indian world into their
lifestyle. The chances that expanding sales of NTFPs in the raw form will be able to meet the increasing consumer
desires of people in this situation seem very slim.
The amount of raw material per plant is the starting point for assessing an NTFP’s productive potential,
but the amount of material that can be sold after the raw material is collected is what counts in determining its
actual yield per area of forest. Converting raw product to a saleable product can also significantly add to the
overall handling time for a particular product. Some exudates, such as copaiba and amapá latex, can be sold in
essentially the same form they come out of the tree. Breu resin had a modest conversion loss of 17% in the drying
process, while saleable titica roots that were reduced to node-free segments and stripped of their cortex and dried
only retained 20% of the original weight (Commercial Product/Raw Product Table 6.1). The most extreme
example of processing loss among the products investigated in this study was the preparation of oil from andiroba
(Carapa guianensis Aublet) seeds. In one sample prepared at Tekohaw it took an estimated 14.4 kg of seeds to
produce one liter of oil showing a conversion ratio of 3% by weight (Appendix 6-B). Even a commercial
manufacturer of andiroba oil in Belém reported he only obtained a 10% yield in his mechanized process (Morães,
personal communication 2000).
REGENERATION RATE OF THE RESOURCE AFTER HARVEST
The sale of NTFPs may generate less revenue per hectare of forest than other land uses such as logging in
the short-term but be more profitable in the long-term because the harvest cycle is shorter (Peters et al., 1989).
Fruits can be potentially collected every year with large harvests available on a more irregular basis from trees that
have a mast fruiting pattern (one year of peak fruit production out of many years with little or no fruit production).
The NTFPs examined in this study showed that the time a harvester would need to wait between successive
harvests to achieve a steady harvest level varies tremendously with the product being harvested.
Harvesting latex from amapá trees at Tekohaw showed that some products can be obtained from the same
trees on a relatively frequent basis, but there are limits to the amount of material a tree can produce in a short
period of time. C. guianensis and P. amapa trees both yielded latex twice a week for five weeks in 1999, but in
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both species the average amount collected per tree dropped off significantly by the fifth week of harvesting
(Appendix 6-A). A similar series of harvests conducted on the same trees in 1998 showed that latex harvest levels
had returned to full capacity after a year of rest. Experience with rubber tree (Hevea brasiliensis) latex extraction
has shown that an improper harvesting regimen can reduce a tree’s productive capacity (Oliveira, 1998) so
additional studies with amapá latex species are needed to determine what method and frequency of cutting are
tolerated by them.
Quantifying the appropriate harvest cycle with breu trees was complicated because resin build-up was
determined by the growth rate and colonization patterns of Sternocoelus weevils as well as the resin generation
capacity of the trees. While resin could be harvested from the same tree once or twice a year, the study estimated
that initial harvest levels could only be repeated if trees were harvested once every four to five years. There are
probably many other NTFPs based on plant defensive compounds whose production and availability are based on
insect and microbial attacks that have not yet been described.
The renewable harvest frequency for titica vines will probably be measured in decades rather than years.
This study in the Tembé reserve reinforced results of other researchers (Hoffman, 1997; Durigan, 1998) who
found that some severed roots do grow back slowly, but many roots and possibly many plants die as a result of
harvesting. It is, therefore, possible that an area that has been intensively harvested will not offer commercial
densities of titica vines again until young plants grow up and have mature roots fixed in the ground. In such cases,
calculating NTFP yields and devising sound harvesting strategies should look to timber management for models
that assume harvested plants will die and that product for the next round of harvesting will need to come from the
next generation of plants.
The copaiba case study demonstrated another NTFP where the first harvest from individual plants could
be the only harvest even if harvesting does not mortally wound the tree. This study confirmed results of research
on C. multijuga (Alencar, 1982) that showed successive harvests from the same trees conducted six months to a
year after the initial one yielded little or no oleoresin. There are many published statements that copaiba trees
yield large amounts of oleoresin on a repeated basis, but these cases have yet to be scientifically documented.
Managing NTFP species like copaiba could also benefit from long-term timber management models, but it should
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not be assumed that copaiba trees that have lost oleoresin yielding capacity have exhausted their ecological role in
their community in part because they can remain reproductively active for many more years.
HARVESTING AND PROCESSING TIME
Harvesters will obviously make more money for each day invested in an NTFP enterprise by minimizing
time spent finding plants, harvesting the raw material and processing it into a marketable form. While copaiba
oleoresin requires no processing once it is obtained, finding trees in a low density population, drilling into the
heartwood of prospective trees, and possibly returning several days later to collect the product all involve a lot of
time. Selling breu is relatively easy because trees with resin are moderately abundant, resin is usually quickly
removed from the bark and processing only requires minimal drying outside. Titica vines are the most common
NTFP in the Tembé forest area, but stripping the cortex from the roots requires as much time as harvesting them
(Processing Day/Harvesting Day Table 6.1). This low conversion ratio of freshly harvested roots to clean root
stems and the large amount of time required to carry out this processing made the yield of marketable titica
product for one day’s work (Commercial Product per Day Harvesting & Processing Table 6.1) much lower than
for breu resin. Preparing andiroba oil was the most labor consuming NTFP enterprise I observed in this study. It
first required finding and visiting fruiting trees every day to gather seeds that had fallen before too many rotted or
were consumed by wildlife. Secondly, seeds needed to be boiled, dried, dehulled, mashed and put out in the sun
daily for three to six weeks to leach the oil from them (Appendix 6-B).
MARKET PRICE AND DEMAND
This study confirmed the finding of many previous studies that the price offered to harvesters for raw or
even semi-processed NTFPs is very low in relation to the amount of time that must be devoted to procuring any of
them. The sobering result of the production ecology studies and low market prices of NTFPs examined in this
study was that collectors needed ideal and in some cases unattainable circumstances to make commercial
harvesting of these products worthwhile to them. Titica was the only plant that offered the possibility for
harvesters working in areas where roots had not been intensively harvested in recent years to earn as much as the
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Brazilian minimum wage (about $US 3.50 per day in 1999). It was, therefore, not surprising that this was the most
important economic NTFP for the Indians in the Gurupi Region. Some Tembé men considered commercial breu
collection worth their time, while others did not. Copaiba tree density and/or oleoresin yield per tree would have
to be ten times higher than current levels for a person at Tekohaw to make minimum wage collecting this
oleoresin. People who tried to make money selling andiroba oil would be the hardest pressed to make the
endeavor worthwhile. Given the $US 1.50 per liter paid for the raw oil and a 10% conversion ratio, a harvester
collecting and making the oil would need to collect about 47 kg of seeds per day to make minimum wage of $US
3.50 per day. Collectors who sell seeds directly to an oil press factory have the same collection burden since they
are paid only $US 0.07 per kg for raw seeds. This would have been impossible at Tekohaw in 1999 when
harvesters visiting 47 trees every day collected only 0.6 kg of seeds per day in the peak month of the fruiting
season (Appendix 6-B).
