Biodiversity Assessment and Long-Term Monitoring, Lower Urubamba Region, Perú 1997

Biodiversity
Assessment
and
Long-term
Monitoring,
Lower
Urubamba
Region, Per
San Martin-3
and
Cashiriari-2
Well Sites
SI/MAB
Series #1
Edited by
Francisco Dallmeier
and Alfonso Alonso
Smithsonian Institution
Monitoring & Assessment
of Biodiversity Program
(SI/MAB)
Acknowledgments
We first express our sincere thanks
to Mr. Alan Hunt, general manager,
and Murray Jones, manager of
Health, Safety and Environment, of
Shell Prospecting and Development
(Per) B.V. (SPDP) for their support
of the biodiversity assessment and
monitoring project in the lower
Urubamba region of Per. Gert
VanderHorst, Peter Dushe, Craig
Schenk, Allan Sayers, Miguel RuizLarrea, Mary Malca, Augusto
Baldoceda, Lincoln Williamson, and
Alonso Zarzar of SPDP devised
solutions to all logistical, operational,
medical, and safety challenges.
Annelise and Machy were indispensable when Gert and Murray had to be
located. Alejandro Alvarez, Martha
Rojas, and Victor Grande facilitated
work with Native communities and
found excellent field assistants. Alan
Dabbs of Pro-Natura USA was
helpful concerning sustainable use of
natural resources. SPDP provided
transportation to the well sites, and
we traveled safely in the helicopters
of Canadian Helicopters, Ltd. and
Fuerza Aerea Peruana. We also thank
site supervisors Aarnoud Smit and
Robert Smit at Nuevo Mundo for
logistical and supply arrangements.
Field work was greatly enhanced by
field guides Federico Ramrez, Antonio Gmez, Jacobo Pacaya, Justino
Martin, Jos Tenteyo, and Ismael
Pascal. We thank the Native communities for collaborating with us and
SPDP on this project. Employees of
the Peruvian construction company
COSAPI, led by Ing. Roberto Arbulu
at San Martin-3, supported our field
operations. Concession and Catering
(Per) provided extremely highquality food. The Asociacion Peruana
para la Conservacion de la Naturaleza
(APECO) handled administrative
matters and some logistics from Lima.
The Instituto Nacional de Recursos

Naturales (INRENA) granted the
permits for sampling scientific specimens. The Museo de Historia Natural,
Universidad Nacional Mayor de San
Marcos, directed by Dr. Neils
Valencia, played an important role in
developing this project, and several
Peruvian counterpart researchers
come from that institution. It is also a
depository for specimens we sampled.
We are very appreciative of the work
of Judy Sansburry, Shana Udvardy,
and Tatiana Pacheco of the
Smithsonian Institutions Biodiversity
Programs. Alfredo Sandoval provided
logistical support in Lima. Deanne
Kloepfer reviewed and provided
comments on the papers.
SI/MAB Biodiversity Program
Institute for Conservation Biology
SI/MAB Series #1
1997 by SI/MAB Biodiversity Program
All rights reserved
ISBN # 1-893912-00-0
Library of Congress Catalog
Card Number: 99-61103
Cover design: Mitchelle Stephenson
Cover photographs: Severo Balden with
Discorea sp.; San Martin-3 well site (F.
Dallmeier.).
Opinions expressed in the SI/MAB Series
are those of the authors and do not
necessarily reflect those of the Smithsonian or partner organizations.
Suggested citation: F. Dallmeier and A.
Alonso (eds). 1997. Biodiversity Assessment of the Lower Urubamba Region,
Per: San Martin-3 and Cashiriari-2 Well
Sites. SI/MAB Series #1. Smithsonian
Institution/MAB Biodiversity Program,
Washington, DC.
Printed in United States of America by
Charter Printing, Alexandria, Virginia,
on recycled paper. This publication was
made possible through a grant from Shell
Prospecting and Development (Per) B.V.
Contents
Executive Summary........................i
Francisco Dallmeier and
Alfonso Alonso
Resumen Ejecutivo..................xxxv
Alfonso Alonso
Introduction.....................................1
The Lower
Urubamba Region........................17
Photo Gallery.................................21
Biodiversity
Assessment and Monitoring
for Adaptive Management...........29
Francisco Dallmeier
Assessment of Biological
Diversity and Long-term
Monitoring Plan for the
Lower Urubamba Region............41
Thomas Stohlgren and Geneva Chong
ASSESSMENT
AND MONITORING:
VEGETATION
Flora I............................................45
Pedro Acevedo, Deborah Bell,
Katherine Rankin, and Stephen Smith
Flora II...........................................59
Percy Nez, Severo Balden, and
Hamilton Beltrn
Floristic Composition,
Structure and Diversity
Assessment in the Lower
Urubamba Region.......................81
Alfonso Alonso, Francisco Dallmeier,
Shahroukh Mistry, Percy Nez, Jos
Santisteban, Gorky Valencia, Severo
Balden, Hamilton Beltrn,
Christopher Ros, and James Comiskey
Long-term Vegetation
Monitoring Plan............................91
Francisco Dallmeier, Shahroukh
Mistry, and James Comiskey
ASSESSMENT
AND MONITORING:
ARTHROPODS
Arthropods.....................................101
Jos Santisteban, Gorky Valencia,
and Alfonso Alonso
Diurnal Butterflies
(Lepidoptera: Rhopalocera).......115
Alfonso Alonso and Gorky Valencia
Nocturnal Butterflies
(Lepidoptera: Heterocera:
Ctenuchinae)...............................127
Juan Grados
Ants (Hymenoptera)...............131
Leeanne Alonso and
Alfonso Alonso
Bees and Wasps
(Hymenoptera: Aculeata).......141
Manuel Laime
Dragonflies and Damselflies
(Odonata: Anisoptera and
Zygoptera)..................................149
Jerry Louton
Spiders
(Arthropoda: Arachnida)...........155
Saida Crdova and Janine Duarez
Snails
(Mollusca: Gasteropoda)............161
Rina Ramrez and Saida Crdova
Beetles
(Coleoptera: Scarabaeidae).......169
Gorky Valencia and Alfonso Alonso
Long-term Monitoring
of Arthropod Fauna in the
Lower Urubamba Region..........177
Albert Finnamore
ASSESSMENT
AND MONITORING:
AMPHIBIANS AND REPTILES
Amphibians and Reptiles I........213
Robert Reynolds, Thomas Fritts,
Steve Gotte, Javier Icochea, and
Guillermo Tello
Amphibians and Reptiles II......223
Javier Icochea and Joseph Mitchell
Genetic Structure of
Amphibians and Reptiles...........231
Jesus Crdova and Cesar Aguilar
of Amphibians in the
Joseph Mitchell
ASSESSMENT
AND MONITORING: BIRDS
Birds...............................................247
George Angehr and
Constantino Aucca
of Bird Fauna in the
Jim Siegel and George Angehr
ASSESSMENT
AND MONITORING:
MAMMALS
Small, Non-volant Mammals.....281
Sergio Solari and Juan Jos Rodrguez
of Small, Non-volant
Mammals in the
Sergio Solari
Bats................................................293
Don Wilson, Robert Baker, Sergio
Solari, and Juan Jos Rodrguez
of Bats in the Lower
Urubamba Region.....................303
Don Wilson
Medium
and Large Mammals..................311
Major Boddicker
of Medium and Large
Mammals in the Lower
Urubamba Region......................341
Major Boddicker
Ecto- and
Endoparasites in Mammals......345
Ricardo Guerrero
Ectoparasites in Mammals........351
John Chavez
San Martin-3
and Cashiriari-2
Well Sites
Alfonso Alonso
Monitoring and Assessment
of Biodiversity Program (SI/MAB)
Synopsis
The biologically rich Amazonian lowlands of Per represent an
exciting opportunity to integrate
science, conservation, and development through careful planning,
assessment, monitoring, and decision-making. In the Lower
Urubamba region, the Smithsonian
Institutions Institute of Conservation Biology has joined Shell Prospecting and Development (Per)
B.V. (SPDP) in a unique venture
aimed at achieving environmentally
sensitive development of natural
Biodiversity Assessment & Monitoring, SI/MAB Series #1 (1997)
Executive Summary
Biodiversity
Assessment and
Long-term
Monitoring of
the Lower
Urubamba
Region, Per
gas and condensate resources.

Collaborators include Peruvian
scientists from the Museo de
Historia Natural of the Universidad
Nacional Mayor de San Marcos,
Universidad San Antonio Abad del
Cusco, and Universidad Nacional
de la Libertad (Trujillo).
SPDP is currently appraising
gas reservoirs in the study area (at
the San Martin-3 and Cashiriari-2
well sites) that were discovered
during exploration in the mid1980s. This work consists of
drilling four wells and then completing a logistics base. Plans for
additional development of the gas
reservoirs call for flow lines, a gas
plant located in the study area, and
a pipeline across the Andes to the
coast where coastal facilities will
also be built.
The overall goals of
Smithsonians work are to link
vegetation information to data
about other taxa that are being
assessed and monitored and apply
the results to management decisions at the well sites and, later, in
the larger area. Multi-disciplinary
teams of researchers are carrying
out the project under the research
and training framework devised by
SI/MAB (Smithsonian Institution/
Monitoring and Assessment of
Biodiversity Program) for longterm, multi-taxa forest biodiversity
monitoring at permanent research
sites.
Two phases of the project have
been completed. First, a relatively
quick assessment of the biological
and cultural diversity in the Lower
Urubamba region was completed.
This was followed by a workshop
about the area in September of
1996 to set the parameters for a
biodiversity assessment and monitoring program. Participants in-
Executive
Summary
We conducted
a baseline
biodiversity
assessment of
target groups
at the
San Martin-3
and
Cashiriari-2
well sites.
ii
cluded representatives of government institutions, non-governmental organizations, the scientific

community, and local Native
groups. A report on Phase I was
published in early 1997.
Phase II included initial assessments of biodiversity in the study
region and establishment of multiscale plots and permanent
biodiversity monitoring plots.
SI/MAB Series #1Biodiversity
Assessment and Long-term Monitoring of the Lower Urubamba
Region, Per: San Martin-3 and
Cashiriari-2 Well Sitesprovides
the basis for this executive summary. The study addresses the
findings of the biodiversity assessment and presents a preliminary
monitoring plan.
The objectives of Phase II were to:
* conduct a baseline
biodiversity assessment of target
groups at the San Martin-3 and
Cashiriari-2 well sites,
* select and establish multiscale vegetation test plots to estimate species/area relationships and
determine the number and size of
long-term biodiversity research
plots needed to monitor vegetation
and other selected groups, and
* study the area and gather data
from trails and other survey sites.
The assessment was targeted to
five biologically diverse groups
vegetation (trees, shrubs, ferns),
invertebrates (insects, snails, spiders), amphibians and reptiles
(frogs, snakes, lizards), birds, and
mammals. Fifty-two people participated in Phase II. Thirty-nine
researchers from several Peruvian
and foreign institutions worked on
the field assessments, seven focused
on data management and interpretation, and six provided technical
support. Six guides from nearby
Machiguenga communities assisted

in the field research.
This executive summary begins
with a description of the well sites
where the biological assessments
were conducted, followed by a list
of primary findings and recommendations concerning current operations and for the longer term. The
remainder of the document presents
the long-term monitoring plan.
The Well Sites
The San Martin-3 and

Cashiriari-2 well sites are located
near the Camisea River in La
Convencin Province, Department
of Cusco, Per. Brief descriptions
of the two sites follow.
San Martin-3
San Martin-3 is found approximately five kilometers (km) north
of the Camisea River at an altitude
of 474 m (geographical reference
coordinates are 11 47' 09.8"S, 72
42' 05.3"W). The area contains
steep hills (slopes range from 20 to
70) crossed by numerous rivers and
streams and an abundance of dense,
spiny, bamboo stands (Guadua
sarcocarpa) known locally as
"pacales." This terrain makes
walking difficult. For example, it
took researchers an entire afternoon
to obtain one undescribed species
of Pleurothyrium, a plant of the
avocado family (Lauraceae). The
site can be described as non-flooded
and wet and a canopy emerging
grassland with very few large trees,
a relatively abundant understory
layer, and numerous epiphytes and
lianas. The Cashiriari community is
located approximately 15 km southwest of the drilling site. We conducted the biodiversity assessment
at this site from January through
SPDP & Smithsonian/MAB Biodiversity Program, Lower Urubamba, Per
April of 1997, at the same time that

SPDP was constructing a 3.5-ha
platform for natural gas exploration.
Cashiriari-2
Cashiriari-2 is located approximately four km south of the
Camisea River and three km northeast of the Cashiriari River at an
altitude of 579 m (geographical
position is 11 51' 51.3"S, 72 46'
45.6"W). This site is characterized
by a hilly terrain with rather gentle
slopes (no rocky outcrops) dissected
by small creeks. Soils are, for the
most part, deep and clayey. The
vegetation surrounding the drilling
site is covered by primary, wet,
closed forest with numerous representatives of different botanical life
forms such as trees, herbs, epiphytes, and lianas. Large trees are
common, the understory vegetation
is moderately abundant, and there
are no bamboo stems. Within the
area of the platform, adult pioneer
species were observed. They apparently colonized the site after it was
cleared in the 1980s. The Cashiriari
community is located approximately
six km northwest of the drilling
site. We carried out the biodiversity
assessment at Cashiriari-2 from May
through June of 1997, while SPDP
was conducting drilling operations.
Findings
Overall, we found that the

biological diversity at each of the
two sites was extraordinarily high.
In addition, for each of the taxa
studied, the biological community
appeared to be in nearly pristine
condition. There was virtually no
evidence that human activities have
had significant impacts. The main
perturbations that may affect
biological communities are 1) direct
habitat losses, 2) changes in the

quality of the habitat, and 3) siltation. Direct habitat losses are the
obvious result of constructing
operational facilities. To date, the
areas occupied by those facilities
are small islands in a sea of green
vegetation. Changes in the quality
of habitat occur in the areas that
are opened and along their edges
where pioneer forest species are
exposed to more sunlight and higher
levels of temperature, humidity,
and wind. These factors may change
the species composition of the
disturbed areas or the surrounding
forest, and changes in habitat will
affect the structure of biological
communities. Soil washing into
streams from the construction sites
may cause abnormal levels of
siltation, although this does not
now appear to be a problem in the
small streams near the well sites,
based on the presence of tadpoles
and invertebrate aquatic fauna.
More specifically, our findings
indicate the following:
* Tree species densities proved
to be among the highest known to
date anywhere in the world. It is
well understood that the flora of
southeastern Per comprises a large
number of species. At our onehectare (ha) permanent biodiversity
monitoring plots, we found, on
average, 251 species of trees represented by 1500 individuals with
diameters equal to or greater than
four centimeters (cm). Many species
of herbs, lianas, and epiphytes were
also abundant in the forest.
* The Lower Urubamba may be
one of the most diverse areas on
earth for nocturnal butterflies. The
region contains a high diversity of
invertebrates, including many in the
groups we are studying (wasps,
bees, butterflies, dragonflies, ants,
Executive
Summary
For each of
the taxa
studied, the
biological
community
appeared to
be in nearly
pristine condition.
iii
Executive
Summary
Jaguar, ocelot,
and tapir
activity was
evident very
close to the
well sites.
iv
spiders, and snails, among others).