I did not gather information on the amount of agricultural products sold by the Tembé, but informal
interviews indicated that titica sales were a larger source of income for most people than farinha (manioc flour),
rice, or beans. Most families only planted enough manioc to meet their family’s anticipated food needs for the
year and only sold farinha if they had a surplus and the price was at least $US 7.50 per sack. At this price, they
would earn the equivalent of $US 2.17 per person per day for their efforts making this labor-intensive crop
(Appendix 6-C). This was lower than several of the NTFP case study species (Table 6.1), but the value of the
agricultural product per hectare of land was far greater than any of these NTFPs in the short-term or long-term.
The regional office of the Brazilian Federal Indian Agency (FUNAI) was very supportive of my efforts to identify
additional marketable NTFPs for the Tembé, but they have prioritized their technical assistance to help villages
increase their production of agricultural cash crops such as farinha, rice and beans (Galvão, personal
communication 1999).
RESEARCH IMPLICATIONS FOR THE ROLE OF NON-TIMBER FOREST
PRODUCTS IN FOREST COMMUNITY DEVELOPMENT
The fact that products such as copaiba oleoresin, andiroba oil, amapá latex, breu resin and titica roots are
readily available in regional and some national markets shows that the Tembé and other harvesters are willing to
228
collect them in spite of their low price. It seems that people living in the forest need or want money so badly that
they are willing to accept these prices because they perceive they have no better alternative. The Tembé do so
rather grudgingly because they are aware that middle-men and retailers are re-selling these products in the same or
slightly altered forms for four to more than 100 times as much as they were paid for them. While the need to
redress this imbalance in compensation to harvesters in the NTFP marketing chain has been repeatedly stated
(Clay and Clement, 1993; Warner and Pontual, 1994), I saw no easy ways that the Tembé could sell their products
to retail vendors or end consumers or take on value-added processing, and they do not control enough of the
resource base to simply demand higher prices.
It is clear that the Tembé are going to expand their production of agricultural cash crops as a primary
means of increasing village income. The question then remains whether or not NTFPs can provide other economic
resources. Continuing to sell breu resin and titica vine roots at current levels provides a modest but at least fairly
dependable source of cash to buy certain staples. The other product that can be taken directly from the forest that
seems to offer possible economic reward is fruit from açai (Euterpe oleracea) palm tree. In early 2000, an açai
fruit buyer sent a boat up the Gurupi River to buy several hundred kilograms of the berries. The Tembé could
conceivably take these fruits to market themselves, but the collecting and delivery would have to be highly
organized because the berries need to be ripe when picked and reach the market within 24 hours before they spoil.
It will always be easier for people living in flooded varzea forest islands near Belém to harvest and sell açai fruit
bunches from nearby trees and bring them to market only one to two hours away than it will be for the Tembé or
anyone who lives in place that requires a 12 – 16 hour boat ride and 4 hours by bus or truck to get to market. One
potential problem with such a commercial venture is that it could lead to a competition between people wishing to
harvest açai for consumption in the home and those wishing to collect it for sale. The fruits are abundant during
the peaks of the fruiting season, but in off-peak times, people travel farther from their home village in order to
collect the berries.
While NTFPs are sometimes thought of exclusively as products from wild populations, many commonly
marketed ones, particularly fruits, are the result of intentional planting. It is common for Tembé to plant cajú
(Anacardium occidentale L.)(cashew) trees in the midst of their manioc fields. After the manioc field has gone
through one or two cycles of planting, it is left fallow, but the cajú trees continue to grow amidst the pioneer trees
229
that take over the plot. The masterful timing of this agroforestry practice is that the juicy fruits ripen in these
fallow plots late in the dry season when people are laboring in the hot sun in the new fields and need liquid
refreshment. The nuts are roasted outside the village so people will not be unnecessarily exposed to the toxic
fumes that emanate from their hull. The Tembé man who has been buying breu resin and titica roots started
buying cajú nuts from villagers since he found a dealer willing to buy this product in batches of several hundred
kilograms (Kapara’i Tembé, personal communication 2000). There are many other trees and herbaceous plants
that Tembé plant in their backyard gardens and fields for personal use, so there are probably other plants with a
marketable product such as cupuaçu that could also be integrated into an agroforestry system.
The major source of income for many Tembé in Tekohaw is the sale of handicrafts (Mitchell, personal
communication 1999). These items are not always thought of as NTFPs, but since they are made with a diverse
array of plant materials including seeds, leaves, roots, fibers, wood, and resins and animal materials including
feathers, fur, and hide they should qualify even though they occasionally include synthetic materials such as
plastic beads and fishing line. One typical Tembé man might toil for three days in the forest and three more days
in the village to process 100 kg of titica roots and make $20 for his effort while another man might spend much
less time to collect one kg of the same roots and turn them into a basket that could also earn him $20. Skilled
Tembé women can collect one bunch of nuts from a tucumã palm tree (Astrocaryum vulgare Mart.), carve tiny
pieces into the shapes of turtles, frogs, and fish, and string them into necklaces that earn them as much as selling a
60 kg sack of farinha (manioc flour).
The limitation of handicraft sales as source of income for community development is that it mostly
benefits a few people who are creative artisans and assertive entrepreneurs. Many Tembé who churned out
mediocre items were frustrated when the government shop in Belém would not even pay them the bottom price
because it already had a hundred items just like it. The handicraft business was unusually good for a year when an
American intermediary took responsibility for buying top-quality items in Tembé villages and personally shipped
them to a socially-motivated wholesale buyer in San Francisco. This channel fell apart after the American left
Brazil because it proved too difficult to maintain good communication between leaders in the village, the new
intermediary in Belém and the buyer in the U.S.
230
The Tembé and some other Amazonian communities have so far ruled out logging as a means to generate
community income. Where logging will occur in well-planned and executed forest management settings, there
may be ways that NTFP harvests could be integrated into such plans to increase forest product revenue before any
cutting occurs. This study has shown that there are cases when an NTFP can only be profitably harvested from a
tree one time or only once during a long period of regeneration. In the case of copaiba trees that only yield a
significant amount of oleoresin one time, cutting down a tree could be postponed at least until it reaches the size
class of maximum oleoresin production (55 – 65 cm DBH in the trees at Tekohaw) and has its oleoresin removed.
Depending on the circumstances, NTFPs may or may not be able to offer significant economic
opportunities for forest communities. The most important role for these products may not be the amount of
revenue they generate for a community, but the amount of money they save people who would otherwise need to
buy food and other items from stores. People are able to survive in remote parts of the jungle without much cash if
they can procure game, fish, fruit, medicines and materials to make shelter, tools and other basic necessities from
the forest and streams around them. Where forests are destroyed and these NTFP resources are lost, forest peoples
are put into a double bind. They first lose a direct source of food and other materials that they would then need to
buy, and in the process of losing the forest they have also lost many of the resources that allowed them to make
any money at all. The Tembé communities living along the Guamá River provide a striking contrast to the ones
living along the Gurupi. In the Guamá area, the Tembé now have to grow cash crops so they can buy meat from
the city. So much of their forest has been burned by non-Indians that hunting game animals is almost impossible,
and fishing is rarely worthwhile. The Gurupi Tembé are far from rich in monetary standards, but they directly
meet most dietary and other material needs with a diverse variety of fish, wildlife, agricultural and forest products.