We found, for example, 87 species
of nocturnal butterflies (subfamily
Ctenuchinae) in just 18 days of
field work, or approximately 85%
of the 103 species found for
Tambopata, Per, during a much
more intensive study. We also
discovered new supra generic
taxaSclerogibbidae and
Scolebithydaefor Peruvian entomofauna. This finding enhances
understanding of biogeography in
the study area since the
Scolebithydae is also reported in
tropical Africa and Brazil.
* Amphibians (e.g., frogs and
toads) and reptiles (e.g., lizards and
snakes) are very important components of all tropical forests. We
documented more than 80 species
of amphibians and reptiles at the
San Martin-3 and Cashiriari-2 well
sites, which included 43 amphibians
and 44 reptiles.
* Bird species richness at the
well sites may equal or exceed that
of equivalent habitats in Manu
National Park, Per, one of the
most diverse areas for birds known
in the world. We recorded 198 bird
species at San Martin-3 and
Cashiriari-2. In addition, the bird
communities (including game birds)
at the well sites appear to be largely
undisturbed.
* We sampled 51 bat species at
the well sites during just one month
of field work. This total is very high
when compared to the 55 bat
species recorded at Manu National
Park over five years and the 57
found in French Guyana after 10
years.
* Neither the well operations
nor the noise they generate seemed
to limit large mammal activity,
except directly on the drilling pad.
We found that 58 species of me-
dium- and large-sized mammals

were potentially present at the well
sites. Jaguar, ocelot, and tapir
activity was evident close to the
sites. Big cats are often the first
animals to flee human development,
but these animals were active to
within 100 meters of the San
Martin-3 well site.
Key
Recommendations
Based on our findings to date,

we recommend the following
actions.
Current Operations
* Helicopter transport, key to
SPDPs commitment to minimize
environmental impacts, should
continue. Our findings show that
noise produced by vehicles and
machinery at the well sites is having
few, if any, effects on game animals,
which exhibit a distinct lack of
wariness upon encountering humans. (Part of this behavior is likely
due to low hunting pressure in the
vicinity of the well sites.)
* Botanical studies should be
continued. Research on the plants
used by local Native communities
should be added to the results on
current growth rates and distribution and should increase understanding of the natural cycles of
species abundance and scarcity.
* Fisheries studies should also
be continued. Fish are particularly
important because they are a major
source of protein for Natives. A
biological assessment of fish fauna
along the Urubamba and Camisea
rivers will provide baseline data for
work on fish migration cycles,
reproductive patterns, and growth
rates, as well as understanding of
the natural cycles of fish species.
The Longer Term

The following outline lists
important recommendations for the
next phase of the Lower Urubamba
biodiversity project. The final
section of this executive summary
(pages xxvi-xxxiv) presents a more
detailed description of the second
recommendation concerning establishment of a monitoring program
for the five target groups.
* Complete baseline
biodiversity assessments at the well
sites.
* Establish a long-term monitoring program for target groups
vegetation, invertebrates, amphibians and reptiles, birds, and mammalsat all of the well sites.
* Conduct a biodiversity assessment of the pipeline area and
design a monitoring program for
selected points along the pipeline.
* Complete an assessment of
selected points within six geological
regions identified on the satellite
images.
* Use the study process as an
opportunity to train Natives in
natural resource management.
* Develop educational materials and films along with a training
package for use with local communities and the public in the region.
* Convene a symposium in
1998 to present project findings;
publish the proceedings of the
symposium.
A. Introduction:
The Lower
Urubamba Region
The 170,000-ha study area

(defined as the area that includes
the four proposed appraisal well
sites and the proposed centers for
logistical activityNuevo Mundo
and Camisea) is located east of the

Andes Mountains in the Urubamba
River valley of the Ucayali watershed (see map, page 28). It is
bounded by the northern spurs of
the Vilcabamba Mountains to the
west and Urubamba Mountains to
the east between 10 and 13
latitude and 72 and 74 longitude.
The areas lowlands, hills, and
mountains vary in elevation, from
less than 500 m (lowland rain
forest) and 500 to 1,000 m (highland rain forest) to 1,000 m or more
(montane cloud forest). Slopes
range from 25% to 70%.
The Pongo de Mainique along
the Urubamba River separates the
upper and Lower Urubamba regions.
The Urubamba River flows north
into the Ucayali River, a major
tributary of the Amazon, and the
Camisea River drains the Urubamba
Mountains into the Urubamba. In
the southern portion of the area,
the Urubambas gorges intersect the
land as it rises to the highlands
around Cusco.
The great hilly landscape
consists of steep slopes located in
the Amazonian plains. This landscape stretches across 71% of the
study area. Its rolling hills, averaging 300 m in height, originated from
orogenic and tectonic events combined with the shaping action of
erosive process. The great mountainous landscape occupies about
20% of the study area. It comprises
relief forms of large dimensions
and slopes of up to 70%. Soils are
superficial with abundant extrusion
of meteorized rocks visible at the
surface. The mountains were
formed from tectonic processes that
occurred during the Jurassic and
Cretaceous periods. These areas are
characterized by a wide variety of
lithologic material. The largest
Executive
Summary
Executive
Summary
A factor that
may contribute to high
species
diversity for
both plants
and animals
in the
Urubamba
River valley is
its status as a
potential
Pleistocene
refuge.
vi
distribution of this landscape is in

the southern portion of the study
area.
Little is known about most of
the plant and animal species that
inhabit the Lower Urubamba region. The study area compresses
many different climatic zones, soil
types, and eco-geological patterns
in a relatively small space, suggesting an ecosystem that is complex
and high in heterogenity. A factor
that may contribute to high species
diversity for both plants and animals in the Urubamba River valley
is its status as a potential Pleistocene refuge. It is thought that
during the Pleistocene ice ages, the
climatic conditions in the Amazon
Basin changed so that the forest
retreated into isolated pockets and
was replaced by savannas. The
remaining forest refuges served as
incubators for new species of
rainforest organisms, accounting for
some of the observed biodiversity
in the Amazon. The Urubamba
valley (including the study area) has
been proposed as one such site.
As noted above, the highland
rainforest starts at around 500 m in
altitude and extends upward to the
cloud forest at 1,000 m. Floristically it is similar to the adjacent
terra firme lowland forests, but it is
characterized by higher rainfall
(more than 3,200 millimeters per
year), a more fractured topography
and, consequently, better drainage.
There are no flooded areas, and the
torrential white-water streams
contain rapids where the underlying
rock surfaces are exposed. Ridges
of native rock may occur. The socalled Fitzcarrald vegetation
apparently also reaches into areas
above 500 m.
The area in which the well sites
are located is peopled by indigenous
groups, most of whom are

extablished in Native communities,
with some colonist settlements
along the Urubamba River. The
communities of Nuevo Mundo,
Camisea, Shivankoreni, Segakiato,
and Cashiriari are all within the
study area. Just to the west (south
of Nuevo Mundo) and east of the
study area are the communities of
Kirigueti and Montetoni, respectively. Most of the inhabitants are
Machiguenga. The four communities closest to the well sites are
Cashiriari, Segakiato, Shivankoreni,
and Camisea. They are recent
settlements; all were established
since 1958. The combined population of these communities is slightly
less than 1,000 inhabitants. In
Cashiriari and Segakiato, the populations are relatively young (52% of
Cashiriari inhabitants are under the
age of 14, while in Segakiato, about
45% of the people are younger than
14).
Over the course of time, the
Machiguenga people have undergone a process of settlement into
distinct populations centers that are
now recognized as Native communities under Peruvian law. Each
community has ownership of land
with defined boundaries. The
Machiguenga usually settle along
river and stream courses that provide a means of navigation and
grounds for hunting and fishing.
Different populations have retained
strong bonds. Thus, it is not unusual for a family to visit other
families in different communities
for periods of several months,
traveling upstream on the Camisea
and Cashiriari Rivers. Travel for
commerce is usually along the
Urubamba River.
Executive
Summary
Figure 1. Adaptive management cycle for developing biodiversity assessment

and monitoring plan frameworks.
B. Biodiversity
Assessment
and Monitoring for
Adaptive
Management
Aspects of the adaptive management processsetting objectives, carrying out an assessment

and monitoring plan for forest
biodiversity, evaluation, and decision-makinghave been incorporated into the Lower Urubamba
project. This process can be represented as a cycle (Fig. 1), which is
periodically calibrated to ensure
that appropriate information from
each component feeds the next
level. The cyclical nature of the
process is maintained through
refining the objectives and management decisions based on the ongoing results. Thus, adaptive manage-
ment gives managers the flexibility

to adjust on-the-ground practices.
Adaptive management is well
suited to the type of project underway in Urubamba. The process
serves as a guide in planning and
carrying out a sound forest
biodiversity monitoring program. As
hydrocarbon exploration continues
in Urubamba, pressures to provide
adequate development of the
identified gas resource in conjunction with conservation of the
Urubamba region will likely increase, resulting from greater public
and private interest and involvement in use of the local resources.
An assessment and monitoring
process can aid in evaluating existing management approaches and
their impacts on the Urubamba
forest ecosystems and define new
approaches, if needed.
Management approaches should
be viewed as a means to test
vii
Space
Executive
Summary
Mammal
Community
Bird
Community
Amphibian
Community
Arthropod
Community
Weather Info.
Acid Rain
Soil/
Topography
Vegetation
T1=Baseline Measurement
T2=Second Census
Tn=Nth Census
Time
Figure 2. SI/MAB framework for forest biodiversity monitoring.

hypotheses to reach operational
goals. Through evaluation of the
monitoring data, managers receive
feedback as these hypotheses are
tested. Thus, evaluation is a tool for
improving management by checking
on management actions and providing guidelines for adjustments. If
the findings determine that
biodiversity trends are within the
expected values, monitoring can
continue without substantial alterations. If significant changes in the
trends are observed, managers or
decision-makers need to design the
most appropriate response, thus
restarting the cycle of setting
objectives, conducting assessments
and monitoring, and so forth. The
reasons for monitoring are also
evaluated at this stage. Is the
monitoring still required, and, if so,
do the objectives remain the same?
Long-term monitoring requires
a base or expected norm against
viii
which all future changes may be

compared. Baseline information is
normally taken from the literature
and/or initial surveys of the areas
under study. At San Martin-3 and
Cashiriari-2 no biological information was available before this study.
The initial assessments conducted
at the two well sites over the past
six months represent one of the
most comprehensive such efforts
ever undertaken by a large group of
scientists.
At the well sites, biodiversity
multi-scale and monitoring plots
have been incorporated along with
multi-taxa forest assessments to
construct the first biological picture
of the area. Subsequent monitoring
over time will provide a biological
movie of natural and anthropogenic changes (Fig. 2).
These long-term data are
helpful in detecting the magnitude
and duration of changes, how
related taxa are changing, and early

warnings of forest health. They also
serve as the basis for formulating
additional research and management hypotheses within the adaptive management framework. Any
results which are inconclusive
require adjustment of the objectives
and sampling approaches. However,
careful planning and design of the
monitoring program can reduce the
risk of inconclusive information.
It is no simple task to acquire
the knowledge necessary to understand changes in ecosystems,
especially considering that forest
biodiversity monitoring is a science
in the process of development. For
complex tropical forest ecosystems
such as in the Urubamba region,
researchers are still defining the
most appropriate methodologies to
sample and monitor biodiversity.
However, tools such as the adaptive
management process are improving
monitoring programs and their
capacities for making relevant and
reliable information available to
managers.
C. Assessment of
Biodiversity and
Long-term
Monitoring in the
Lower Urubamba
Region
The Lower Urubamba project

involves an assessment of
biodiversity in a large, natural
tropical forest and monitoring over
time to detect potential changes in
biological communities resulting
from local disturbances caused by
constructing and operating wells for
natural gas and condensate evaluation. Normally, only a small portion
of any given landscape can be
monitored because of cost constraints, but scientists and project

managers can make valid inferences
about larger, unsampled landscapes.
The location of long-term
monitoring plots should be based
on: 1) a better understanding of the
spatial distributions of species,
vegetation types, and habitat types;
2) spatially explicit models of biotic
and abiotic components and processes; and 3) quantitative information to determine how representative the long-term plots are compared with other plots and the
larger, unsampled landscape. Monitoring programs normally benefit
from a preceding systematic inventory of biotic and abiotic resources.
Revised protocols for vegetation
sampling, renewed economic incentives to improve monitoring programs, and recent advances in the
design of ecological assessments
have made it easier to improve the
design of integrated monitoring
programs. The Lower Urubamba
project includes a landscape-scale
assessment with the establishment
of monitoring sites and long-term
monitoring.
Landscape-scale assessments
begin by using remotely sensed data
to stratify a site into relatively
homogeneous ecological land units.
Randomly selected sampling sites in
each stratum allow for an unbiased
assessment of biodiversity hot
spots and identify habitats that are
less sensitive to land-use change
because of high redundancy (i.e.,
where the same habitat is abundant
nearby). Similar to findings of the
Oficina Nacional de Evaluacin de
Recursos Naturales of Per
(ONERN 1990), eight ecological
land-unit types were identified at
the Lower Urubamba site. These
appeared to differ in topography
Executive
Summary
ix
Executive
Summary
Multi-scale biodiversity
assessment plot
SI/MAB biodiversity
monitoring plot
Figure 3. Configuration of SI/MAB biodiversity assessment and monitoring test plots.

and drainage patterns (based on
elevation contour maps) and color
and texture (based on satellite
imagery). These sites are likely
different in vegetation composition,
successional stages, soils, and site
history.
In the Lower Urubamba, the
biodiversity plots are being established near the drilling sites to study
local changes in biodiversity in
habitats located in the selected
stratum. Here, the internationally
accepted SI/MAB design (one-ha
plot; Fig. 3) combined with multiscale assessment plots are followed
for assessing species diversity and
monitoring site-specific changes in
tree demography and changes in
plant and animal species diversity.
Additional long-term monitoring
x
plots will be needed in other strata

to evaluate changes in biodiversity
at landscape scales. Decisions
concerning the number of plots
needed are based on information
gained from the biological assessment and baseline work
The long-term monitoring
phase includes periodic remeasurement of the monitoring plots. The
time between measurements may
vary. A few test plots may be
assessed yearly at first to evaluate
seasonal variability.
D. Biodiversity
Assessment of
Vegetation in the
Lower Urubamba
Region
The flora of southeastern Per

comprises a large number of species. Topographic irregularities of
the eastern side of the Andes create
different micro-habitats where local
plants and animals survive. A high
plant diversity is usually correlated
with a high diversity of the organisms that depend on them. Most
animal species feed on the leaves,
flowers, and fruits that plants
produce. The greater the variety of
plants in an area, the greater the
number of animals they are likely to
support.
To date, most studies of
floristics in the Urubamba region
have been done in the upper portion. As part of the biodiversity
assessment of the Lower Urubamba
region, the research team sampled
the vegetation near the San
Martin-3 and Cashiriari-2 well sites.
Ground-truthing surveys as far as
one days walk from the well sites
were conducted. The assessment
entailed sampling trees, epiphytes,
lianas, herbs, and ferns that were
found in a fertile condition (i.e.,
flowering or fruiting). Trees, lianas,
and epiphytes were sampled by
climbing or with telescoping pruning poles.
Establishing the One-ha
Permanent Biodiversity Plots
To initiate the assessment and
monitoring of vegetation, multiscale sampling plots and one-ha
permanent biodiversity test plots
were established at San Martin-3
and Cashiriari-2. Plots #1 and #2
were set up at San Martin-3; plot #3

was set up at Cashiriari-2 (Fig. 4). A
brief description of the protocols
used in establishing the plots follows.
Topography teams delineate a
one-ha plot (10,000 m2) and divide
it into 25 quadrats, each 400 m 2 in
size, taking level measurements to
produce a detailed topographic map
of the plot. The boundaries of the
quadrats are demarcated using
string, which makes orientation
easier within the plot. Exact coordinates are determined with the aid of
a geopositioner. Close supervision is
needed to ensure the least amount
of disturbance to the vegetation
when the plots are set. After the
quadrats are delineated, the field
team locates, measures, marks, and
maps each tree that is at least four
cm in diameter at breast height
(dbh) in each quadrat. Diametric
tape is used to measure dbh, avoiding any protrusions on the trunk.
Where multiple stems occur, individual diameters are measured.
Trees are tagged with an aluminum
label that should face toward the
base line of the plot and set with a
nail above the point of measurement. The nails serve as a general
guide for future measurements.
Trees are tagged with a different
number consisting of a sequence of
three double digits. Using (01-2409) as an example, the first two
numbers (01) designate the zone
where the plot is located. The
second set (24) identifies the quadrat. The last two numbers (09)
represent an individual tree within
the quadrat. No other tree receives
this number. In each quadrat, the
tree numbers start at 1 and continue
until the last tree is labeled.
A team of three people maps
the position of the trees within the
Executive
Summary
At the well
sites, multiscale sampling
plots and oneha permanent
biodiversity
test plots were
established at
the two well
sites.
xi
Executive
Summary
quadrats. Two team members stand

at the corners of the quadrat, while
the third moves among the trees.
Electronic range finders measure
the distance from each tree to two
adjacent corners. The A and B
values recorded are later used,
along with the diameter, to calculate the exact position of the tree.
The field mapping teams use different preprinted data sheets for each
quadrat to record each trees position, dbh, and height as well as
pertinent notes about any tree or
quadrat. After the mapping teams
complete data gathering, botanists
record the identities of the trees on
the sheets, normally using six-letter
codes for the different species
mapped in the plot. Information

from the finished sheet is then
entered into BioMon (a program
developed by SI/MAB for data
management) by data entry teams.
BioMon generates a map of each
quadrat that field personnel use for
on-site verification of the correct
location of the measured trees.
Results of Vegetative
Sampling at the
Permanent Biodiversity Plots
Of the samples taken in the
study area, some exhibited geographical and morphological differences within the same species. The
flora of the Lower Urubamba
region is composed of a mixture of
Figure 4. Diagram showing the location of the biodiversity monitoring test plots, well sites, and
local settlements in the Lower Urubamba region.
xii
species that are commonly found in

other areas (e.g., species typical of
dry forests found at altitudes of 700
m in Kiteni to 1100 m in Potrero,
close to Quillabamba, Department
of Cusco, Per). Less than 10% of
the tree species found in the area
were deciduous (trees that drop
their leaves seasonally). Several of
the species sampled were classified
only as morphospecies (individuals
that share the same morphological
characters are grouped in the same
species). Several individuals of the
genus Inga spp. (Fabaceae) were
very difficult to classify. However,
with the existing comprehensive
database, literature, and collection
of photographs, the most common
species were determined.
Not all samples have been
identified to date. A few species
appear to be new to science, including one in the Acanthaceae family
and two terrestrial ferns. Other
records of interest are a possibly
new color form of Episcia fimbriata
and a species of Phinaea, relatively
rare in Per. In the Heliconiaceae
family, two of the Heliconia species
sampled seem to occur rarely in
Per.
Because the biodiversity plots
at San Martin-3 and Cashiriari-2
have different topographic features,
one would expect different vegetation types. Plots #1 and #2 at San
Martin are located in rugged terrain
with many hills and valleys. On the
ridges of these hills, thickets of
bamboo were often encountered. In
fact, bamboo accounted for up to
59% of all individuals in the plots
(when considering bamboo stems as
individuals). Its role in determining
the structure of the forest is likely
to be quite strong. During field
work, it became apparent that
heavy rains and wind knocked bam-
boo down and that this in turn had

an effect on neighboring tree species. Trees without crowns were
often found; the immediate impact
of this condition is a reduction in
photosynthetic products, thus
reducing tree growth. In addition, it
was apparent that several individuals damaged by bamboo died as a
result.
Biodiversity plot # 3 at
Cashiriari-2 is typical lowland forest
with few ridges or hills. The flora of
this site was somewhat different
from that at San Martin-3. In particular, the frequency of the species
seemed to be different (i.e., some
species found to be common at San
Martin-3 were rather rare at this
site). The vegetation lacked bamboo, while species such as Grias
peruviana and Calatola venezuelana
were quite common.
The numbers of tree species
were different at each permanent
plot (Fig. 5). Plots #1 and #3 had a
similar number of species (258 and
271, respectively) while plot #2 had
a lower count224 species. The
number of individuals in the plots
was markedly different (Fig. 6). Plot
#3 had a lower number of individuals, but many of those had large
diameters and were generally taller
than trees in the plots at San
Martin-3. A larger diameter resulted
in a larger basal area (Fig. 7). A
total of 16 species occurred at
densities of more than 1% (of all
non-bamboo individuals) in the
plots. The plot at Cashiriari-2
displayed much more equity in
densities among species than did
the plots at San Martin-3, although
at San Martin-3, there were only a
few common species, with the rest
occurring at low densities.
Certain similarities existed
among the forest plots in the Lower
Executive
Summary
With the
existing
comprehensive database,
literature,
and collection of photographs, the
most common species
were determined.
xiii
3500
250
3000
200
150
100
50
E. Biodiversity
Assessment of
Arthropods in the
Lower Urubamba
Region
In discussions of animal diversity, most people tend to think of

vertebrate species such as large
mammals, birds, and reptiles. These
constitute what have been called
the macroscopic fauna of tropical
forests. However, they represent
only a small fraction of forest
biological richness.
It is the less glamorous microscopic fauna that comprises the
largest portion of forest biodiversity. These organisms are
generally small to very small and
lead secretive lives.
The principal element of the
microfauna is the group known as
arthropods, which represent the
largest component of biological
diversity on the planet. Approximately 64% of all animal species
described to date belong to this
group (Fig. 8), a proportion that
continues to increase as more
studies in tropical regions are
40
Total Basal Area
300
# of Individuals
# of Species
Executive
Summary
Urubamba and SI/MAB plots in the

nearby Manu National Park. In
particular, the number of species
and their densities were quite
comparable. The major difference
was tree size. At Manu, many
species were 0.8 m or more in dbh.
In the Lower Urubamba, the tree
species were smaller with only a few
rare species such as Ficus mathewsii
exhibiting large stems. This difference may have to do with the
correlation between diameter and
bamboo. The presence of bamboo
can drastically reduce the girth of
co-existing tree species. In plots
where bamboo was absentthe
Manu plots and plot #3 at
Cashiriari-2the girth of trees was
considerably larger. Additional
effects of bamboo on the distribution and growth of other forest
plants will be clarified through
future research and monitoring.
2500
2000
1500
1000
500
0
0
P lot 1
(S M -3)
P lot 2
(S M -3)
S ite
P lot 3
(CA-2)
35
30
25
20
15
10
5
0
Plo t 1
(SM -3)
Plo t 2
(SM -3)
S ite
Plo t 3
(C A-2 )
Plo t 1
(SM -3)
Plo t 2
(SM -3)
Plo t 3
(C A-2 )
S ite
Figure 5 (left). Number of woody plants at each of the three plots (SM-3 = San Martin-3 and
CA-2 = Cashiriari-2). Figure 6 (center). Number of individual woody plants at the three sites.
The bars for Plots 1 and 2 show the number of individuals with and without the bamboo species
included. Plot 3 had no bamboo. Figure 7 (right). The total basal area of the three plots.
xiv
O t he r
Inve r t e b r a te s
19%
V e r te b r a t e s
3%
P la nt s
14%
A r t hr o p o d s
64%
Figure 8. The known global biota.

undertaken and new species are
discovered.
Arthropods are characterized
by bodies covered with hardened
walls and moving appendices that
are divided into distinct segments.
These animals include butterflies,
flies, mosquitoes, bees, wasps, dung
beetles, plant bugs, spiders, scorpions, mites, crabs, crayfishes, and
lobsters as well as a large number
of less well-known organisms.
Their generally diminutive size
does not stop arthropods from
performing critical functions in
forest ecosystems. They are important as herbivores, consuming the
living tissue of higher plants and
therefore regulating plant growth; as
predators in a broad sense, including as parasites that help regulate
animal populations; as pollinators,
assuring plant reproduction; and as
nutrient recyclers involved in
decomposition of dead organic
matter. Arthropods are also very
important as vectors of diseases
and pathogens to animals and
humans.
Executive
Summary
Approximately
64% of all
animal species
described to
date throughout the world
belong to the
arthropod
group.
Most arthropod species are

sensitive to environmental changes,
both spatial and temporal. Environmental gradients such as changes in
water availability and small differences in temperature will limit
species distribution. Temporal
changes in arthropod activity occur
daily, as different species are active
during the day and night. The
composition and density of arthropod communities for a given place
also change throughout the year;
many species have higher densities
during the rainy season, while
others are more abundant during the
dry season. Habitat heterogeneity
increases the number of species at a
larger scale, and structural patterns
of the forest have a profound
influence on arthropod composition
and abundancethey are found
from deep in the soil up to the top
of tree canopies.
In the study area, arthropod
sampling and preparation were
completed following standard
methods. The strata selected for
sampling were the forest floor and
leaf-litter layer immediately above
xv
Executive
Summary
The Lower
Urubamba
could be one
of the most
speciose
regions in
Per for
nocturnal
butterflies.
it, the understory layer, and the first

layer of vegetation. Special habitats
were also surveyed, including fallen
logs and fungi. Techniques were
selected so as to minimize overlap.
Sorting of sampled material is a
very time-consuming and laborintensive activity when dealing with
forest arthropod samples. We
attempted preliminary sorting and
identification of specimens in the
field. Once completed in a fully
equipped laboratory, the initial
sorting to ordinal level will enable
targeting of selected taxa for further
processing and study, while at the
same time yielding a gross estimate
of the arthropod biodiversity in the
study area. Voucher material will be
deposited in the collection at the
Museo de Historia Natural,
Marcos, Lima, Per, and at the
National Museum of Natural
History, Smithsonian Institution,
Washington DC.
Butterflies
Diurnal butterflies constitute
one of the most studied groups of
animals among the invertebrate
fauna. Adult butterflies are popular
with people because of their
brightly colored wings and their
observable daily activities that
include feeding on nectar as they
visit gardens. As with other insects,
diurnal butterflies have a life cycle
that can be completed in a few
months. By feeding on plants,
butterfly caterpillars have to deal
with the chemical composition of
the secondary compounds that
plants produce to reduce herbivore
attack. Thus, caterpillar studies
have enriched understanding of
animal/plant interactions, symbiotic
association, and the ecology and
behavior of parasitoids and preda-
xvi
tors. Butterflies are highly dependent on the microclimate conditions

of their habitats, and they play
important roles in ecosystems
because they serve as plant pollinators and plant feeders (i.e., herbivores) as well as prey for hundreds
of parasitoid species of wasps and
flies.
At San Martin-3 and Cashiriari2, the principal objectives for
studying diurnal butterflies were to
document the species present at
these sites and provide basic information about the natural history of
some of the species. This study is
of particular interest since the
highest diversity of butterflies in
the world has been documented in
the upper part of the Amazon River
in Colombia, Ecuador, Per, and
Brasil. As other sites in tropical
areas are studied, it is likely that
new species of this diverse groups
will be found.
At the well sites, we sampled
176 diurnal butterflies, including
more than 15% of the butterfly
species expected to be found.
Additional sampling must occur
during several seasons and over
several years to construct a picture
of total butterfly diversity in the
area and detect natural patterns of
change.
Nocturnal butterflies (i.e.,
moths) have been studied less than
diurnal butterflies, although we
know they are more diverse. There
are no significant samples of nocturnal lepidoptera for the Lower
Urubamba region. The fact that
researchers were able to sample 87
of the 103 species of the subfamily
Ctenuchinae reported for
Tambopata in southeastern Per
over just 18 days of field work
indicates that the Lower Urubamba
could be one of the most speciose
regions in Per for nocturnal butterflies.

Ants
Ants are particularly well suited
for inventory and monitoring programs. They are ideal bio-indicators
because they are diverse (approximately 9500 described species) and
found in abundance in almost every
habitat in the world. Ants are
numerically dominant in many
tropical forest canopy studies,
comprising 19% to 50% of the
individual arthropods. A study in
the Brazilian Amazon found ants to
contribute 80% of the insect biomass and more than 30% of all
animal biomass. These creatures are
important ecologically because they
function at many levels in an ecosystemas predators, prey, and
detritivores. Ants also have diverse
associations with plants; they are
seed dispersers, predators, and
herbivores.
Since ants are common and
apparent in almost every habitat,
they are readily sampled. Several
techniques can be used, depending
on the purpose of the project, the
habitat, and available resources.
Many ant species are highly sensitive to microclimate and habitat
structure and thus respond rapidly
to environmental changes. Since
they are social insects, ants live in
colonies that range from a few to
several million sterile worker ants
serving one to several queens. The
social structure of ant colonies
allows sampling of numerous
worker ants without significantly
affecting the reproductive unit of
the colony.
Researchers worked in 30
quadrats at the San Martin-3 well
site. Sampling methods used were
quantitative and can be repeated
over time, which makes them

appropriate for monitoring. Future
sampling of ants in the same location using these methods can help
detect and monitor changes in the
ant community at the biodiversity
plots and surrounding areas.
Nests of 55 ant species were
sampled, with a mean of 2.2 species
per 1-m 2 quadrat. A few species
were fairly common, but 27 (49%)
of the 55 species were sampled only
one time. This indicates that these
species are not common in the
forest near San Martin-3. Further
sampling in the area should provide
more information.
A mean of 4.3 ant nests were
found per 1-m 2 quadrat for a total
of 120 nests in 30 m 2. While 13
(43%) of the 30 quadrats had only
two or three ant nests, one quadrat
had nine nests and another had 11.
With a mean of 4.3 nests per m2,
one can estimate 43,000 ant nests
per ha. The number of worker ants
per nest varies greatly between
species; some have nests of 10 ants,
while others have nests of more
than a million individuals. Therefore, it is difficult to estimate the
number of individual ants per area,
but it is likely that their numbers
and biomass significantly exceed
that of all other animal species.
Given their importance in the
ecosystem through their various
roles, ants are certainly an essential
element in the functioning of the
forest at San Martin-3.
The 30 quadrats were not
sufficient to sample all ant species
in the leaf litter and soil. With each
new quadrat sampled, more species
were added, except that no new
species were added from quadrats
27 through 30. This leveling off of
the species accumulation curve
does not mean that no new species
Executive
Summary
xvii
Executive
Summary
New supra
generic taxa
for the
entomofauna
of Perthe
Sclerogibbidae
and
Scolebithydae
were
recorded.
xviii
will be added by sampling additional quadrats. Furthermore,

singleton speciesthose found in
only one quadratwere added as
more quadrats were sampled,
indicating that more rare species
will likely be sampled with each
additional quadrat. This type of
species accumulation curve, which
does not level off, indicates that
there are more species in the area
than were sampled and that additional sampling is needed.
hymenoptera fauna is even in this

areaone of the most biologically
rich places in the world. Interestingly, Trigona spp. bees were seen
gathering minerals from pipes at the
well sites. Further study of this
observation may prove helpful in
determining whether it is useful as
an indicator of pollutant levels, and
genetic studies of these bees can
give us an idea of the effects that
the chemicals may have on this
group.
Bees and Wasps

Bees and wasps are extremely
important in ecosystem functioning
since they are the primary pollinators of hundreds of flowering
plants. Many species of wasps
parasite other species of insects,
serving as population regulators of
potential pest species, and several
species of bees have evolved social
traits that makes them very competitive. The honey that bees
produce is used by indigenous
peoples in remote areas as an
important source of sugars and
proteins.
We sampled 54 species of bees
and wasps, likely just a small fraction of the diversity of the bees and
wasps that live at San Martin-3 and
Cashiriari-2. Twenty of the wasp
species and four species of bees
were found in the low stratum near
the ground and the leaf litter, while
24 wasp and six bee species were in
the medium and high strata.
The researchers reported new
supra generic taxa for the entomofauna of Perthe Sclerogibbidae
and Scolebithydae. This finding
enhances understanding of the
biogeography of the area since the
Scolebithydae is also reported in
tropical Africa and Brasil, and it
confirms how poorly known the
Dragonflies and Damselflies