They readily acknowledge that their strength as a culture depends on access to all of the plants and animals they
use and share a home with in the forest. It’s not surprising that preserving the forest from destruction is their
major concern.
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TABLE 6.1 PRODUCTION ECOLOGY SUMMARY FOR CASE STUDY NONTIMBER FOREST PRODUCTS AND MANIOC
VARIABLE
Density (Harvested Plants/ha)
Raw Product/plant
Raw Product/ha
Plants Harvested/day
Area (ha) Harvested/day
Raw Product Harvested/day
Comm.Prod./Raw Product
Comm. Prod./ha
Comm. Prod./Day Harvesting
Processing Day/Harvesting Day
Comm. Prod./Day Harv.& Process.
$US/Comm. Product
$US/Day Harv.&Process.
$US/ha (1st Harvest)
Harvests/30 Years
$US/ha (30 Years)
COPAIBA
(Prod. in liters)
0.5
0.1
0.05
5
5
0.25
100%
0.05
0.25
0
0.25
3.50
0.88
0.18
1 - 10
0.18 – 1.75
BREU
(Prod. in kg)
10
0.5
5.0
30
3
15.0
83%
4.15
12.45
0
12.45
0.25
3.11
1.04
6 – 7.5
6.24 – 7.80
TITICA
(Prod. in kg)
50
0.5*
25.0
50
1.5
37.5
20%
5.0
7.50
1
3.75
1.00
3.75
5.00
1-2
5.00 – 10.00
MANIOC
(Prod. in kg)
6032
3.3
17743
X
0.02**
376.2
23%
4216
X
4.1
17.43
0.13
2.17
527.00
1-3
527 – 1581
Values are considered “typical” based on findings of this study and reports in the literature. They could be higher or lower in
different regions.
Notes: * Raw Product/plant for titica refers to weight of harvestable roots per host tree with the vine.
** - Harvest of manioc includes time devoted to forest clearing, burning, planting, weeding and harvesting roots
X – variable not applicable to manioc
232
REFERENCES
Albuquerque, Milton de. 1969. A Mandioca na Amazônia. SUDAM, Belém.
Albuquerque, Milton de and Eloisa Maria Ramos Cardoso. 1980. A Mandioca no Trópico Úmido.
EMBRAPA/CPATU, Belém.
Alencar, Jurandyr da Cruz . 1982. Estudos silviculturais de uma população natural de Copaifera multijuga
Hayne (Leguminosae) na Amazônia Central. 2. Produção de óleo-resina. Acta Amazonica 12(1): 7589.
Baksh, Michael. 1995. Changes in Machigeunga quality of life. pp. 187-205 in Leslie E. Sponsel (ed.)
Indigenous Peoples and the Future of Amazonia: An Ecological Anthropology of an Endangered
World. The University of Arizona Press, Tuscon.
Balée, William. 1994. Footprints in the Forest: Ka’apor Ethnobotany - the Historical Ecology of Plant
Utilization by an Amazonian People. Columbia University Press, New York.
Carvalho, José Edmar Urano de., Francisco José Câmara Figueirêdo, and Carlos Hans Müller. 1996.
Comportamento ortodoxo em sementes de sorva, Couma utilis. Boletim de Pesquisa 168, EMBRAPA,
Belém.
Clay, Jason W. and Clement, C.R. 1993. Selected species and strategies to enhance income generation from
Amazonian forests. FAO Forestry Paper (Final Draft). United Nations Food and Agricultural
Organization, Rome.
Duke, James. 1970. Ethnobotanical observations on the Chocó Indians. Economic Botany 24:344-366.
Durigan, Carlos C. 1998. Biologia e Extrativismo do Cipó-Titica (Heteropsis spp. – Araceae) – Estudo para
Avaliação dos Impactos da Coleta sobre a Vegetação de Terra-Firme no Parque Nacional do Jaú. M.S.
Thesis, Instituto Nacional de Pesquisas da Amazônia (INPA) and Universidade do Amazonas (UA).
Grimes, A., S. Loomis, P. Jahnige, M. Burnham, K. Onthank, R. Alarcón, W. P. Cuenca, C. C. Martinez, D.
Neill, M. Balick, B. Bennett, and R. Mendelsohn. 1994. Valuing the rain forest: the economic value of
nontimber forest products in Ecuador. Ambio 23(7):405-410.
Hoffman, Bruce. 1997. The Biology and Use of Nibbi Heteropsis Flexuosa (Araceae): The Source of an Aerial
Root Fiber Product in Guyana. M.S. Thesis, Florida International University, Miami.
Lancaster, P.A., J.S. Ingram, M.Y. Lim and D.G. Coursey. 1982. Traditional cassava-based foods: survey of
processing techniques. Economic Botany 36(1):12-45.
Nepstad, Daniel C., Adriana G. Moreira, and Ane A. Alencar. 1999. Flames in the Rain Forest: Origins,
Impacts and Alternatives to Amazonian Fire. The Pilot Program to Conserve the Brazilian Rain
Forest, Brasilia.
Oliveira, Ronaldo L. 1998. Extrativismo e meio ambiente: conclusões de um estudo sobre a relação do
seringueiro com o meio ambiente. pp. 93-117 in Alfredo Kingo Oyama Homma (ed.), Amazônia:
Meio Ambiente e Desenvolvimento Agrícola. EMBRAPA-SPI, Brasília.
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Peters, Charles M. 1994. Sustainable harvest of non-timber plant resources in tropical moist forest: an
ecological primer. Biodiversity Support Program c/o World Wildlife Fund, Washington, D.C.
Mors, Walter B. and Carlos T. Rizzini. 1966. Useful Plants of Brazil. Holden-Day Inc., San Francisco.
Robinson, John G. and Kent H. Redford. 1991. eds. Neotropical Wildlife Use and Conservation. The
University of Chicago Press, Chicago.
Stearman, Allyn MacLean. 1995. Neotropical foraging adaptations and the effects of acculturation on
sustainable resource use. pp. 207-224 in Leslie E. Sponsel (ed.) Indigenous Peoples and the Future of
Amazonia: An Ecological Anthropology of an Endangered World. The University of Arizona Press,
Tuscon.
van den Berg, Maria E. 1984. Ver-o-Peso: The ethnobotany of an Amazonian market. Advances in Economic
Botany 1: 140-149.
Warner, P.D.III. and Pontual, A.C. 1994. Manual de Comercialização de Produtos Florestais. GENESYS
(Gênero em Sistemas Econômicos e Socias), The Futures Group, Washington, D.C.
234
APPENDIX 6-A PRODUCTION ECOLOGY STUDIES ON AMAPÁ LATEX AT
TEKOHAW
INTRODUCTION
In 1998, I conducted a pilot harvesting study on 17 trees near Tekohaw that the Tembé classified as
“amapá branca” (white amapá), “amapá vermelha” (red amapá) and “amapá preta” (black amapá). An
examination of leaf specimens from each study tree revealed that all white and red amapá trees were
Parahancornia amapa (family Apocyanaceae) – the species most commonly assigned the amapá common name
(van den Berg, 1984). The tree called black amapá by the Tembé was Couma guianensis Aubl. – another
species in the Apocyanaceae family. Other species in this genus have been well-known producers of latex
used for industrial purposes, chewing gum and medicinal purposes (Mors and Rizzini, 1966; Duke, 1970;
Carvalho et al., 1996). All types of these trees were occasionally harvested by some people from the village to
obtain the latex used as a general tonic, particularly by women feeling weak after childbirth. The traditional
method of harvesting is making a diagonal gash in the tree with two narrowly spaced cuts of a machete. The
latex is then collected in a container that is held or fixed at the bottom of the gash. In order to standardize latex
harvest studies for this tree, I used a short handled tool with a curved sharpened point (“faca de borracha”rubber knife) that is typically used for collecting latex from rubber trees (Hevea brasiliensis).
METHODS
In the pilot study, all 17 amapá trees ≥30 cm DBH that were found during the search for copaiba trees
in 1998 were included in the study. It included eight C. guianensis trees (average DBH 38.1 ± 2.5 cm) and
nine P. amapa trees (average DBH 40.8 ± 2.5). Each tree was first cut with a rubber knife at a 45 degree angle
to a length that was one-third of its circumference at a height approximately 1.2 m above the ground. Each cut
was approximately one cm wide and one cm deep. This depth was sufficient to remove the bark and expose
the latex producing tissues and vessels. The next five cuts were made four cm above the previous cut. The
sixth cut was made on the opposite side of the tree, and then subsequent cuts were made above it. Trees were
235
harvested twice a week for five weeks. Latex was collected in graduate cylinders until latex flow diminished
to a drip rate of ≤1 ml in a five minute interval. The study showed that C. guianensis trees yielded more latex
than P. amapa trees, but results lacked some precision because there was considerable variation in the width
and depth of cuts made by different Indians harvesting the latex.
The amapá latex harvest study was repeated on the same trees in April and May, 1999 with three
modifications. Each cut was 25 cm every time a tree was harvested, and all cuts were made by the same
harvester. The pilot study showed that on average 98% of the latex that emerged from a cut did so in the first
45 minutes; the 1999 study only collected latex from each tree after cutting for this amount of time. This study
was also conducted for five weeks with twice-weekly harvesting.
RESULTS
The results of the 1999 amapá harvest study showed distinct patterns of resin production and yield
between the two species of trees. The C. guianensis trees yielded an average of 100 ± 27 ml per tree during the
first cut. This average steadily declined to 22 ± 6 ml per harvest as new cuts were made every three or four
days above the previous one. The yield rebounded to 75 ± 19 ml per tree when a new series of cuts was begun
to the side of the first ones, but it, too, steadily declined to 21 ± 8 ml per tree during the tenth and final harvest.
The P. amapa trees yielded 19 ± 6 ml during the first cut, but this average dipped and rose to a peak of 43 ± 12
ml per tree during the sixth round of harvesting (week 3 of the study) and then declined to an average yield of
28 ± 6 ml per tree in the tenth and final harvest. The Couma harvest pattern indicates that latex that had
already been produced was simply being drained away in successive rounds of harvesting. The Parahancornia
harvest showed that harvesting may have at least temporarily stimulated latex production.
The relationship between latex harvest and the size of the tree was also different between these two
species. Regression analysis that compared tree diameter (DBH) of C. guianensis trees to the amount of latex
harvested from them fell far short of significance (R2 = 0.2; p = 0.687) while the same comparison for P.
amapa trees did show a slight but significant correlation (R2 = 6.4; p = 0.26).
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APPENDIX 6-B PRODUCTION ECOLOGY STUDIES ON ANDIROBA OIL AT
TEKOHAW
METHODS
I studied andiroba (Carapa guianensis) fruit production and seed oil processing between March and
July of 1999. During this time, Tembé colleagues and I visited 47 trees in the “baixo” (forest subject to
occasional seasonal flooding) and “terra firme” forest around Tekohaw. These trees had a diameter range of
20.9 to 70.5 cm DBH with an average DBH of 38.1 ± 1.6 cm. We visited study trees every other day in March
when fruits were just beginning to mature on some trees. Between April and July when fruits matured and fell,
trees were visited every day. Mature andiroba fruits have a hard outer pod with an average of nine seeds per
fruit that each weighs an average of 16.7 ± 0.1 g (n = 2134 seeds). During these visits we collected all fruits
(including loose seeds and pieces of the outer pod) found underneath the canopy of every tree. Each seed was
weighed and classified as bad (rotten or largely consumed by insect larvae) or good (lack of these “negative”
qualities). An attempt was made to match loose seeds with loose pod pieces so the total number of fruits (and
seeds) that fell from each tree per day could be estimated. Teeth marks on some seeds indicated that many
were partially or fully consumed by wildlife including agoutis, pacas, and brockett deer.
RESULTS
Results of this study showed that these andiroba trees produced an estimated average of 8.0 ± 1.8
fruits per tree, 73.1 ± 16.2 seeds per tree, and 1222 ± 271 g of seeds per tree during the five months of
observations. We collected an average of 807 ± 179 g of seeds per tree (66% of the estimated total weight of
seeds that fell) with an average of 573 ± 127 g of seeds per tree (71% of seeds that were found) that were
considered to be in “good” condition. The greatest number of seeds found was in June and July. In July we
found an average of 616 g of seeds per day from all 47 trees combined (13.1 g of seed per tree). Observations
of the trees indicated that very few trees had any fruits left in the canopy by the end of the harvesting period.
237
Seeds collected in this harvest were processed into oil by one Tembé woman using her traditional
method. Her method was to boil the seeds, store them in the shade until some oil appeared, break open the
seed’s outer cover, scoop out the seed meat, mash the seed meat, and then placed the mashed seed meat on a
tilted metal surface in the sun with a container at the bottom of the apparatus to catch the oil that slowly
emerges. Depending on the size of the seed batch and weather conditions, it took three to six weeks to process
the oil from three to nine kg batches of seeds. In one sample, 2569 g of seeds yielded 178 ml of oil that
weighed 93.3 g. This means that 14.4 kg of seeds would be needed to produce one liter of oil (6.9%
conversion ratio). The conversion from weight of seeds to weight of oil from was 3.6%. The process of
preparing oil took so long that I was unable to get results from five other batches of seeds that were still being
processed when my time in the field ended.
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APPENDIX 6-C MANIOC AND FARINHA PRODUCTION STUDIES AT
TEKOHAW
METHODS
Like many Amazonian people, the Tembé grow manioc (Manihot esculenta Crantz) and convert the
roots to farinha flour that is their most important staple food and occasional cash crop. Since the varieties of
manioc that often grow best in poor soils have lethal levels of cyanide, roots need to be detoxified before the
product is safe to eat (Lancaster et al., 1982). I did a study in Tekohaw to measure the production of manioc
roots, the conversion ratio of processing the roots to farinha, the time involved in the different stages of
handling the plant and final product, and the economics of the process. I wished to compare the value of this
crop on a per hectare basis and revenue per day basis with the NTFPs examined in this study.