Dragonflies and damselflies are
excellent subjects for biodiversity
studies. They are relatively
well-known compared to other
groups of tropical insects, they are
easily sampled and studied, and
they are important in benthic
communities that are routinely used
to assess the quality of surface
water. Adults are large, colorful,
day-flying insects that can often be
identified to genus as they fly.
Larvae are aquatic and often encountered in benthic studies. Both
adults and larvae are large predators
near the tops of their respective
food chains. Sampling is relatively
nondisruptive because local populations of these insects generally have
high numbers of individuals, and
only a small proportion is taken.
At San Martin-3, we sampled
the larvae of these species by
cutting into bamboo internodes and
then examining the contents in
enameled pans. It was confirmed
that possibly two species of helicopter damselflies occupied the top
predator niche in the system, and
that a number of mosquitoes, other
flies, and coleoptera were present.
The amount of water available in
the bamboo internodes was estimated to be to be about 10,000 1/
ha. A cross-tabulation of the raw

data yielded a listing of about 40
morphospecies by nine habitat
types.
Spiders
Spiders are found worldwide in
all terrestrial environments. Members of this invertebrate group have
a great ability to find prey and to
disperse. Individuals are nocturnal
predators, mostly of other invertebrates. They have developed a
variety of ways to capture their
food: some are orb-weavers, others
are sheet-web weavers, others are
cursorial hunters, and still others
are expert at ambush. All play an
important role in controlling the
populations of their prey. Spiders
are particularly numerous in the
tropics, where the abundant,
species-rich vegetation supports
great numbers of their prey. Habitat
complexity and changes in temperature and precipitation (i.e., seasonality) have strong influences in the
distribution and abundance of
spider communities in ecosystems.
Land-use practices also affect the
species composition of spiders.
Spiders are very diverse, as
evident from the fact that 34,000
spider species have been reported
around the world. At least one-half
of them are found in neotropical
America. Some 1120 genera of
spiders have been recognized in
neotropical areas, and at least onehalf of those genera are found in
Per. In the first assessment of the
spider diversity in the Lower
Urubamba, we identified a total of
69 morphospecies of spiders.
Snails
Terrestrial mollusks include
snails, sloths, and pseudo-sloths
(veronicellidos). The estimated
number of these species in the

world is between 30,000 and
35,000, or more than the total
combined number of species of
mammals, birds, reptiles, and
amphibians. Most known mollusk
biodiversity is found in the tropics;
the number could be even larger
because much of the tropics remains unexplored.
The terrestrial mollusks from
San Martin-3 were classified in 34
species33 snails (mollusks with
shells) and one veronicellid
(pseudobabosa with no shell).
Thirty-two species were found in
biodiversity plot #1. Most of the
species were in a reproductive state
as indicated by the presence of
eggs. Variation in species abundance was high, from abundant
species with 207 individuals to four
species that were found only once.
More than half (19) of the species
were represented by fewer than 10
individuals. More than 70% of the
species registered at San Martin-3
lived on the leaf litter and the
humus. Only three species of
arboreal snails were found.
Executive
Summary
In the first
assessment of
the spider
diversity in
the Lower
Urubamba,
we identified
a total of 69
morphospecies.
Beetles
Beetles have great potential for
biodiversity monitoring because of
the multiple roles they play in
ecosystems and because of their
sensitivity to environmental
changes. Beetles are well adapted
for excavation; that is, their legs are
very strong to help them move large
quantities of soil and food. They
are very important for nutrient
recycling, leading to a great deal of
interest in studying them. Their
ability to recycle animal dung has
been used to improve the cultivation of grasses and has been connected to disease transmission
because they move eggs of para-
xix
Table 1. Comparison of the known amphibian fauna for tropical sites
Executive
Summary
in Per (the number of amphibian species found in the Lower

Urubamba region at San Martin-3 and Cashiriari-2 [data from Reynolds et al. and Icochea and Mitchell this volume] are compared to
Manu [data from Morales and McDiarmid 1996] and Iquitos [data
from Rodriguez and Duellman 1994] )
Family
Bufonidae
Dendrobatidae
Centrolenidae
Hylidae
Leptodactylidae
Microhylidae
Pipidae
Ranidae
Urubamba
Manu
Iquitos
3
4
3
6
1
27
26
4
7
8
11
30
1
sites that affect human health to the

soil surface, thus enhancing parasite
transmission to hosts. Beetles are an
important component of the diet of
many insectivore birds. For example, birds consumed up to 85.4%
of these animals in the terra firme of
central Amazonia and 49% in Manu
National Park. Fluctuations in
beetle populations have been
correlated to behavioral changes in
several species of mammals. Fortyseven beetle morphospecies distributed in 13 genera in three subfamilies were sampled in the study area.
F. Biodiversity
Assessment of
Amphibians and
Reptiles in the
Lower Urubamba
Region
Amphibians and reptiles comprise an important component of

the vertebrate fauna in tropical
forest regions. They exhibit high
diversity and ecological specialization with habitat preferences, and
they are generally amenable to
xx
50
37
7
2
1
quantitative sampling. The primary

objective of the two amphibian and
reptile assessments conducted in
the study area was to begin establishing baseline data on amphibians
and reptiles present at the two well
sitesinformation that can be
incorporated into the multi-taxa
monitoring program to help detect
changes in the ecosystem and assist
in management of natural resources
in the area.
Selected specimens of amphibians and reptiles encountered during
the two assessments were sampled
and preserved as vouchers to
document individual species occurrence and to begin examining and
clarifying distributions and taxonomy of the herpetofauna in the
region. In the first assessment, a
total of 504 specimens were
sampled, documenting more than 80
species of amphibians and reptiles
from San Martin-3 and Cashiriari-2.
There were substantially more
specimens per taxa of amphibians,
but more families and genera of
reptiles even though reptiles constituted only 18% of the total number
of specimens sampled. In addition,
researchers recorded four additional

taxa at San Martin-3 that were not
sampled. They also captured and
released one caiman from a small
creek east of camp and positively
identified the call of a tree frog.
That vocalization constituted the
only record for this species. The
botanists reported that they encountered a tortoise while mapping trees
in biodiversity plot #1. Several
species of lizards were captured
with the sticky traps.
Several of the amphibian
species reported in this study have
been found at other lowland tropical sites in southeastern Per91%
in Cocha Cashu and 78% in
Pakitza. Table 1 compares the
amphibian faunas of the study area
and at nearby Manu National Park
and in Iquitos. The reptile fauna
also overlaps with those at Cocha
Cashu and Pakitza72% and 67%,
respectively. Fair comparisons of
the results from San Martin-3 and
Cashiriari-2 are not possible at this
time because the sampling effort
was more extensive at San Martin-3.
Researchers also sampled 84
specimens of amphibians and
reptiles to determine the genetic
diversity of the species of the area.
Fifty blood samples (for the micronucleus test) of 21 species were
taken, 3000 nuclei per individual
are being processed, and 51 samples
from 18 species were taken to test
for chromosomic alterations.
G. Biodiversity
Assessment of Birds
in the Lower
Urubamba Region
Birds are probably the most
widely studied vertebrate animal
group, primarily because of their
great diversity, their considerable
influence on the environment, and

their place in human and cultural
history. In all communities, even
those dominated by humans, birds
are important consumers, predators,
and prey of other organisms. Their
role in controlling insect populations, including agricultural pests, is
well known. In many environments,
birds are the key dispersal agents of
plants, usually by spreading seeds
after feeding on a variety of fruits,
cones, and nuts. A few important
groups of birds have evolved as
exclusive plant pollinators, feeding
on flower nectar and pollen. In their
roles as seed dispersers and pollinators, birds serve as mobile links
among various food webs. In this
capacity and as predators and
consumers, some birds are keystone species that largely determine the structure and function of
natural communities. Ultimately,
birds act to shape habitat for other
organisms, including other birds. In
addition, the nests of birds serve as
habitat for a wide range of organisms, and the birds themselves are
hosts to a suite of specialized
parasites and commensals.
The bird communities of the
Amazonian lowlands of Per are
among the most diverse in the
world. However, relatively few sites
within this vast area have been
surveyed by ornithologists. In
southeastern Per, good information exists for several sites in Madre
de Dios province, including Manu
National Park, the Tambopata
Reserve, and the Reserva Cusco
Amaznico. In contrast, little work
has been done in the lowlands of
Cusco Province. The upper course
of the Urubamba River, the areas
major drainage, has been extensively surveyed; the Lower
Urubamba region has not.
Executive
Summary
xxi
100
90
Executive
Summary
# of Species
It appears that
the species
accumulation
curve for birds
at San Martin-3
exceeds that of
Manu
National Park.
80
70
60
50
40
30
20
10
San Martin
Cashiriari
Manu
Panama
Brazil
Costa Rica
0
0
61
87
100
109
171
200
248
283
300
323
359
400
440
550
Captures
Figure 9. Species accumulation curves for birds at San Martin-3 and

Cashiriari-2 (diamonds = San Martin-3, squares = Cashiriari-2). The line
with triangles indicates the species accumulation curve from Manu National Park, derived from Karr et al. 1990. The line with the x represents
Panama, the asterisk Brazil, and the circle Costa Rica.
We sampled a total of 198
species at San Martin-3 and
Cashiriari-2, 65 of which were
observed at both sites, 100 only at
San Martin-3, and 33 only at
Cashiriari-2. The lower number of
species observed only at Cashiriari
is likely a result of the shorter
amount of time researchers sampled
there. An examination of the
number of species recorded just
within a single habitat indicates that
the Lower Urubamba sites are
indeed similar to others in southeastern Per. Data from mist-net
sampling at San Martin-3 and
Cashiriari-2 show that at San
Martin-3, 87 species were sampled
in a total of 440 captures (representing 360 individuals). At
Cashiriari, 62 species were sampled
in a total of 248 captures (representing 224 individuals).
xxii
As with many other tropical

sites, there were only a few common species and many rare ones in
the samples. At San Martin-3, 83%
of the species captured each made
up 2% or less of the total captures,
while at Cashiriari-2 the corresponding figure was 77%. It appears
that the species accumulation curve
at San Martin-3 exceeds that of
Manu National Park (Fig. 9).
There was little evidence of
breeding activity at either site. In
the mist-net samples, no birds were
observed with brood patches. In
contrast, many individuals were in
molt at both sites. At San Martin-3,
64% of all mist-netted birds examined (316) showed some sign of
molt. The percentage at Cashiriari2, 43%, was somewhat lower.
Large game birds such as
tinamous, guans, curassows, and
trumpeters are typically among the
first species to disappear in response to human activity in an area,

chiefly as a result of over-hunting.
At San Martin-3, small groups of
Spixs guan were noted regularly,
and the blue-throated piping-guan
was seen on three occasions. The
razor-billed curassow and palewinged trumpeter were reported in
the area by other observers. Three
species of tinamougreat, cinereous, and black-cappedwere
observed. Tinamous are secretive
and generally recorded primarily by
call. Calling activity by many species is low during the late rainy
season, and it is possible that
surveys at another season would
record additional species of
tinamou. The frequency of encounter of these game species plus their
relative lack of wariness when
encountered indicate that hunting
pressure in the immediate area of
San Martin-3 is fairly low.
Large parrots, macaws, and
toucans also are liable to disappear
rapidly because of human activity;
they are hunted not only for food
but also for the pet trade. As mentioned above, these species were
rare or uncommon at San Martin-3.
However, their low density at this
site is more likely to be the result of
forest structure than of hunting
pressure, which is almost nonexistent.
The avifauna of San Martin-3
and Cashiriari-2 is highly diverse,
and the species richness at these
sites may equal or exceed that of
equivalent habitats in Manu National Park, one of the richest
known sites for birds in the world.
At present, the bird community
appears to be in nearly pristine
condition, with little evidence that
human activity to date has had
significant impacts. Every effort
should be made to see that this

exceptional avifauna does not suffer
degradation as a result of future gas
development in the area.
H. Biodiversity
Assessment of
Mammals in the
Lower Urubamba
Region
Small Non-volant Mammals
The small, non-flying mammals
are estimated to constitute between
15% and 25% of the mammal fauna
in tropical rainforests. These widespread, abundant animals occupy a
variety of habitatsarboreal,
terrestrial, and semi-aquatic
depending on their different morphological adaptations and thus are
excellent candidates for assessment
and monitoring studies.
At San Martin-3 and Cashiriari2, this group was sampled along
trails, in streams, and near the
drilling platforms using baited traps.
Six species of marsupials and 13
species of small rodents were found
(Fig. 10). Despite the relatively
short amount of time spent sampling at the two well sites (15 days
at each location), the findings were
sufficient to allow comparisons with
the nearby areas of Manu and
Cusco Amazonico. At San Martin-3,
the sample rate was 5%; at
Cashriari-2, it was 2%. The sample
rates for Manu and Cusco
Amazonico were 1% and 5%,
respectively.
The higher number of species
at San Martin-3 than at Cashiriari-2
may be due to the presence of
bamboo at San Martin. In addition,
the forest structure at San Martin-3
may provide more places for shelter,
Executive
Summary
Every effort
should be
made to see
that this
exceptional
avifauna does
not suffer
degradation
as a result of
future gas
development
in the area.
xxiii
20
18
Executive
Summary
16
# of Species
14
12
10
8
6
4
San Martin - 3
C ashiriari - 2
0
0
10
11
12
13
Sampling Days
Figure 10. Species accumulation curves for small, non-volant mammals at

San Martin-3 and Cashiriari-2.
sources of food, and ways to reduce
predation.
Bats
Bats are the only group of
mammals that can fly. They comprise the second largest order of
mammals with 177 genera and 925
species. They occur throughout the
tropical and temperate regions of
the world, where they occupy a
wide array of ecological niches.
Bats are mostly active at night,
finding shelter during the day in
caves, tree cavities, and crevices. In
cities, they can be found roosting
on buildings, under bridges, and in
tunnels. Bats are known to
echolocate (i.e., navigate and find
food through a system of high
frequency sounds and their echoes).
They can catch insects while flying
using echolocation, but they also
feed on mammal blood, fishes,
small vertebrates, and plant material such as pollen, nectar, and
fruits.
Bats play essential roles in
ecosystem functioning. They consume large quantities of insects,
they aid in the pollination of hunxxiv
dreds of flowering plant species

that have evolved to open their
flowers at night, and they serve as
seed dispersers by feeding on fruits.
Farmers use the nutrient-rich guano
(feces) produced by bats to fertilize
fields.
A total of 51 different species
of 31 genera and five families of
bats (379 individuals) were sampled
at San Martin-3 and Cashiriari-2.
Thirty species were located at San
Martin-3 and 43 at Cashiriari-2. Of
these, 22 were found in both localities, eight were found only at San
Martin-3, and 21only at
Cashiriari-2. The species accumulation curve for data gathered at each
site (Fig. 11) suggests different
patterns of diversity for each
locality. As a comparison, 55 bat
species were recorded over five
years at Manu National Park and 57
in French Guyana after ten years.
Medium and Large Mammals
Because of growing concerns
about the management of biological
systems possibly impacted by oil
and gas development, biodiversity
surveys have been suggested as a
method to develop baseline data

about those systems. Monitoring
processes to follow up on the
baseline surveys are being developed and tested. At the well sites in
the Lower Urubamba, the objective
of the assessment was to conduct
medium- and large-mammal surveys
in preparation for monitoring.
A list of species that could be
expected to occur at the two well
sites was compiled from range maps
and distribution descriptions. The
species list reflected the sum of
four sources, which were then
confirmed as present or not by the
score derived from accumulated
evidence at each site. Fifty-eight
species of mammals were considered potentially present at the well
sites. Confirmation was achieved
for the presence of 25 species at
San Martin-3 and 26 species at
Cashiriari-2 for a total of 33 nonduplicated species. The total number of confirmed species was 57%
of the potential list. Twenty-four
or 41% of the possible species
were not confirmed as present, but
could be present, even common.
Some monkeys, canopy-inhabiting

species, species with large territories and/or low mobility, and
nocturnal animals can easily avoid
survey methods in rainforest. All
species listed have a reasonable
probability of living on, or occasionally moving through, the study
area. Furthermore, the agouti, small
cats, and armadillo species have
very similar tracks and sizes and are
difficult to distinguish without
specimens. These species have a
tendency to be lumped.
Difficulties were encountered
in distinguishing the signs of the
small cats, particularly margays,
ocelots, and jaguarundis. During
this study, measurements were
taken of four sets of tracks, all
identified as ocelot tracks. They
were found within two hours over
three km and were likely different
individual cats. All tracks were
fresh on wet sand or clay and of
excellent quality. Cat tracks with
different heel pad configurations
and ranging in size from 2.5 to 5.0
cm across were also observed. It
was assumed that at least two
Executive
Summary
50
45
40
# of Species
35
30
25
20
15
10
San Martin - 3
Cashiriari - 2
0
0
10
11
12
13
14
Sampling Days
Figure 11. Species accumulation curve for bat species sampled at San
Martin-3 and Cashiriari-2.
xxv
Executive
Summary
Ocelot, tapir,
and peccary
tracks, beds,
and other
signs were
found within
100 meters of
the well sites.
species of small cat were present,

probably margay and ocelot; however, further studies are needed to
verify these assumptions.
General observations indicated
that the species expected at the
sites were present and that numbers
or densities of those populations
were in expected ranges. Large
mammal activity around the San
Martin-3 construction area and the
Cashiriari-2 drilling site was close
and continuous; the noise and
operations did not appear to have
any severe impacts other than
directly on the drilling pad. Red
brocket deer and Brazilian rabbit
tracks were present at both well
sites. Ocelot, tapir, and peccary
tracks, beds, and other signs were
found within 100 m of the drilling
pads. Cat species that are often
considered the first to flee human
development (ocelots, pumas, and
jaguars) were also active to within
100 m of the pad at San Martin-3.
A major loafing and toilet area for
jaguars and pumas was found
approximately 2.5 km from
Cashiriari-2 well site, where the
engines could clearly be heard.
Sightings and signs of large mammals did not appear to be any less
numerous near the drilling activity
than 2.5 km away. The observations
and signs at the well sites were
sufficient to develop a density
estimate for 28 species. San
Martin-3 clearly had a higher abundance index (1643) for most animals than did Cashiriari-2 (1028).
Parasites of Mammals
Of the sampled bats, 207
specimens were reviewed for ectoparasites. A 68.5% rate of infestation was found. Approximately 93%
of the small mammals analyzed
were also infested. One hundred
xxvi
fifty-five specimens of ectoparasites were sampled from small