I accompanied Tembé through each of the five phases of manioc and farinha production: field
clearing, field burning, manioc planting, manioc weeding, manioc root harvesting, and root processing to
farinha. Fields are created in patches of either secondary or primary forest by cutting down most or all large
trees (usually those ≥ 10 cm DBH) in the middle of the dry season (August or September). Based on
interviews with 25 household heads in Tekohaw, each family cleared an average of 1.03 ± 0.10 ha of forest in
1997. Since groups of men typically take turn clearing each other’s fields, I measured clearing time by
marking out five 25 x 25 meter (0.0625 ha) plots in areas destined for clearing and measured the amount of
time it took five men using axes and machetes to clear each plot. The average amount was divided by five to
estimate the clearing rate per person. I watched field burning, manioc planting, and manioc weeding several
times and made an estimate of the amount of time involved in traveling to the site and conducting these
operations per hectare based on conversations with the families involved and my own observations.
Depending on the variety of manioc, roots are first ready for harvest between six and fifteen months
after planting. On the harvest day, people need to travel to their field, pull up manioc plants, cut off the roots,
carry the roots to a stream to soak (to initiate cyanide leaching), and return to the village. I timed one family
239
harvesting manioc roots from a 20 x 20 meter section (0.04 ha) of their field and approximated travel times
based on my observations and conversations with several families.
I measured manioc productivity by accompanying owners to 13 different fields. All but one of these
fields had been planted 15 months earlier and weeded three to four months after planting. Harvesting was done
by pulling up plants in a contiguous series of 2 x 2 meter plots in a harvest area designated by the field owner.
The number of manioc stems and weight of manioc roots was recorded for each plot. The study team
harvested 12 to 40 plots per field per harvest. The number of plots sampled per field in one day depended on
how much manioc the field owner needed to produce a two to three week supply of farinha for his family. The
average area harvested per field was 119.4 ± 13.0 m2 (0.01 ha).
Farinha production day begins three to five days after manioc roots have been soaked in a stream
when the owner determines the roots have reached the proper degree of softness. He then gathers fire wood
and brings it to the shelter that stores the “forno” – the ceramic oven used to cook the manioc in a 1.5 to 2
meter wide round steel pan. The soft manioc roots are taken out of the water and have their outer shells
removed. Root interiors are then grated and mashed and have hard pieces removed. The mass is then placed
in a “tipiti” (a five-foot long flexible tube woven from local reeds) or other type of press to squeeze out as
much free water as possible. The strained mass is then placed in the pan of the hot forno to drive off any
remaining cyanide and produce the dry crunchy farinha flour. I quantified the conversion ratio and time
required for a family group to process manioc roots to farinha flour on five occasions.
RESULTS
Results of yield studies showed that the Tembé fields produced an average of 17743 ± 1892 kg
manioc roots/ha (n = 13 fields). This average places it in the “regular” yield range (15 to 20 metric tons/ha) for
manioc harvests in the Amazon where root production between 20 and 25 tons per hectare is considered
“good” and yield less than 15 tons per ha is rated “low” (Albuquerque, 1969). It took an average of 4.21 ±
0.26 kg of manioc roots to make 1 kg of farinha (n=5) so the Tembé conversion rate of 23.7% for this process
was lower than the 32 to 35% range cited in other Amazon manioc studies (Albuquerque, 1969 and
Albuquerque and Cardoso, 1980). Using the Tembé conversion rate, the fields around Tekohaw produced an
240
average of 4216 ± 450 kg of farinha per ha. Farinha is typically sold in 60 kg sacks that sold for about $R 15
per sack ($R 10 to $R 20 range) between 1997 and 1999 (Galvão, personal communication 1999). Tembé
fields were, therefore, producing an average of 70.3 sacks of farinha/ha worth an estimated $R 1054/ha.
The time required to clear a forest with hand tools averaged 0.033 ± .006 ha per hour per person with
the assumption that two hours were spent in transit to the field. Using the farinha per ha average value
mentioned above, the clearing rate averaged 0.47 ± 0.09 hours per person per sack of farinha produced. This
rate was 45% longer in the two plots that contained undisturbed primary forest than in the three plots of 20year-old secondary forest because one large tree took much longer to bring down than many smaller pioneer
trees. Since 1999, the village has had greater access to chainsaws so the clearing rate will be much faster.
Field plots were burned two to three months after they were cleared. Manioc stems were planted
several weeks after clearance. When new manioc plants were three to four months old, the fields were weeded.
Each of these operations required the owner and several friends or family members to spend about one day to
go to and from the site and carry out the required task. I estimate that each of these operations added about the
same amount of time needed to make the final product (0.5 hour per person per sack of farinha) as did field
clearing activities. Results of manioc harvesting showed that it required about 3.2 hours per person per sack of
farinha produced. Processing manioc roots into farinha was by far the most time consuming phase of the entire
process since it took an average of 19.4 ± 4.6 hours per person per sack of farinha produced.
Combining the five elements of farinha production showed that a total of 24.1 hours of time per
person was needed to produce one 60 kg sack of the product. This means a Tembé farmer could make $R
2.87, $R 4.34, or $R 5.81 per seven hour day as the selling price for a sack of farinha increased from $R 10 to
$R 15 to $R 20. While the Tembé have not quantified the value of their time in these terms, they clearly
carefully assess their opportunity costs in deciding whether or not to sell farinha (and NTFPs as well) since
most men were not willing to sell it unless the price was at least $R 15 per sack. One man who made a highquality farinha wouldn’t sell a sack for less than $R 18. Tembé farinha sellers reaped the full sales price of
their product because the government Indian agency (FUNAI) usually took responsibility for arranging its sale
and transportation. FUNAI was also trying to make farinha production more efficient by providing electric
manioc grinders and new farinha roasting ovens.
EPILOGUE
242
CHALLENGES AND LESSONS STUDYING NON-TIMBER FOREST PRODUCT
ECOLOGY IN AN INDIGENOUS COMMUNITY
Learning how to do publishable quality scientific work is not easy under any circumstances; learning
how to do it in a different culture, a different environment and speaking a different language makes the
learning curve even steeper. Scientific journals, though, tend to present the successful parts of researchers’
work, and relatively few researchers write about the non-scientific challenges or failures they faced. As I
stated in the introduction to this dissertation, my experiences in the field taught me many things that I cannot
condense into a case study table, so I will present some of these thoughts here since I believe they, too, might
be valuable to other researchers.
OFFICIAL CHALLENGES
The official challenges of doing NTFP research in Brazil and many countries begin long before one
sets foot in the forest. Getting a research visa is the first major hurdle. This first requires finding a “scientific
counterpart” who will vouch for the integrity of your work and financial solvency. I was fortunate to find a
professor at the agricultural university (FCAP) in Belém to take on this role of handling paperwork with the
federal government. Getting the visa next required the approval of my research proposal by the national
research council (CNPq), natural resources agency (IBAMA), the Indian agency (FUNAI), three independent
scientists, and Minister of Technology. This took “only” nine months because I spent hundreds of dollars in
phone calls to Brasília pleading with various officials to move my application on to the next person. Patience
and persistence are required to succeed.
HEALTH CHALLENGES
I got every immunization required or recommended ahead of time and brought a small pharmacy’s
worth of prescription and over-the-counter drugs with me into the field. “Where There is No Doctor” (Werner
et al., 1992) was my family’s most well-read book during our time in the village. I drank water that had been
243
passed through a ceramic filter, applied andiroba oil to my skin to ward off biting bugs during the day and slept
in a hammock surrounded by mosquito netting at night. In spite of these precautions, there were other irritants
and more serious maladies we had to regularly confront.
After getting badly infected bug bites during my first trip, my family adopted an evening first aid
ritual, we took turns treating bites and scrapes with alcohol, Mercurochrome, and anti-biotic ointment. Tembé
neighbors showed us how to pluck burrowing fleas from tender toe areas with an orange tree spine before they
festered. One day my wife and daughter removed several hundred tiny ticks from my body after I had
stumbled into a nest of them in the forest. Other nasty encounters with invertebrates included wasp stings, ant
bites, and accidentally squishing a venomous hairy caterpillar on my arm. Tall rubber boots saved my field
crew and me several times from poisonous snake bites, and I was lucky not to get malaria when many others in
the village did. My body was not always strong enough or well adapted to the environment, though, so I
periodically got sick with a fever or malaise of unknown origin that caused me to miss days in the field. I left
the village for a whole month once when bad headaches and pain in my joints became an unrelenting part of
my day. I did not completely recuperate until I came home for an extended period of time. I loved this field
work as much as anything I’ve ever done, but taking care of my health in this climate required a lot of
vigilance.
LOGISTIC, COMMUNICATION AND TECHNICAL CHALLENGES
Doing research in primary tropical forests almost by definition requires working in remote places
because most places that are easy to get to have been significantly altered. Traveling from Belém to my field
site at Tekohaw was almost always a saga. Scheduling a trip to the reserve took anywhere from two days to
two weeks. Once a journey began, it first involved a four hour bus ride from Belém to the town of Gurupi at
the intersection of the main Belém highway with the Gurupi River. One often had to spend one or two nights
in a rickety store room until a government or Tembé boat arrived and was provisioned for the trip up river.
Traveling in an open motorboat involved a 6 to 10 hour trip to the village if the motor did not break and
gasoline did not run out on the way. Going by covered motor launch saved one from exposure to rain, but the
trade-off was spending 12 to 24 hours in transit with the constant noise and fumes of a diesel engine. Travel
244
during the dry season was particularly tricky since boats had to navigate through rocky rapids, and several
boats capsized in the process. Tekohaw now has an airstrip that can handle small planes, but chartering a
private plane for this one hour trip costs about $800, so I took whatever boat was available and did my best to
protect equipment for whatever might happen on the way.
Once I made it into the village, it was not easy to communicate with anyone outside it. The life line
of Tekohaw to other villages and FUNAI is a ham radio. There were daily check-ins to share important news,
pass along requests for transportation, and arrange meetings. Once a month or so I could send out a letter with
someone going to the city or receive mail brought in by the FUNAI agent. I looked into a cell phone with
international capabilities but the licensing procedure and costs were prohibitive. In my second year I
periodically listened to BBC International, but in general once I was in the reserve I pretty much had to accept
that I was not going to have any contact with the rest of the world for six to eight weeks. This was a relaxing
blessing in many ways and tough in others.
My brand of ecological field work fortunately did not rely on a lot of sophisticated equipment. I
brought standard 50 meter measuring tapes, tree diameter tapes, compasses, tree tags and GPS units from the
U.S., and I easily got basic tools like machetes, hammers, drills in Belém hardware stores. Accidental machete
cuts through measuring tapes were remedied with duct tape. Setting up my house as a lab and office that could
function at night was a bigger challenge. Three unbreakable solar panels worked flawlessly for years, but I
went through three expensive car batteries before I finally worked out (with the help of a digital voltmeter)
how to budget my energy consumption for a couple of compact fluorescent light bulbs, a laptop computer and
portable printer. I did without the computer for the first few months since its screen was damaged in the first
boat trip to the reserve, but once it was repaired, it became a vital tool for creating data collection sheets,
entering data, and writing reports in the field.
BLENDING INDIGENOUS KNOWLEDGE WITH SCIENTIFIC INQUIRY
I intentionally wanted to study NTFPs with indigenous people because I believed that I could learn a
lot more about these plants working with people who were familiar with them than if I simply set off with a
tape measure into a pristine area of forest by myself. It took less than a few days to confirm that my Tembé
245
colleagues knew more about the forest they had grown up in than I could learn in my lifetime, but I was eager
to learn as much as I could from them. It took a good bit longer to realize that their understanding and beliefs
did not always translate well into my desire to assign numerical values to some things or describe natural
history relationships in traditional biological terms. One lesson was learning how to gauge the response to a
question about the abundance of a particular plant or how much product it typically yielded. Beginning with
the copaiba work, the Tembé elders I first worked with assured me that we would find lots of trees and told me
about times when they had filled large cans with oleoresin. It was rather sobering to them and me when we
found that it took several weeks of daily searching just to find 50 trees and that we could only gather a few
liters of oleoresin after drilling most of them.
Another important lesson I learned working in the villages was that the level of skills and knowledge
varied dramatically between individuals. Some people were very adept at finding certain kinds of trees, but not
others. Elders could generally identify hundreds of types of trees but were unable to read, write or count
higher than ten. Younger men generally knew relatively few trees, but a few were quick to learn how to read
instruments and reliably record the measurements on data sheets. Research teams for any given NTFP almost
always had people with a blend of these skills, so the research could provide opportunities for crossgenerational learning and appreciation.
When I started asking about breu, Tembé and Ka’apor collectors told me about the “tapuru” (larvae)
they found in the lumps and related stories about filling entire sacks of resin from single trees. As the research
progressed, it became apparent that the Indians’ observation of the “tapuru” was fundamental to the resin’s
formation, and we proceeded to probe this relationship together in great detail. On the harvest side, it again
became apparent that the huge yields people described were the result of their best days, not their normal ones.
This reinforced the recommendation by Grimes et al. (1994) that estimates of NTFP harvest amounts obtained
through ethnobotanical inquiry be followed up with actual production studies in order to develop reliable
quantitative estimates.
The most culturally challenging situation I confronted related to work in the field involved the
regeneration of titica vine roots. As mentioned in the case study, one Tembé elder told me that the slender
titica vines climbing some host trees were a different kind of vine than the ones whose aerial roots were
246
harvested. Other Tembé told me that these aerial roots were created when the legs of certain dead ants become
elongated. It was a believable story to some since they had not seen flowers or fruits of the part of the plant
that is largely out of site in the host tree’s upper trunk. I was skeptical about the Indian version of titica
regeneration, but I tried to keep an open mind. Before I found out that titica was a secondary hemiepiphyte
that actually starts its life on the ground, I was wondering if ants might play some role in the germination of
titica seeds in the canopy. When I later harvested several whole plants at a different site, it was immediately
apparent that the roots were attached to the branches off the main stem of the titica plant and that normal
reproductive structures were held on the branches’ tips. I regret that I did not harvest some plants with the
Tembé since the plant could have provided us with a concrete example to discuss what I felt awkward saying
based on book knowledge. My hope is that harvesters who have a better understanding of the plant’s
regeneration mechanisms will be better able to regulate the harvest to preserve the resource for the future.