mammals at the two well sites.
These specimens were classified
into eight families and twenty-two
genera. Forty-five species were
found for the first time in the Lower
Urubamba.
I. Biodiversity
Monitoring Plan
for the Taxonomic
Groups Under
Study in the
Lower Urubamba
Region
As noted throughout this

document, there is a need for a
scientifically sound monitoring
program at the SPDP well sites and
along the proposed pipeline. The
program will help track trends in
the groups under study in the Lower
Urubamba and aid in determining
changes in forest structure and
ecosystem functions that may occur
as a result of construction and
operations at the well sites. The
following discussion centers around
the rationale for monitoring in each
of the groups and provides specific
recommendations for inclusion in
the monitoring program design.
Vegetation
Biological systems, such as
tropical forests, are extremely
complex networks of biotic and
abiotic interactions. While all
components of a complex system
are essential, plants comprise
perhaps the most important structural and functional component of a
forest ecosystem. Their roles are
many, from the obvious to the
subtle. Forest plants are crucial to

multi-taxa monitoring as well.
Some plants such as fig trees
are considered keystone forest
species because of their role in
supporting numerous species of
birds and bats during seasons when
fruits are not readily available.
Without keystone species, many
bird and bat population levels
would decline, leading to decreases
in populations of plants that depend on these animals for seed
dispersal or pollination.
The scientific implications of
forest biodiversity and the critical
role that forest plants play with
respect to humans cannot be overstated. There are tremendous
economic, medical, and agricultural
benefits to studying forest plants. In
addition to their value in finding
cures for diseases that plague
people, tropical forest plants provide a natural genetic warehouse
that helps maintain many domesticated agricultural species.
Work to date in the study area
has provided a baseline distribution
of tree species and a description of
habitats at a particular site. The
process of monitoring change in
these plots over time will enable an
deeper understanding of the impacts of natural and human-caused
disturbances on species and on
community composition. Data from
the monitoring program can also be
used to predict future changes,
based on current knowledge of
ecosystem processes, structures,
and functions.
The monitoring program objectives include the following:
* Continue obtaining vegetation baseline information at the
well sites and in the general area
Block #75 where SPDP is currently
operating. This objective includes
continuation of the assessments

and surveys already initiated, thus
allowing quick identification of
different forest sites, estimates of
species diversity, and descriptions
of vegetative characteristics. The
amount of data will increase
through the incorporation of more
sites and the collection of data on
species dynamics, as we sample in
different locations and at diferent
times.
* Gain a better understanding
of habitat types, vegetation types,
and spatial distributions of species
at the well sites, along the pipeline,
and in the general area of influence
of the gas development project. It
is important that the implications
of human activity on vegetation
patterns and forest regeneration be
thoroughly studied. Changes in
forest structure caused by human
activity at the well sites and along
the pipeline will induce regeneration of plants in the disturbed areas.
The species that dominate these
sites will be pioneering secondgrowth species that will form a
forest quite different from the
surrounding undisturbed forest. The
influence of this successional
sequence will have significant
impacts on the animals currently
inhabiting the area. It is thus recommended that the vegetation in the
undisturbed forest and regenerating
forest, both near communities and
SPDP operation facilities, be studied and analyzed.
* Obtain quantitative information about how representative the
forest monitoring plots are compared to the multi-scale vegetation
plots and the larger unsampled
landscape. Vegetation studies
require data from numerous sites in
the forest to obtain a representative
sample. Since this forest is very
Executive
Summary
One hundred
fifty-five
specimens of
ectoparasites
were sampled
from the
small mammals studied
at the two
well sites.
xxvii
Executive
Summary
An important
issue in
monitoring
and assessing
biotic shifts
in ecosystems is the
ability to
provide spatial/temporal
referencing
for biotic
data.
xxviii
large and it is possible to sample

only a few hectares, it is important
that the sites be representative of
the larger area.
Arthropods
Arthropods (insects, spiders,
mites, and relatives) are the most
biologically diverse group of organisms in terrestrial ecosystems.
Recent estimates place the global
arthropod fauna between 10 and 30
million species. Arthropods can
provide information on virtually all
macro- and micro-habitats within an
ecosystem. They cover several size
classes; exhibit a range of ecosystem requirements (highly specific to
generalist), dispersal abilities, life
cycles, and development times;
assist in mediating ecosystem
functions such as decomposition
and in maintaining soil structure
and soil fertility; regulate populations of other organisms (including
arthropods, vertebrates, and plants);
respond quickly to environmental
changes; and act as mobile links
essential to the reproduction of
many flowering plants. Information
derived from arthropod species
assemblages can be used to characterize accurately almost any aspect
of an ecosystem.
An important issue in monitoring and assessing biotic shifts in
ecosystems is the ability to provide
spatial/temporal referencing for
biotic data. To analyze biotic shifts
associated with climate or anthropogenic change, biotic data must be
comparable in space and time and
placed within the context of natural
ecosystems (i.e., data from modified
systems must be compared with
data from the appropriate natural
counterpart). Both sampling protocols and the methods used to record
and structure abiotic data associ-
ated with the protocols must be

standardized. The ecosystem framework used to implement a sampling
and monitoring program must also
be standardized.
Vegetation structure, considered as multiple-scale canopies,
provides the ecosystem framework
that can be used to organize monitoring of arthropods. Data acquired
using appropriate protocols within
this framework may then be compared to data from communities
across gradients in and among
ecosystems or to analogous communities in highly disparate ecosystems. The recognition that ecosystems possess several scales of
canopies has implications both for
the design of a sampling program
and for the protocols used to conduct it. For instance, canopy sampling protocols should be applied
vertically along the aridity gradient
within sampling sites (i.e., low, mid,
and high canopy of trees and
herbaceous vegetation) and horizontally along the hydrological
gradient represented by the transition from riparian zone to high
elevation points. The pulse fauna
(locally transient fauna), the most
diverse arthropod community
assemblage, can then be characterized using specific protocols and
not be confused with the arthropod
fauna endemic to the litter, soil,
canopy, and aquatic communities.
The latter groups can also be characterized with appropriate protocols
used in association with the respective vegetation canopy.
It is not possible to monitor all
species or to assess changes in a
few species without a baseline
reference to the appropriate natural
ecosystem. The acquisition of
biodiversity baselines from unmodified sites is an extremely important
component of a monitoring strategy. Baselines are data sets against

which similar, usually smaller,
collections of data can be viewed in
perspective to provide an interpretation that reflects ecosystem
reality. Baselines set the stage for
development of monitoring technology while acquiring a database
that functions as a powerful tool in
the analysis of ecosystem change.
Implementation of monitoring
programs can then be based on a
broad and detailed knowledge of
the species.
Arthropod Monitoring
Strategy
For the Lower Urubamba
project, researchers have recommended that the arthropod baseline
be restricted to those groups that
are taxonomically feasible (i.e. their
taxonomy is known and the taxonomist is available to determine the
specimens). The arthropod baseline
for each site should consist of at
least two biodiversity plots: one
situated in the forest type most
likely to be affected by gas well
operations; the other, a control plot,
situated in the same forest type in a
nearby area unlikely to be affected
by gas well operations or pipeline
construction. Subsequent monitoring programs must use the same
biodiversity plots from which the
baselines were acquired. Each
modate available expertise from the
international systematics community.
Arthropod assessment and
monitoring is underway in the study
area, following the basic precepts
described above. The objectives are
to:
* acquire arthropod faunal
baselines for at least two
biodiversity plots, including one
control plot, at each of gas well

sites;
* in subsequent years, monitor
selected components of the arthropod fauna representative of size
classes, dispersal abilities, and
trophic levels in the soil, litter, and
canopy communities existing across
major physical and chemical gradients in the biodiversity plots;
* quantify the rate of
biodiversity change in arthropod
species assemblages that can be
associated with natural gas extraction;
* assess the impact of changes
in arthropod species assemblages in
the study area; and
* develop the arthropod faunal
inventory of the region using data
obtained from the baselines and the
monitoring program.
Substantial progress has been
made in acquiring initial data for
the baselines. During the next
phases of the project, additional
work on baseline acquisition and
the main thrust of the monitoring
program will take place in the three
stepsstaff training, acquiring the
arthropod baseline, and monitoring.
The assessment and monitoring
strategy presented in this plan can
be expected to generate information
on most levels of the ecosystems
surrounding the gas well sites.
Information will be gathered on
biodiversity in the soil, litter,
vegetation, and canopy communities at the sites. An estimated
15,000 species of arthropods will
be included in the biodiversity
analysis in the most cost-effective
manner possible.
Executive
Summary
One of the
objectives of
the monitoring program
is to assess
the impact of
changes in
arthropod
species assemblages in
the study
area.
Amphibians and Reptiles

Amphibians are a diverse group
of predatory animals that occur on
all continents but reach their
xxix
Executive
Summary
greatest abundance in tropical

regions. Most possess complex life
cycles, an aquatic larval stage, and
a terrestrial adult stage that is
unique among vertebrates. Each
individual is thus exposed to
aquatic, terrestrial, and atmospheric
perturbations during its life time.
This life history, combined with a
moist, sensitive skin used for
respiration and lack of longdistance migratory abilities, renders
individuals and populations susceptible to a variety of environmental
modifications. They are usually the
first vertebrates to respond to
environmental change.
Amphibians worldwide have

responded to large-scale perturbations by undergoing population
decline, range reduction, and extinction. Causes of decline include
habitat loss and degradation, acid
precipitation, UV-B radiation,
pathogens, exotic species, harvesting by humans, and natural population fluctuations. The most common causes of range reduction,
population decline, and alteration
of community structure are habitat
loss and degradation.
The primary perturbations
likely to occur at the well sites that
may affect amphibians are: 1) direct
Figure 12. Example of nocturnal micro-habitats of selected frog species in the

Lower Urubamba region.
xxx
habitat loss at each construction

site 2) changes in the quality of
rainforest habitat adjacent to each
site (i.e. edge effect), and 3) siltation of streams in the watershed
from soil washing from the construction sites. Since most species
are adapted to forest conditions, the
number of species that occurred
originally at each construction site
has already declined. However, at
least two species survive in the
open areasthe marine toad and a
leptodactylid frog. Once the
rainforest is opened by a disturbance such as clearing for a well
site, the trees and other vegetation
that occurred inside the forest are
exposed to higher levels of sunlight,
temperature, humidity, and wind.
The composition of the vegetation
at the forest edge changes in response to the new environmental
conditions. Because many rainforest
amphibians are adapted to interior
forest conditions, few tolerate open
areas or forest edges (Fig. 12).
Those that can live in the ground
and breed in open puddles or are
drawn from the edge vegetation by
the lights that attract insects.
Siltation does not appear to be a
current problem in the small
streams at San Martin-3 and
Cashiriari-2 (based on the presence
of tadpoles), although additional
land excavation and site operation
may have greater effects in the
future. Any such changes in habitat
affect the structure of the amphibian communities.
A monitoring program for
amphibians in the Lower Urubamba
will help determine whether
changes in species distribution have
taken place because of habitat
alterations. If conducted for several
years, it can also determine which
species, if any, have been impacted
by past operations and whether

changes in the fauna occur over
time in response to large-scale
environmental alterations. Changes
in the fauna due to climate shifts or
atmospheric perturbations are not
expected in the near future. However, analysis of changes in amphibian abundance and distribution
must take into account possible
changes in the abiotic environment.
The primary goals of the amphibian monitoring plan are to:
* evaluate the effects of the
construction and operation of each
well site and the proposed pipeline
on frog communities,
* evaluate the long-term trend
in amphibian community structure
and species distributions relative to
habitat and climate changes in the
Lower Urubamba, and
* add to the faunal inventories
and knowledge of the natural
history of each species at each site
using data from the monitoring
project. These goals will be met by
using standard monitoring protocols.
Executive
Summary
Analysis of
changes in
amphibian
abundance
and distribution must
take into
account
possible
changes in
the abiotic
environment.
Birds
Although tropical bird communities are typified by high species
richness, many species are rare,
making them vulnerable to local
extinction caused by human activity. Part of the high diversity of
these communities is due to the
high habitat specificity of the many
species that are found in only one
or a few habitats or microhabitats
and are unable to survive if that
habitat is altered or degraded.
It is not feasible to monitor all
conceivable impacts of development on bird populations in the
study area. Therefore, this discussion is limited to the impacts of the
minimal expected future develop-
xxxi
Executive
Summary
xxxii
ment in the area if full-scale gas

production (i.e., installation of
permanent production wells, their
long-term operation, and the construction of an underground gas
pipeline with subsequent reforestation of the pipeline route) is initiated. It is assumed that no permanent access road will be built into
the lower Urubmba from outside,
that SPDP or other authorities will
continue to control access, and that
uncontrolled colonization will not
take place. If these assumptions
prove false, additional monitoring
programs will be needed to examine
the effects of forest fragmentation
or widespread deforestation.
Given these assumptions, the
major impacts of development will
be those caused by the presence of
the well sites and the pipeline. The
actual area of forest that would be
removed for each well site and the
pipeline is small in comparison to
the entire project area, so the direct
effect of this deforestation on bird
populations should also be small.
However, the potential indirect
effects of creating the clearings
must also be taken into account. It
is possible that the presence of
these clearings within the forest
could affect bird populations over a
much wider area, due to the edge
effect and species colonizations.
SPDP forbids hunting near the
well sites, either with firearms or
other by any other means. Once a
pipeline is constructed, controlling
access to the project area may be
more difficult, even if the pipeline
route is reforested. Therefore,
monitoring game bird species along
the pipeline is desirable. In addition, placing patrols in conjunction
with Native communities along the
route may help control unauthorized access. The effectiveness of
this method could be assessed

through a monitoring program.
Indigenous hunters and some
environmental groups have speculated that noise and human activities at the well sites might adversely
affect the forest fauna in the area,
possibly by frightening animals
away. Based on findings from this
study, researchers hypothesize that
noise (including that from helicopters) and human activity in the
absence of hunting, other direct
persecution, and habitat alteration
will likely not have a major longterm effect on the forest fauna.
This question could be examined by
instituting studies of game species
at the well sites and along the
pipeline.
The following priorities are
recommended for the monitoring
program:
Well Sites
Initially, monitoring should
include San Martin-3 and
Cashiriari-2. Additional sites should
be included if new wells are established in substantially different
habitat types. The priorities are:
* Establish a system of survey
areas at each well site to include
two control sites and corresponding
base camps.
* Initiate a mist-net survey in
these survey areas to assess edge
effects.
* Initiate a point-count survey,
also to assess edge effects.
* Establish transect routes and
initiate transect surveys to monitor
game birds and other exploited
species.
Pipeline Route
Monitoring should be carried
out at a minimum of two sites, one
in a few kilometers of the point
where the pipeline starts and one

within intact forest. The priorities
are:
* Establish a system of three
survey areas at each pipeline study
site that includes one control area
and corresponding base camp.
these areas to assess edge effects.
species.
* Initiate a point-count survey
to assess edge effects.
Training
A three-week orientation and
training course conducted by expert
ornithologists from Per and elsewhere will go far in creating a cadre
of experienced field workers for the
Lower Urubamba project and
related bird monitoring programs.
Mammals
Small Non-volant Mammals
The monitoring plan recommends the use of standardized
methodologies for long-term monitoring at the well sites and in areas
that will be affected by development and extension of the gas line.
The principal objective is to determine the distribution and abundance of small, non-volant mammal populations. This will require
efficient trapping procedures to
optimize sampling while also facilitating a rapid and reliable comparison of the studied variables. The
proposal calls for selecting an
adequate number and type of
habitats directly related to the use
of a variety of traps and methods
(audio and visual) used to identify
the mammals.
Bats
In the Lower Urubamba, as
with elsewhere in the world, there is
no single method suitable for
monitoring all bats. A combination
of observation techniques and
sampling methods should be employed for various bat species.
These include direct roost counts,
disturbance counts at roosts, nightly
dispersal counts, direct visual
counts, counts with motion detectors, ultrasonic bat detection, and
direct capture.
Executive
Summary
Medium- and Large-sized

Mammals
Currently, monitoring activities
are tracking what happens to the
ecosystem after planning is completed and gas exploration begins.
That is important. But to ensure
that major challenges resulting from
future exploration and development
are resolved, baseline monitoring
should be accomplished along with
the planning effort.
Researchers emphasize that
significant challenges will likely
arise with the placement of the
pipeline and associated facilities.
The pipeline route from the fields
to Lima and location of the gas
processing plant, pumping stations,
pipeline maintenance facilities, and
access are all critical. Monitoring of
biodiversity will provide useful
information so that decision-makers
can carefully plan and, if needed,
adjust routing and locations to
avoid ecological alterations.
Based on such reasoning, the
following four tasks have been
delineated for large mammals during
the next phases of the Lower
Urubamba biodiversity assessment
and monitoring project. While the
focus is on large mammals, the
xxxiii
Executive
Summary
xxxiv
basic rationale for the tasks holds

for other taxa in the study area.
The following priorities are
recommended for the monitoring
program:
* Conduct monitoring of
mammal diversity at San Martin-3,
Cashiriari-2, and related facilities
associated with gas production.
* Conduct biodiversity monitoring along the proposed pipeline
route from Urubamba to Lima and
at facilities associated with delivery.
* Develop training manuals and

support materials to assist future
researchers in their environmental
assessment efforts; e.g., compile
track, scat, and sign keys and
biomonitoring technique manuals
for South American mammals.
* Train Peruvian biologists in
biodiversity monitoring techniques
for large mammals.
Los Pozos de
Exploracin
San Martn-3 y
Cashiriari-2
Francisco Dallmeier y
Alfonso Alonso
Biodiversity Program (SI/MAB)
Sinopsis
Las zonas bajas de la amazona
peruana son ricas biolgicamente y
proporcionan una gran oportunidad
de integrar ciencia, conservacin y
desarrollo a travs de una cuidadosa
planificacin, evaluacin y toma de
decisiones. En la regin baja del ro
Urubamba, el Institute of Conservation Biology del Smithsonian
Institution se ha unido con Shell
Prospecting and Development
Resumen Ejecutivo
Evaluacin de la
Biodiversidad y
Monitoreo en la
Regin Baja del
Ro Urubamba
(Per) B.V. (SPDP) en un proyecto

nico encaminado a lograr el
desarrollo de los recursos de gas
natural y condensados en una forma
sensible al medio ambiente.
Nuestros colaboradores incluyen
cientficos peruanos del Museo de
Historia Natural de la Universidad
Mayor de San Marcos, Universidad
San Antonio Abad del Cusco, y la
Universidad Nacional de la Libertad
(Trujillo).
SPDP est actualmente
evaluando los reservorios de gas en
los pozos de San Martn-3 y
Cashiriari-2 que fueron descubiertos
durante la exploracin realizada a
mediados de los 80s. Esta
evaluacin consiste en la
perforacin de 4 pozos
exploratorios y el establecimiento
de una base logstica de
operaciones. Los planes para el
desarrollo adicional de los
reservorios de gas incluyen tuberas
de flujo, una planta para el
procesamiento de gas ubicada en el
rea de Camisea, y un gaseoducto
que cruce los Andes hasta el
Ocano Pacfico en donde tambin
se construirn instalaciones
costeras.
Las metas principales del
trabajo del Smithsonian son las de
integrar la informacin recabada de
la vegetacin con datos de otros
grupos taxonmicos que tambin se
estn evaluando y monitoreando, y
aplicar estos resultados a las
decisiones de manejo en los pozos,
y ms tarde, en el rea en general.
Equipos multidisciplinarios de
investigadores estn llevando a
cabo este proyecto bajo el marco de
investigacin y entrenamiento
diseado por SI/MAB (Smithsonian
Institution/Monitoring and Assessment of Biodiversity Program) para
el monitoreo a largo plazo de varios
xxxv
Resumen
Ejecutivo
Llevamos a
cabo la
evaluacin de
la
biodiversidad
de grupos
seleccionados
en los pozos
de San
Martn-3 y
Cashiriari-2.
xxxvi
grupos taxonmicos de gran

diversidad en sitios de investigacin
permanente.
Se han completado dos fases
del proyecto. Primero se realiz una
evaluacin de la diversidad
biolgica y cultural de la regin baja
del Urubamba. Esta fue
complementada con un taller en
Septiembre de 1996 (realizado en
Lima) sobre el rea de estudio, para
establecer los parmetros a seguir
del programa de evaluacin y
monitoreo de la biodiversidad. Los
participantes incluyeron
representantes de instituciones
gubernamentales, nogubernamentales, la comunidad
cientfica, y los grupos nativos
locales. El reporte de la fase I fue
publicado a principios de 1997.
La fase II incluy una
evaluacin inicial de la
biodiversidad en la zona de estudio
y el establecimiento de parcelas
permanentes para el monitoreo de
la biodiversidad. El SI/MAB Series
#1, producto de este trabajo
Evaluacin de la Biodiversidad y
Monitoreo en la Regin Baja del Ro
Urubamba - Pozos de exploracin
San Martn-3 y Cashiriari-2
provee la base para este resumen
ejecutivo. El reporte revela los
resultados de la evaluacin de la
biodiversidad y presenta un plan de
monitoreo preliminar.
Los objetivos de la Fase II
fueron:
* llevar a cabo la evaluacin de
la biodiversidad de grupos
seleccionados en los pozos de San
Martn-3 y Cashiriari-2,
* establecer parcelas de
vegetacin multi-escala para estimar
las relaciones especie/rea y
determinar el nmero y tamao de
parcelas de investigacin necesarias
para monitorear la vegetacin y

otros grupos seleccionados, y
* estudiar el rea y registrar
datos a partir de trochas y otros
lugares de inters.
Esta evaluacin fue enfocada a
cinco grupos biolgicamente
diversosvegetacin (rboles,
arbustos, helechos); invertebrados
(insectos, caracoles, araas);
anfibios y reptiles (ranas, culebras,
lagartijas); aves y mamferos.
Cincuenta y dos personas
participaron en la fase II. Treinta y
nueve investigadores de varias
instituciones peruanas y extranjeras
trabajaron en la evaluacin de
campo, siete se concentraron en el
manejo e interpretacin de datos, y
seis proporcionaron apoyo tcnico.
Seis guas de comunidades
Machiguenga cercanas al sitio de
estudio ayudaron en el trabajo de
campo.
Este resumen ejecutivo
comienza con una descripcin de
los pozos donde se condujo la
evaluacin de la biodiversidad,
seguido por una lista de los
hallazgos principales y
recomendaciones concernientes a
las operaciones actuales y futuras.
El resto del documento presenta
detalles adicionales tomados del
reporte general.
Los Pozos
Los pozos de San Martn-3 y

Cashiriari-2 estn ubicados cerca
del ro Camisea en la Provincia de
la Convencin, Departamento del
Cusco, Per. A continuacin se da
una breve descripcin de los dos
sitios.
San Martn-3
San Martn-3 se encuentra
aproximadamente a cinco
kilmetros (km) al norte del ro
Camisea a una altitud de 474 m (las

coordenadas de referencia
geogrfica son 11 4709.8"S, 72
4205.3"W). El rea contiene
colinas muy empinadas (la
pendiente vara de 20 a 70)
cortadas por numerosas quebradas
y arroyos, y una abundancia de
densos macizos de bamb (Guadua
sarcocarpa) conocidos localmente
como pacales; caminar en este
terreno es difcil. Por ejemplo, a los
investigadores les tom toda una
tarde obtener una especie an no
descrita de Pleurothryrium, una
planta de la familia de la palta
(Lauraceae). Adems del bamb, el
rea tiene una cubierta de bosque
abierto que puede describirse como
hmedo no-inundable, y un dosel
emergente con relativamente pocos
rboles grandes sobresaliendo del
gramadal conformado por el
bamb; una capa de sotobosque
relativamente abundante, y
numerosas lianas y epfitas. La
comunidad nativa de Cashiriari est
ubicada aproximadamente a 15 km
al sur-oeste del pozo. La evaluacin
de la biodiversidad de este pozo fue
realizada de enero a abril de 1997,
al mismo tiempo que SPDP estaba
construyendo la plataforma de 3.5
ha, necesaria para la exploracin
del gas natural.
Cashiriari-2
Cashiriari-2 est ubicado
aproximadamente a cuatro km al
sur del ro Camisea y tres km al
noreste del ro Cashiriari a una
altitud de 579 m (su posicin
geogrfica es 11 o5151.3"S, 72o
4645.6"W). Este lugar est
caracterizado por un terreno con
colinas y con laderas relativamente
poco empinadas (sin rocas
descubiertas) disectadas por
pequeas quebradas. Los suelos
son, en su mayor parte, profundos y

de naturaleza arcillosa. La
vegetacin que rodea el sitio de
perforacin est conformada por
bosque hmedo cerrado con
numerosos representantes de
diversas forms de vida botnicas
tales como rboles, hierbas, epfitas
y lianas. Los arboles grandes son
comunes, el sotobosque es
moderadamente abundante y no hay
bamb. Dentro del rea de la
plataforma se observaron especies
adultas de rboles pioneros. Estos
aparentemente colonizaron el sitio
despus de que fue deforestado en
los 80s. La comunidad nativa de
Cashiriari est ubicada
aproximadamente a seis km al
noroeste del punto de perforacin.
Efectuamos la evaluacin de
biodiversidad en los alrededores de
Cashiriari-2 de mayo a junio de
1997, mientras SPDP estaba
realizando operaciones de
perforacin.
Resumen
Ejecutivo
La
composicin
de cada uno
de los grupos
taxonmicos
estudiados se
encontr en
condicin
pristina.
Hallazgos
Realizamos la evaluacin de la
biodiversidad mientras se estaba
construyendo la plataforma en San
Martn-3 y durante las operaciones
de perforacin en Cashiriari-2. En
general, la diversidad biolgica en
cada uno de los dos lugares fue
extraordinariamente alta.
Adems, la composicin de
cada uno de los grupos taxonmicos
estudiados se encontr en condicin
pristina. Prcticamente no hay
evidencia de que las actividades
humanas hayan tenido un impacto
significativo. Las perturbaciones
principales que pueden afectar a las
comunidades biolgicas son:
1) prdida directa de hbitats,
2) cambios en la calidad del
ambiente, y 3) sedimentacin.
xxxvii
Resumen
Ejecutivo
xxxviii
La prdida de hbitat es el resultado

obvio de las operaciones de
construccin de la plataforma.
Hasta la fecha, las reas ocupadas
por stas son pequeas islas en un
mar verde de vegetacin. Cambios
en la calidad del habitat ocurren en
las reas que estn abiertas y a lo
largo de sus bordes, donde especies
pioneras del bosque estn expuestas
a mayor cantidad de luz y niveles
ms elevados de temperatura,
humedad y viento. Estos factores
pueden cambiar la composicin de
las especies del rea perturbada o el
bosque circundante, y los cambios
en hbitat tendrn efecto en la
estructura de las comunidades
biolgicas. El lavado del suelo hacia
cursos de agua desde los lugares de
construccin puede causar niveles
anormales de sedimentacin, pero
sto no parece ser un problema por
ahorra dada la presencia de
renacuajos e invertebrados
acuticos en los arroyos cercanos a
los pozos.
Ms especficamente, nuestros
hallazgos indican lo siguiente:
* La densidad de especies de
rboles estan entre las ms altas
conocidas hasta ahora en el mundo.
Se conoce bien que la flora del
sureste del Per incluye un gran
nmero de especies. En nuestras
parcelas permanentes de una
hectrea (ha) para el monitoreo de
la biodiversidad, encontramos en
promedio 251 especies de rboles
representados por 1500 individuos
con dimetros iguales o mayores a
cuatro centmetros (cm). Muchas
especies de hierbas, lianas, y
epfitas fueron tambin abundantes
en el bosque.
* El bajo Urubamba puede ser
una de las reas ms diversa en el
mundo para mariposas nocturnas. El
bajo Urubamba contiene una alta
diversidad de invertebrados,
incluyendo muchos de los grupos
que estamos estudiando (avispas,
abejas, mariposas, liblulas,
hormigas, araas y caracoles entre
otros). Por ejemplo, en slo 18 das
de trabajo de campo encontramos
87 especies de mariposas nocturnas
ctenuchidas (subfamilia
Ctenuchinae), esto es,
aproximadamente el 85% de las 103
especies encontradas en
Tambopata, Per, en un estudio
ms largo. Tambin encontramos
nuevos grupos taxonmicos
supragenricos para la entomofauna
peruana -Sclerogibbidae y
Scolebithydae. Estos hallazgos
aumentan nuestro entendimiento de
biogeografa ya que los
Scolebithydae tambin estn
distribudos en Africa tropical y
Brazil.
* Los anfibios (e.g. ranas y
sapos) y reptiles (e.g. lagartijas y
culebras) son componentes muy
importantes de todos los bosques
tropicales. Durante el estudio
documentamos ms de 80 especies
de anfibios y reptiles en las reas de
los pozos de San Martn-3 y
Cashiriari-2, las cuales incluyeron
ms de 43 especies de anfibios y 44
de reptiles.
* La riqueza de especies de
aves en las reas que rodean a los
pozos, puede igualar o tal vez
exceder la de hbitats equivalentes
al Parque Nacional Manu, Per, una
de las reas ms diversas en aves
conocidas en el mundo. Registramos
198 especies de aves en San Martn3 y Cashiriari-2. Adems, las
comunidades de aves incluyendo las
aves de caza en los sitios de los
pozos, parecen estar muy poco
perturbadas.
* Registramos 51 especies de
murcilagos en los sitios de los
pozos durante solo un mes de

trabajo de campo. Este total es muy
alto cuando se compara con las 55
especies de murcilagos registradas
para el Parque Nacional del Manu,
durante un perodo de ms de 5
aos, y con las 57 especies
encontradas en Guyana Francesa
despus de 10 aos de estudio.
* Ni las operaciones del pozo
ni el ruido que ellas generan
parecieron limitar las actividades de
mamferos grandes, excepto
directamente sobre la plataforma de
perforacin. Hay 58 especies de
mamferos de tamao medio y
grande potencialmente presentes en
los sitios de estudio. Cerca a los
pozos, la actividad del otorongo,
pecary y sachavaca fue evidente.
Con frecuencia, los gatos grandes
(felinos) son los primeros animales
que huyen de las actividades
humanas, pero en San Martn-3,
stos estuvieron activos a 100
metros (m) de distancia de la
plataforma.
Recomendaciones
Claves
Basados en nuestros hallazgos

la fecha, recomendamos las
siguientes acciones.
A la Fecha
* El transporte en helicptero,
clave del compromiso de SPDP de
minimizar los impactos ambientales,
debe continuar. Nuestros hallazgos
muestran que el ruido producido
por vehculos y maquinaria en los
sitios de los pozos est teniendo
poco, si algn, efecto en animales
de caza, los que no se alertan al
encuentro con humanos (es posible
que este comportamiento en parte
se deba a la baja presin de caza en
la vecindad de los pozos).
* Los estudios botnicos

debern continuarse. Se deben de
investigar las plantas que las
comunidades nativas locales usan,
para enriquecer los estudios
actuales. Esta actividad tambin
proveer una oportunidad para
incrementar la participacin de la
gente local en las evaluaciones
biolgicas.
* Estudios sobre la pesca
tambin debern continuarse. Los
peces son particularmente
importantes porque son fuente
principal de protenas para los
nativos. La evaluacin biolgica de
la fauna de peces de los ros
Urubamba y Camisea proveer
informacin de base para trabajos
sobre ciclos migratorios, tasas de
crecimiento, as como un
entendimiento de los ciclos naturales de abundancia y escasez de las
especies acuticos.
Resumen
Ejecutivo
A Largo Plazo
A continuacin se presentan
recomendaciones importantes para
la siguientes fases del proyecto
sobre la biodiversidad del bajo
Urubamba. La ltima seccin de
este resumen ejecutivo presenta una
descripcin ms detallada en
relacin al establecimiento de un
programa de monitoreo para los
grupos taxonmicos seleccionados.
* Completar las evaluaciones
de biodiversidad para obtener
informacin de base en los pozos de
exploracin.
* Establecer un programa de
monitoreo a largo plazo para los
grupos seleccionados - vegetacin,
invertebrados, anfibios y reptiles,
aves y mamferos - en cada uno de
los pozos.
* Conducir una evaluacin de
puntos seleccionados en las 6
xxxix
Resumen
Ejecutivo
regiones identificadas en las