EXPECTATIONS OF A RESEARCHER
Working with the Tembé provided me an opportunity to learn about the forest in ways I never could
have done on my own and gave me some of the best and worst experiences of my life. Many of these
successes and disappointments relate to the fact that my relationship with the community went far beyond my
role as a researcher studying NTFPs. As mentioned in Chapter 1, my initial contact with Tembé from the
Gurupi River villages occurred in the U.S. when I escorted a leader on a 10-day long tour of the U.S. that gave
him many opportunities to discuss his people’s needs in the areas of health care, education and land rights. I
was welcomed to Tekohaw not just as someone who would study the economic potential of a few plants but as
someone who would be an active participant in many aspects of village life and concerns. For the better part of
four years, I was known to everyone in the community as “Zui’zu” (white frog). I was determined not to be
another scientist who came into the community, gathered whatever data and materials I wanted and left without
giving much in return.
I tried to help direct resources to the Tembé community in a variety of ways. The most supportive
people were from the State College Friends Meeting that formed the Tembé Indian Support Committee. This
committee secured several large donations from one individual and did fundraising projects that included:
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young children making and selling buttons they made with Indian designs, older children making dinner and
offering babysitting services, and all children selling flower bulbs. I bought and traded for Tembé handicrafts
in the village and sold them at the Friends Meeting and at slide shows I did about the Tembé in the U.S. I
arranged for a Tembé leader from Canindé village to attend an Amazonian indigenous conference in New York
and recruited five people from the U.S. to attend a traditional festival in the reserve in return for a donation to
the community. I obtained donations of clothing, hand tools, medical and school supplies, soccer balls and
sports equipment for six Indian villages along the Gurupi River. Funds raised in these projects were used to
finish construction of a large wooden launch, buy a new motorboat and diesel engine, repair two gasoline
engines, produce a book of 200 traditional Tembé chants in Tembé and Portuguese, and possibly finance a
major land rights project.
While living in the village I tried to contribute to the community by bringing in ample quantities of
fish hooks, fishing line and ammunition for trade, loaned out my tools, fixed broken water pipes, tended sick
children, took family photos, and gave evening classes in math, geography, science, and environmental
advocacy. When I left the village I donated my solar panels, furniture and remaining first aid and paper
supplies to the schools and infirmaries of three different villages.
Trying to get my research done and be a good employer presented many challenges. The first
summer in Tekohaw the elder Tembé worked with me without formal wages, but at the end of the summer they
each made a list of items they wanted from the city in compensation for their time. I initially liked this system
because it felt more like a cooperative venture than a business arrangement, but it was not fair because a few
people who worked little wanted more than others whose daily efforts were integral to the project. When I
returned in 1997 I paid everybody who worked with me the same amount for each day they worked and raised
this amount every year. This system in general worked well, but there was social pressure to hire as many
people as possible. This was difficult because my funds were limited, and I mostly needed people who either
knew forest trees very well, could read and write numbers well or both. At one point I created a special project
to measure manioc productivity more as a way of paying some extra people than because the information was
vital to my research goals.
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I strove for an elusive balance between firmness and flexibility in dealing with research needs versus
other needs of the village and my helpers. I got very little work done in the fall of 1997 because most men
were away from the village cutting boundary lines for the reserve. Shortly after they came back they all got
engaged in the 10-day long St. Benedict’s Festival. I had to plead with the community to release a few people
to me or I would not be able to accomplish anything. A year and a half later the situation was reversed. I
sometimes had four crews of people going out on the same day to get production data on andiroba, breu, amapá
and copaiba. It seemed wonderful until the chief came to me and asked me to release some of these men so
they could work on the community garden and building projects. It became apparent that while many people
appreciated the opportunity to work, the community was suffering because their desire to do work for the
community for free had begun to wane. Some of these people at least willingly made donations to a travel
fund for the leader when there was an important meeting for him to attend in Belém or Brasília.
There is no doubt that taking on such a broad range of research topics had other consequences for my
work and relationships. By trying to gather production data about so many plants during a two-year period, I
inevitably sacrificed gaining a deeper understanding of a few of them. Over time, I became more of a field
manager than an ecologist slowly building my insights through day by day observations in the field. Part of
this seemed unavoidable since there were many days when I felt too weak or sick to go in the forest, but this
mode gradually altered my relationship with my Tembé field assistants. I knew next to nothing about the
forest, but when I faced the mud, bugs, poisonous snakes, rain and shared one can of sardines with them for
lunch I earned their respect. When I stayed in the village more often during the day, I had fewer chances to
personally share their successes and travails in the forest. My interactions with them became more centered
around checking data sheets at the end of the day and paying them minimum wage every few weeks. Our
relationship had uncomfortably shifted from friends and colleagues to employer and employees.
At the end of the project it was clear that the expectations the community had of me (and that I had for
myself) were more than I could possibly meet. By hiring people from one family in Tekohaw, other families
sometimes felt slighted if I did not hire someone from theirs. Since I lived in Tekohaw it was natural that I
hired mostly people from there and provided other types of assistance to that community. This made some
people in the other Tembé villages upset that I was not paying more attention to them. Since my field work
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was carried out exclusively in Tembé villages and fundraising projects brought benefits mostly to them, the
Ka’apor Indian villages on the other side of the Gurupi River began to feel increasingly angry that I was
unjustly ignoring them. I was aware that such feelings could arise and tried to deal with it by hiring
representatives from several villages to work in projects at Tekohaw, but it was difficult for people with
families to stay away from their home villages for long. I did one well-received side project in Cajueira
village, but it was not possible to do so in others without fracturing ongoing research at Tekohaw. Boats and
boat equipment were theoretically used by all of the villages, but over time, each village wanted their own
craft. I tried to channel funds earmarked for a land right’s project to a committee of village leaders, but the
delay in releasing funds due to inter-village politics led to recriminations against me by individual villages that
wanted me to release the funds directly to them.
I did one round of visits to all of the Gurupi villages to explain the goals of the research project and
did regular updates at Tekohaw. It was evident, however, that even at Tekohaw the level of understanding of
all of these studies was not very high. The simplest explanation was that I was studying plants that might lead
to additional sources of income, and I used the most creative ways I could think of to convey quantitative
concepts to people who knew little or no math. When the research began to show that the new products being
investigated were not very promising in this respect, it became harder to explain to most people why
continuing such studies was still important or relevant to them. Information on some study trees was lost
because people removed numbered tags or harvested trees for themselves in between research harvests. Some
people who worked with me took a genuine interest in probing topics like the relationship between the breu
resin and the weevil larvae that lived in it, but many young men worked with me mostly because it was a
relatively easy and interesting way to make some money. My family was warmly embraced with no
reservations, but I believe most of the village accepted me as a friendly foreigner with quirky interests and was
content with my presence as long as I provided some tangible benefits to the community on a regular basis.
The increasing interest that people had in making money seemed to fuel speculation that I hoped to get rich
taking photos, recording chants and collecting plants around the village. These issues were periodically
discussed, but it became increasingly difficult to work without unspoken suspicions and jealousies clouding
many interactions. By the time I returned to Brazil in the spring of 2000 to wrap-up field work for my
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dissertation and to present some preliminary research results, the cord of trust between several villages and me
had badly frayed. I left with my head high but a very heavy heart.