imgenes de satlite.
* Entrenar algunos miembros
de la poblacin nativa en el manejo
de recursos naturales.
* Desarrollar materiales
educativos y documentales como un
paquete de entrenamiento para uso
de las comunidades locales y el
pblico interesado en la regin.
* Organizar un simposium en
1998 para presentar los hallazgos
del proyecto y publicar las
memorias.
A. Introduccin: La
Regin Baja del Ro
Urubamba
Las 170,000 ha del rea de

estudio (definidas como el rea que
incluye los cuatro pozos propuestos
para la evaluacin y los centros
propuestos para la actividad
logstica- Nuevo Mundo y Camisea)
estn localizadas al este de los
Andes en el valle del ro Urubamba
en la cuenca del Ucayali (ver mapa
1, pgina 28). La regin est
limitada hacia el oeste por las
vertientes norte de las montaas de
Vilcabamba y hacia el este por las
montaas del Urubamba entre los
10 y 13 de latitud, y los 72 y 74
de longitud. El rea contiene valles,
colinas y montaas que varan en
elevacin, desde menos de 500 m
(zonas bajas de bosques lluviosos);
de 500 a 1,000 m (zonas altas de
bosques lluviosos); y de ms
de1,000 m (bosques nublados de
montaa o yungas). La pendiente
vara de un 25% hasta un 70%.
El Pongo de Mainique a lo
largo del ro Urubamba separa la
regin alta y baja del Urubamba. El
ro Urubamba se une al ro Ucayali
por el norte, uno de los mayores
afluentes del ro Amazonas, y el ro
xl
Camisea drena las montaas hacia

el ro Urubamba. En la parte sur del
rea, las quebradas del Urubamba
intersectan el valle a medida que se
elevan hacia las alturas cercanas al
Cusco.
Este gran paisaje montaoso
est formado por quebradas
empinadas que se encuentran en los
llanos de la Amazona. Este paisaje
se extiende en un 71% del rea de
estudio. Sus loms onduladas, de
una altura promedio de 300 m, se
originan por cambios orognicos y
tectnicos combinados con la
accin modeladora del proceso
erosivo. El paisaje de las grandes
montaas ocupa un 20% del rea de
estudio. Comprende los relieves de
las grandes extensiones y declives
hasta del 70%. El suelo es superficial, con frecuente exposicin de
rocas meteorizadas visibles en la
superficie. Las montaas se
formaron por procesos tectnicos
que ocurrieron durante los perodos
Jursico y Cretcico. Estas reas se
caracterizan por una amplia
variedad de material litolgico. La
distribucin ms amplia de este
paisaje se encuentra en la zona sur
del rea de estudio.
Poco se sabe de muchas de las
especies de plantas y animales que
habitan en la regin del bajo
Urubamba. El rea de estudio
agrupa zonas climticas, tipos de
suelo, y zonas geogrficas muy
diferentes en una area relativamente
pequea, creando un ecosistema
complejo y muy heterogeneo. Uno
de los factores que puede contribuir
a la gran diversidad de especies,
tanto de plantas como de animales
en el valle del ro Urubamba, es su
estatus como un posible refugio
Pleistocnico. Se sabe que durante
el Pleistocno, las condiciones
climticas en la cuenca del
Amazonas cambiaron de tal forma

que los bosque se retiraron hacia
lugares aislados y fueron
reemplazados por savanas. Los
refugios forestales que quedaron
sirvieron como almacn para el
desarrollo de nuevas especies de
organismos del bosque lluvioso, que
resultaron en los patrones de
biodiversidad observados hoy da
en la Amazona. El Valle del
Urubamba (incluyendo el rea de
estudio ) ha sido propuesto como
uno de estos sitios refugio.
Como mencionamos
anteriormente, las zonas altas del
bosque lluvioso comienzan a una
altitud aproximada de 500 m, y se
elevan hacia los bosque nubosos
hasta los 1,000 m. En cuanto a flora
se refiere, sta es similar a los
bosques bajos adyacentes de la terra
firme, pero se caracteriza por lluvias
ms fuertes (ms de 3,200
milimetros por ao), por una
topografa ms accidentada y
consecuentemente, por un mejor
drenaje. No hay reas inundadas, y
los cursos torrenciales de aguas
blancas contienen corrientes
rpidas, donde las superficies de las
rocas subyacentes son visibles; hay
laderas con rocas expuestas. La
llamada Vegetacin Fitzcarrald se
puede encontrar en reas sobre los
500 m.
El rea de localizacin de los
pozos est poblada por indgenas, la
mayora de los cuales estn
establecidos en comunidades
nativas con algunos asentamientos
colonizados a lo largo del ro
Urubamba. Las comunidades de
Nuevo Mundo, Camisea,
Shivankoreni, Segakiato y Cashiriari
se encuentran en el rea de estudio.
Hacia el oeste (al sur de Nuevo
Mundo) y al este de la zona de
exploracin estn las comunidades
de Kirigueti y Montetoni,
respectivamente. La mayora de los
habitantes son Machiguenga. Las
cuatro comunidades ms cercanas a
la zona de los pozos son Cashiriari,
Segakiato, Shivankoreni y Camisea,
cuyos pobladores son relativamente
jvenes (52 % de los habitantes de
Cashiriari tienen menos de catorce
aos de edad, mientras que en
Segakiato cerca del 45 % de la
poblacin tienen menos de catorce
aos).
Con el transcurso del tiempo,
los Machiguenga se han asentado en
distintos centros de poblacin que
son reconocidos como
Comunidades Nativas bajo la ley
peruana. Cada comunidad tiene
propiedad de la tierra con fronteras
definidas. Los Machiguengas
generalmente se establecen a lo
largo del ro y de las corrientes que
les proveen un medio de navegacin
y lugares para la caza y la pesca. Las
diferentes poblaciones han
mantenido fuertes lazos entre ellas.
No es raro que una familia visite
otras familias en diferentes
comunidades por perodos de varios
meses, viajando corriente arriba en
los ros Camisea y Cashiriari. Los
viajes comerciales son generalmente
a lo largo de ro Urubamba.
Resumen
Ejecutivo
Uno de los
factores que
puede
contribuir a la
gran
diversidad de
especies, tanto
de plantas
como de
animales en el
valle del ro
Urubamba, es
su estatus
como un
posible refugio
Pleistocnico.
B. Evaluacin y
Monitoreo de la
Biodiversidad como
parte del Manejo
Adaptativo
Los aspectos del proceso del

manejo adaptativo- tales como la
definicin de objetivos, la
evaluacin y monitoreo de la
biodiversidad, la evaluacin de
resultados y la toma de decisiones se han implementado en el proyecto
xli
Resumen
Ejecutivo
del bajo Urubamba. Este proceso se

puede representar como un ciclo
(Fig. 1), el cual es calibrado
peridicamente para asegurar que la
informacin apropiada de cada
componente alimente al siguiente
nivel. La naturaleza cclica del
proceso se mantiene a travs de la
evaluacin de los objetivos y de las
decisiones de manejo a nivel
gerencial con base en los resultados
que se van obteniendo. Por tanto, el
manejo adaptativo provee a los
gerentes la flexibilidad para hacer
los ajustes necesarios.
El manejo adaptativo es un
buen diseo para el tipo de proyecto
puesto en marcha en Urubamba. El
proceso sirve como gua para
planear y llevar a cabo un fuerte
programa de monitoreo de la
biodiversidad del bosque. Conforme
la exploracin de hidrocarburos
contine en Urubamba, las
presiones para proveer un desarrollo

adecuado del recurso demen
gas afn
con la conservacin de Ejecutivo
la regin,
seguramente se incrementarn,
como resultado de un mayor inters
y compromiso pblico y privado en
el uso adecuado de los recursos
locales. El proceso de evaluacin y
monitoreo puede ayudar en la
evaluacin de las condiciones de
manejo existentes y en sus impactos
en los ecosistems de bosque del
bajo Urubamba, definiendo nuevas
propuestas si son necesarias.
Las propuestas de manejo
deben ser vistas como un medio
para probar hiptesis para alcanzar
metas operacionales. A travs de la
evaluacin de los datos de
monitoreo, los gerentes reciben
informacin constante a medida
que estas hiptesis son puestas a
prueba. Es por esto, que la
evaluacin es una herramienta para
Figura 1. El ciclo del manejo adaptativo integrado al programa de

evaluacin y monitoreo de la biodiversidad.
xlii
Space
Resumen
Ejecutivo
Mammal
Community
Bird
Community
Amphibian
Community
Arthropod
Community
Weather Info.
Acid Rain
Soil/
Topography
Vegetation
T2=Second Census
Tn=Nth Census
Time
Figura 2. Esquema del programa SI/MAB para el monitoreo de la

biodiversidad.
mejorar el manejo a travs de las
acciones a nivel gerencial y del
abastecimiento de lineamientos para
su ajuste. Si los resultados
demuestran que la biodiversidad se
inclina hacia los valores esperados,
el monitoreo puede continuar sin
cambios substanciales. Si se
presentan cambios, los
investigadores y los gerentes
encargados de tomar decisiones
necesitan disear una respuesta ms
apropiada, tal como sera el reinicio
del ciclo de objetivos establecidos.
En esta etapa tambin se evalan
las metas para el monitoreo.
El monitoreo a largo plazo
requiere de una base o patrn que
sirva de referencia para futuros
cambios. La informacin de base es
tomada normalmente de la
literatura y/o de investigaciones
previas del rea de estudio. Para
San Martn-3 y Cashiriari-2 no se
encontr informacin disponible
previa a este estudio. La evaluacin

inicial conducida en las reas de
perforacin durante los ltimos seis
meses representa una de los
esfuerzos ms importantes
realizados hasta ahora por un grupo
de cientficos.
En las zonas de perforacin se
han establecido parcelas multiescalas de biodiversidad para la
evaluacin de varios grupos
taxonmicos para elaborar la
primera fotografa biolgica del
rea. El monitoreo a travs del
tiempo proveer la pelcula
biolgica con cambios naturales y
antropognicos (Fig. 2).
Esta informacin a largo plazo
es til para detectar la magnitud y
duracin de los cambios, cmo los
grupos taxonmicos relacionados
estn cambiando, y cules son los
sntoms de salud de los bosques.
Estos tambin sirven de base para
la organizacin de la investigacin
xliii
Resumen
Ejecutivo
requerida y de las hiptesis de

manejo a probar dentro del marco
de trabajo del manejo adaptativo.
Los resultados inconclusos
necesitarn de un ajuste de
objetivos y de muestreo. Sin embargo, la planificacin y el diseo
cuidadoso de un programa de
monitoreo pueden reducir los
riesgos de una informacin
inconclusa. No es una tarea fcil
adquirir los conocimientos
necesarios para entender los
cambios del ecosistema,
especialmente considerando que el
monitoreo de la biodiversidad de los
bosques es una ciencia en proceso
de desarrollo. Para ecosistems de
bosques tropicales complejos, tales
como los de la regin del
Urubamba, los investigadores estn
definiendo los mtodos ms
apropiados para el muestreo y
monitoreo de la biodiversidad. Sin
embargo, herramientas como el
proceso de manejo adaptativo estn
mejorando los programs de
monitoreo y su capacidad para
proveer informacin importante y
precisa a los gerentes.
C. Evaluacin de la
Biodiversidad y
Monitoreo a Largo
Plazo en la Regin
del Bajo Urubamba.
El proyecto del bajo Urubamba

involucra evaluacin de la
biodiversidad en bosques tropicales
naturales de gran tamao y
monitoreo a lo largo del tiempo para
detectar los cambios potenciales
que se den en las comunidades
biolgicas como resultado de las
perturbaciones locales causadas por
la construccin y operacin de
pozos de evaluacin de gas natural
xliv
y condensados. Generalmente, solo

una pequea porcin del terreno
puede ser monitoreado debido a
limitaciones de costos. Sin embargo,
con un buen diseo, tanto
cientficos como gerentes pueden
hacer inferencias vlidas acerca de
otras reas de gran tamao que no
han sido muestreadas.
Para determinar la ubicacin de
las parcelas a largo plazo se debe
tomar en cuenta lo siguiente: 1) un
mejor entendimiento de la
distribucin de las especies en el
espacio, los tipos de vegetacin, y
los tipos de hbitat; 2) los modelos
explcitos de espacio referentes a
componentes y procesos biticos y
abiticos; e 3) informacin
cuantitativa de cuan representativas
son las parcelas permanentes
comparadas con otras parcelas, y
con otras reas de gran tamao que
no han sido muestreadas. Los
programs de monitoreo
normalmente se benefician de un
inventario sistemtico previo de
recursos biticos y abiticos. La
revisin de protocolos para el
muestreo de la vegetacin, la
renovacin de incentivos
econmicos para mejorar los
programs de monitoreo, y recientes
avances en el diseo de
evaluaciones ecolgicas han hecho
ms fcil el mejoramiento del
diseo de programs de monitoreo
integrados. El proyecto del bajo
Urubamba incluye una evaluacin a
gran escala del paisaje, as como la
determinacin de lugares de
monitoreo, y el monitoreo a largo
plazo. La evaluacin del rea
comienza con el uso de sistems de
informacin geogrfica por medio
de sensores remotos para
estratificar el terreno en unidades
ecolgicas relativamente
homogneas. La eleccin de sitios
de muestreo al azar en cada estrato

permite una evaluacin imparcial de
lugares de alta biodiversidad, as
como la identificacin de hbitats
que son menos sensibles a los
cambios por el uso de la tierra
debido a que son frecuentes y se
repiten en otras partes del bosque.
Observaciones similares fueron
realizadas por la Oficina Nacional
de Recursos Naturales del Per
(ONERN 1990), identificando en la
zona del bajo Urubamba ocho tipos
de unidades ecolgicas. Estas zonas
muestran diferencias en cuanto a
patrones topogrficos y de drenaje
(basado en las lneas de elevacin
de los mapas), y en color y textura
(basado en imgenes de satlite).
Tambin se observa una posible
diferencia en la composicin de la
vegetacin, etapas de sucesin,
tipos de suelo, e historia del lugar.