Indigenous people have an understandable expectation that researchers they welcome to their
communities will carry out their work with full consultation, conduct themselves with integrity and fairly
compensate people who work with them and the villages that host them. These were my expectations going
into this project, but it is apparent I needed to communicate more openly and realistically about these subjects
more often. My dialogue with leaders sometimes worked and sometimes did not because I had to operate in
the context of intra- and inter-village politics that were often beyond my understanding or control. I could
speak Portuguese well enough to do my work in the forest and conduct daily life in the village, but at times I
was simply unable to communicate effectively about complex and sometimes volatile emotional and cultural
issues because I couldn’t use my native language.
I admit that for a time I enjoyed the feeling that I could do fascinating work in the forest and make
contributions to the community in so many ways. The needs of the people seemed deep and diverse, and they
stimulated creative energies to help however I could. Trying to be a researcher, medic, teacher, employer,
banker, fundraiser, archivist, handicraft buyer, hardware supplier, father and husband were too many roles to
keep up for the long haul. My inability to balance these roles led me to make mistakes that hurt my
relationship with some members of the community. The cumulative weight of these roles almost stretched the
patience of my family and my health to their breaking points.
The issue that I will ponder long and hard as a result of my time with the Tembé is the role of money
in the community. I was initially attracted to studying the economic potential of NTFPs because I thought that
increasing revenue would help community improve health care, education and support forest protection. Over
time these beliefs steadily eroded. Most people wanted “modern” medicines because they had lost some of the
knowledge or faith to use rainforest plants, but they didn’t want to spend their money on these things because
they expected the government to supply them. The Tembé wanted to have Tembé teachers in their schools, but
they consistently failed to honor commitments to compensate the local people who devoted long hours to
teaching the village children. Young leaders were unanimously praised for their efforts to defend their land
rights, but raising donations from the community to send these people to important meetings was a perennial
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struggle. People had not abandoned their desire to help the welfare of their community, but they clearly chose
to spend most or all of what little money they made on material goods for their family. The appeal of cooking
food with a clean flame from a gas stove instead of cutting fire wood, making charcoal and living in a sootfilled house is obvious. People needed cash to purchase certain foods and materials like salt and fish hooks to
survive. Given my affluence as a middle-class American, it would be hypocritical to criticize anyone for
wanting a new shirt or CD player, but I left with no illusions that increasing village income would necessarily
improve the Tembé quality of life, preserve its traditional culture or save its forest. Money can sometimes play
a vital role in supporting such objectives, but mechanisms need to be in place in the community to handle such
funds in a positive way. My feeling was that the struggle to accomplish these goals was being waged from
people’s hearts and didn’t depend on how much money they had or the size of their radio.
I hope I will have the chance to work with another community in the Amazon. There is an important
role that researchers can play to promote forest conservation and support rural community development. They
can continue to study the complexities of nature and help develop ways for people to coexist with the full
diversity of a tropical ecosystem in a world where social and economic conditions are rapidly changing. I
would ideally like to take on this kind of challenge again as part of an integrated group of researchers and
development practitioners. I do not wish to fall back into a narrowly defined role of a “specialist,” but I realize
that I can’t do all the things that I’d like to do working alone. I would seek to do my part in an interdisciplinary team well without apology for what I can’t do, and maintain as much humility as possible.
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REFERENCES
Grimes, A., S. Loomis, P. Jahnige, M. Burnham, K. Onthank, R. Alarcón, W. P. Cuenca, C. C. Martinez, D.
Neill, M. Balick, B. Bennett, and R. Mendelsohn. 1994. Valuing the rain forest: the economic value of
nontimber forest products in Ecuador. Ambio 23(7):405-410.
Werner, David, Carol Thuman, and Jane Maxwell. 1992. Where There is No Doctor: A Village Health Care
Handbook. Hesperian Foundation, Berkeley.
VITA
JAMES CAMPBELL PLOWDEN
EDUCATION:
1975: University of Pennsylvania, BA Biology
1993 - 1994: University of Maryland, one year grad. Non-degree; one year in Environmental Science Master's program
2001: Penn State University, PhD Ecology
WORK HISTORY:
2000 (8/00-8/02): Penn State University, University Park, PA; Intercollege Graduate Degree Program in Ecology
Program; Acting Head (7/01-8/02); Research Assistant (1/01-6/01); Biology Dept. Instructor/Graduate Teaching
Assistant (8/00-12/00; 8/01-12/01).
1995 (3/95 - 7/95): Environmental Investigation Agency, Washington, DC; Wildlife Trade Investigator
1994 (1/94 - 1/95): Univ. of Maryland, College Park, MD; Natural Resources Management Program, Graduate Teaching
Assistant and Program Assistant
1993 (1/88 - 4/93): Greenpeace International, Washington, DC; Tropical Forests Campaign, International
Coordinator/Special Projects Manager; International Whale Campaign Coordinator
1985 (1/85 - 12/87): The Humane Society of the US, Washington, DC; Whale Campaign Coordinator
1982 (1/82 - 12/84): Greenpeace USA, Seattle, WA; National Wildlife Coordinator
1979 (5/79 - 5/80; 9/80 - 12/81): Greenpeace International, Tokyo, Japan; Japan Project Coordinator
RECENT PUBLICATIONS
Plowden, C., C. Uhl, and F.A.de Oliveira (in press) Breu resin harvest by Tembé Indians and its dependence on a barkboring beetle. in J.R. Stepp, F.S. Wyndham, and R.K. Zarger (eds.) 2002. Ethnobiology and Biocultural Diversity.
University of Georgia Press.
Plowden, C. (in press) Politics and progress: challenges to sustainable harvest of forest products by Tembé Indians
in the Brazilian Amazon - Response to editorial by Cunningham and Shanley. People and Plants Handbook
special issue Managing Resources: Community-based Conservation.
Plowden, C. (in press). Profiles on Carapa guianensis, Parahancornia amapa, Protium spp., and Copaifera spp. in P.
Shanley, (ed.) Tapping the Green Markets: Management and Certification of Non-timber Forest Products.
Earthscan Publications, London.
Plowden, C. and D. Bowles. 1997. The illegal market in tiger parts in northern Sumatra, Indonesia. Oryx 31(1): 59-66.
AWARDS AND GRANTS:
2001: Penn State Univ. J. Brian Horton Award in Ecology
2000: Penn State Univ. Biology Department Honorable Mention for Best Teaching Assistant Award
2000: Penn State Univ. Ecology Program Professional Meeting Travel Award
1997: Rainforest Alliance Kleinhans Fellowship (2 years)
1997: Food, Conservation and Health Foundation Grant
1995: National Science Foundation Graduate Fellowship (3 years)
1993: The Hotchkiss School Alumni Association Community Service Award
VOLUNTEER ACTIVITIES:
1999 - 2001: Ecology Graduate Student Organization (of Penn State) - Co-founder and President
1997 - 2001: State College Friends Meeting, State College, PA - Co-clerk Tembé Indian Support Committee
1985 - 1993: Tropical Ecosystem Research and Rescue Alliance International, Washington,. D.C. - Founder/President