En la zona del bajo Urubamba se
estn estableciendo parcelas de
biodiversidad cerca de los pozos de
perforacin, para estudiar los
cambios locales de la biodiversidad
en hbitats localizados en los
estratos seleccionados. El diseo
internacionalmente reconocido del
SI/MAB (parcelas de 1 ha; Fig. 3)
se utiliza en combinacin con
parcelas de escala mltiple,
seguidos por inventarios y por el
monitoreo de cambios especficos
dados en la demografa de los
rboles, as como los cambios en la
diversidad de especies de plantas y
animales. Se requerir de parcelas
adicionales de monitoreo a largo
plazo en otros estratos para evaluar
los cambios de la biodiversidad a
escalas mayores. La decisin
Resumen
Ejecutivo
Multi-scale biodiversity
assessment plot
SI/MAB biodiversity
monitoring plot
Figura 3. Diseo de las parcelas del SI/MAB para la evaluacin y monitoreo

de la biodiversidad.
xlv
Resumen
Ejecutivo
Se establecieron parcelas multiescala y

parcelas
permanentes
de 1 ha en
San Martn-3
y Cashiriari-2.
concerniente al nmero de parcelas

que se pueden necesitar est basada
en la informacin recopilada para la
evaluacin biolgica y de lnea de
base.
La fase del monitoreo a largo
plazo incluye la re-medicin
peridica de las parcelas de
monitoreo. El tiempo de medicin
puede variar; algunas parcelas
requerirn al principio ser medidas
anualmente con el fin de evaluar su
variabilidad estacional.
D. Evaluacin de la
Biodiversidad
Vegetal en la
Regin Baja del Ro
Urubamba
La flora del sureste de Per

comprende un gran nmero de
especies. Las irregularidades
topogrficas de la vertiente este de
los Andes crean diferentes microhbitats donde sobreviven plantas y
animales locales. Una alta
diversidad de plantas usualmente se
correlaciona con un gran diversidad
de animales que dependen de ellas.
Muchas especies de animales se
alimentan de las hojas, flores, y
frutos que las plantas producen.
Mientras mayor es la diversidad de
las plantas en una rea, mayor es el
nmero de animales que ellas
pueden soportar.
Hasta la fecha, la mayora de
los estudios florsticos en la regin
del Urubamba se han efectuado en
la parte alta del ro. Como parte de
la evaluacin de la biodiversidad en
la regin baja del ro Urubamba, el
equipo de investigadores muestre
la vegetacin cerca de los pozos de
San Martn-3 y Cashiriari-2. Se
condujeron evaluaciones cercanas a
los pozos localizadas como a un da
xlvi
de camino de los campamentos. La

evaluacin incluy muestreo de
rboles, epfitas, lianas, hierbas, y
helechos que se encontraron en
condicin frtil (i.e. con flores
frutos). Los rboles, lianas, y
epfitas fueron muestreados
subiendo al rbol o muestreandolos
con tijeras telescpicas.
Establecimiento de Parcelas
de Biodiversidad de una
Hectrea
Para iniciar la evaluacin y
monitoreo de la vegetacin, se
establecieron parcelas multi-escala y
parcelas permanentes de 1 ha en San
Martn-3 y Cashiriari-2. Las parcelas
#1 y #2 se establecieron en San
Martn-3, la parcela #3 se estableci
en Cashiriari-2 (Fig. 4). A
continuacin se da una breve
descripcin de los protocolos usados
para establecer las parcelas.
Equipos de topografa
delinearon una parcela de 1-ha
(10,000 m 2 ) y la subdividieron en 25
cuadrantes, cada uno de 400 m 2,
tomando medicions topogrficas
para producir un mapa detallado de
la parcela. Los lmites de la parcela
se demarcan con cuerdas, las cuales
hacen que la orientacin dentro de
la parcela sea ms fcil. Las
coordenadas geogrficas se
establecen con la ayuda de un
geoposicionador. Se requiere una
supervisin estrecha para asegurarse
que cuando se establecen las
parcelas, la perturbacin de la
vegetacin sea mnima.
Despus que los cuadrados son
delineados, el equipo de campo
ubica, mide, marca y mapea en cada
cuadrado cada uno de los rboles
que tengan al menos 4 cm de
dimetro a la altura del pecho (dap).
Se usa cinta diametromtrica para
medir el dap, evitando cualquier
protrusin del tronco. Cuando se

encuentran rams mltiples, se
miden los dimetros
individualmente. Los rboles se
marcan con una etiqueta de
aluminio la cual debe mirar hacia la
lnea de base de la parcela, y se
clavan al rbol por encima del
punto de medicin, por lo que los
clavos sirven como gua general
para mediciones futuras.
Los rboles se marcan con un
nmero nico que consiste de una
secuencia de dgitos dobles. Usando
(01-24-09) como ejemplo, los
primeros dos nmeros (01) designan
la zona donde est localizada la
parcela. Los segundos dgitos (24)
identifican el cuadrado. Los ltimos
dos dgitos (09) numeran al rbol

dentro del cuadrado. Por tanto,
ningun otro rbol recibe el mismo
nmero. En cada cuadrado, los
nmeros de rboles comienzan con
el nmero uno y continan hasta
que el ltimo rbol es marcado.
Un equipo de tres personas
mapean la posicin de los rboles
dentro de los cuadrados. Dos
miembros del equipo se paran en las
esquinas del cuadrado, mientras el
otro localiza los rboles. Con
medidores electrnicos se mide la
distancia de cada rbol a las dos
esquinas ms cercanas. Los valores
registrados de A y B son usados ms
tarde, junto con el dimetro, para
calcular la posicin exacta del rbol.
Resumen
Ejecutivo
Figura 4. Diagrama muestrando las tres primeras parcelas del SI/MAB para la evaluacin y
monitoreo de la biodiversidad.
xlvii
Resumen
Ejecutivo
Con la amplia
experiencia
de los
botnicos, y
con la base
de datos
existente, la
literatura, y la
coleccin de
fotografas,
las especies
ms comunes
fueron
determinadas.
xlviii
Los datos de cada cuadro se

registran en hojas de datos
diferentes en las que se registran la
posicin, dap, y altura, as como
notas pertinentes de los rboles del
cuadrado. Despus que el equipo de
mapeo completa la toma da datos,
los botnicos registran la identidad
de los rboles en las hojas, usando
normalmente cdigos de seis letras
para las diversas especies mapeadas
dentro de la parcela. La informacin
completa de cada hoja se ingresa
luego en BioMon (un programa de
computadora desarrollado por SI/
MAB para manejo de datos) por
equipos. BioMon genera un mapa de
cada cuadrado, el cual es usado por
personal de campo para verificar en
el sitio la ubicacin correcta de los
rboles medidos.
Resultados del Muestreo de
Vegetacin en las Parcelas
Permanentes de
Biodiversidad
De las muestras tomadas en el
rea de estudio, algunas exhiben
diferencias geogrficas y
morfolgicas dentro de una misma
especie. La flora de la regin del
bajo Urubamba est compuesta de
una mezcla de especies que se
encuentran comnmente en otras
reas (e.g., especies tpicas de
bosques secos que se encuentran
desde 700 m en Kiteni a 1100 m en
Potrero, cerca de Quillabamba,
Departamento del Cusco, Per).
Menos del 10% de las especies de
rboles encontradas en el rea
fueron deciduas (rboles que dejan
caer sus hojas en forma estacional).
Varias de las especies muestreadas
fueron clasificadas slo como
morfoespecies (individuos que
comparten los mismos caracteres
morfolgicos son agrupados en la
misma morfoespecie). Varias
especies del gnero Inga spp.

(Fabaceae) fueron muy difciles de
clasificar. Sin embargo, con la
amplia experiencia de los botnicos,
la base de datos existente, la
literatura, y la coleccin de
fotografas, las especies ms
comunes fueron determinadas. No
todas las muestras han sido
determinadas hasta la fecha.
Algunas especies parecen ser
nuevas para la ciencia, incluyendo
una de la familia Acanthaceae y dos
helechos terrestres. Otros
resultados de inters son una
posible nueva forma de color de
Episcia fimbriata y una especie de
Phinaea, relativamnte rara en el
Per. En la familia Heliconiaceae,
dos de las especies de Heliconia
muestreadas parecen ocurrir
raramente en el Per.
Debido a que las parcelas de
biodiversidad en San Martn-3 y
Cashiriari-2 tienen diferentes
caractersticas topogrficas, uno
podra esperar diferentes tipos de
vegetacin. Las parcelas #1 y #2 en
San Martn estn ubicadas en
terreno escabroso con muchas
colinas y valles. En los bordes de
estos valles, se encontraron
frecuentemente macizos densos de
bamb. De hecho, el bamb
constituy hasta el 59% de todos
los individuos en la parcela (cuando
se consideran a rams de bamb
como individuos). Su rol en la
determinacin de la estructura del
bosque parece ser bastante
importante. Durante el trabajo de
campo, result evidente que las
fuertes lluvias y el viento
derribaban las rams de bamb, y
que stas a su vez tienen un efecto
en las especies de rboles
adyacentes. Con frecuencia se
encontraron rboles sin corona, el
efecto inmediato de esta condicin
en general ms altos que los rboles

en las parcelas de San Martn-3. Un
dimetro mayor resulta en una rea
basal mayor (Fig. 7). Un total de 16
especies ocurrieron en densidades
de ms del 1% (de todos los
individuos exceptuando el bamb)
en las parcelas. La parcela en
Cashiriari-2 mostr mucha mayor
equidad en las densidades entre
especies que cualquiera de las
parcelas en San Martn-3, an
cuando en San Martn-3 solo
hubieron unas cuantas especies
comunes, mientras que el resto
ocurrieron en densidades bajas.
Existen ciertas similitudes entre
las parcelas en el bajo Urubamba y
las parcelas SI/MAB de la Zona
Reservada del Manu, Per. En
particular, el nmero de especies y
sus densidades fueron bastante
comparables. La mayor diferencia
fu en el tamao de los rboles. En
Manu, muchas especies tuvieron un
dap de 0.8 m o mayor. En el bajo
Urubamba, las especies de rboles
fueron ms pequeas con slo unas
cuantas especies raras como Ficus
mathewsii que exhibieron troncos
grandes. Esta diferencia puede estar
relacionada con la presencia del
3500
250
3000
200
150
100
50
2500
2000
1500
1000
500
0
0
P lot 1
(S M -3)
P lot 2
(S M -3)
S ite
P lot 3
(CA-2)
Resumen
Ejecutivo
40
Total Basal Area
300
# of Individuals
# of Species
es una reduccin en los productos

fotosintticos, reduciendo as el
crecimiento del rbol. Adems, fue
evidente que varios individuos
daados por el bamb murieron
como resultado de esto. La tercera
parcela de biodiversidad (#3) en
Cashiriari-2 se localiza en un
bosque bajo tpico con pocos filos o
colinas. La flora de este sitio fue un
poco diferente a la de San Martn-3.
En particular, la frecuencia de las
especies vegetales fue diferente
(i.e., algunas especies encontradas
comnmente en San Martn-3
fueron bastante raras en este lugar).
No h aba bamb, y especies como
Grias peruviana y Calatola venezuelana
fueron bastante comunes.
Los nmeros de especies de
rboles fueron diferentes en cada
parcela permanente (Fig. 5). Las
parcelas #1 y #3 tuvieron un
nmero similar de especies (258 y
271 respectivamente) mientras que
la parcela #2 tuvo una cuenta
menor - 224 especies. El nmero de
individuos en las parcelas fue
marcadamente diferente (Fig. 6). La
parcela #3 tuvo un nmero menor
de individuos, pero muchos de stos
tenan grandes dimetros y fueron
35
30
25
20
15
10
5
0
Plo t 1
(SM -3)
Plo t 2
(SM -3)
Plo t 3
(C A-2 )
Site
Plo t 1
(SM -3)
Plo t 2
(SM -3)
Plo t 3
(C A-2 )
Site
Figura 5 (izquierda). Nmero de especies de rboles registradas en cada una de las tres parcelas
(SM-3 es el rea de San Martn-3 y CA-2 es la de Cashiriari-2). Figura 6 (centro). Nmero de
individuos registrados en cada una de las parcelas. Las barras de las parcelas 1 y 2 muestran el
total de individuos con y sin bamb. La parcela 3 no tuvo bamb. Figura 7 (derecha). El rea
basal total de cada una de las parcelas.
xlix
Resumen
Ejecutivo
Aproximadamente un
64% de todas
las especies
de animales
descritas
hasta la fecha
pertenecen al
grupo de los
artrpodos.
bamb, ya que ste puede reducir

drsticamente el grosor de especies
de rboles co-existentes.
En las parcelas donde el bamb
estuvo ausente - las parcelas de
Manu y la parcela #3 en Cashiriari2, el grosor de los rboles fu
considerablemente mayor. Otros
efectos del bamb en la distribucin
y crecimiento de las plantas del
bosque se estudiarn en futuras
investigaciones y monitoreos.
E. Evaluacin de la
Biodiversidad de
Artrpodos en la
Regin Baja del Ro
Urubamba
Al recordar la diversidad del

reino animal, la mayora de las
personas tienden a pensar en
mamferos grandes, aves, y reptiles.
Estos constituyen lo que ha sido
llamado fauna macroscpica de los
bosques tropicales. Sin embargo,
stos representan solo una pequea
fraccin de la riqueza biolgica
animal.
Other
19%
Es la menos glamorosa fauna

microscpica quien comprende la
mayor parte de la biodiversidad
animal. Estos organismos son por lo
general pequeos a muy pequeos y
tienen historias de vida muy
secretas.
El elemento principal de la
microfauna es el grupo conocido
como Artrpodos, los cuales
representan el mayor componente
de la diversida biolgica en el
planeta. Aproximadamente un 64%
de todas las especies de animales
descritas hasta la fecha pertenecen
a ste grupo (Fig. 8), una
proporcin que contina
incrementandose conforme ms
estudios se realizan en las regiones
tropicales y se descubren nuevas
especies.
Los artrpodos se caracterzan
por tener cuerpos cubiertos de
paredes endurecidas y apndices
locomotores que estn divididos en
segmentos distintos. Estos animales
incluyen a las mariposas, moscas,
mosquitos, abejas, avispas,
escarabajos, chinches de las plantas,
Vertebrates
3%
Plants
14%
Arthropods
64%
Figura 8. El conocimiento de la biota global.

l
araas, escorpiones, caros,

cangrejos, camarones y langostas,
as como un gran nmero de
organismos menos conocidos. Su
tamao generalmente diminuto no
evita que los artrpodos cumplan
funciones muy importantes en los
ecosistems del mundo. Ellos son
importantes herbvoros,
consumiendo el tejido vivo de
plantas superiores y de esta manera
regulando su crecimiento; como
depredadores en un sentido amplio,
incluyendo a los parsitos que
ayudan a controlar las poblaciones
de animales; como polinizadores,
facilitando la reproduccin de las
plantas con flores; y como
recicladores de nutrientes al estar
involucrados en la descomposicin
de materia orgnica muerta. Los
artrpodos son tambin muy
importantes como vectores de
enfermedades y patgenos para el
hombre, las plantas y los animales.
La mayor parte de las especies
de artrpodos son sensibles a
cambios ambientales, tanto
espaciales como temporales.
Gradientes ambientales tales como
cambios en la disponibilidad de
agua y pequeas diferencias en la
temperatura limitan la distribucin
de estas especies tan pequeas. Los
cambios temporales en la actividad
de los artrpodos ocurren
diariamente, dado que hay
diferentes especies activas durante
el da y la noche. La composicin y
densidad de las comunidades de
artrpodos para un lugar dado
cambian tambin a lo largo del ao,
muchas especies tienen densidades
ms altas durante la estacin
lluviosa, mientras otras son ms
abundantes durante la estacin
seca. La heterogeneidad del habitat
incrementa el nmero de especies a
una escala mayor, y los patrones
estructurales del bosque tienen una

profunda influencia en la
composicin y abundancia de los
artrpodos - se les encuentran desde
muy profundo en el suelo hasta la
cima del dosel de los rboles.
En el rea de estudio se
complet el muestreo y preparacin
de artrpodos siguiendo protocolos
estandarizados. Los estratos del
bosque seleccionados para el
muestreo fueron el suelo del bosque
y la capa de hojarazca
inmediatamente por encima de ste,
la capa del sotobosque, y el primer
estrato de vegetacin. Tambin se
evaluaron habitats especiales
incluyendo troncos cados y hongos.
Las tcnicas se seleccionarn de
modo tal de minimizar la
sobreposicin entre muestreos.
La separacin de las muestras
es una actividad que toma mucho
tiempo y demanda mucha labor
cuando se trata de artrpodos del
bosque tropical. Una separacin e
identificacin preliminar de las
muestras se hizo en el campo. Una
vez que sto se complete en el
laboratorio, la separacin inicial a
nivel de rdenes permitir enfocarse
en taxa seleccionados para un
mayor procesamiento y estudio,
obtenindose al mismo tiempo un
estimado general de la biodiversidad
de artrpodos en el rea de estudio.
El material muestreado ser
depositado en el Museo de Historia
Natural, Universidad Nacional
Mayor de San Marcos, Lima, Per, y
en el National Museum of Natural
Washington D.C., USA.
Resumen
Ejecutivo
Mariposas
Las mariposas diurnas
constituyen uno de los grupos de
animales ms estudiados dentro de
los invertebrados debido a sus alas
li
Resumen
Ejecutivo
lii
brillantemente coloreadas y a sus

actividades diurnas facilmente
observables, como es el alimentarse
de nctar mientras visitan los
jardines. Como con otros insectos,
las mariposas diurnas tienen un
ciclo de vida que puede completarse
en unos cuantos meses. Al
alimentarse de plantas, las orugas
de las mariposas tienen que lidiar
con la composicin qumica de los
compuestos secundarios que las
plantas producen para reducir el
ataque por herbvoros. As, los
estudios de las orugas de mariposas
han enriquecido el entendimiento
de las interacciones planta-animal,
las asociaciones simbinticas, y la
ecologa y comportamiento de
parasitoides y depredadores. Las
mariposas son altamente
dependientes de las condiciones
microclimticas de sus habitats, y
ellas juegan roles importantes en el
ecosistema debido a que sirven
como polinizadores, se alimentan de
plantas (i.e., herbvoros), y sirven
como presa para cientos de especies
de avispas y moscas parasitoides.
En San Martn-3 y Cashiriari-2, los
objetivos principales de estudiar las
mariposas diurnas fueron el
documentar las especies presentes
en estos sitios y proveer de
informacin bsica acerca de la
historia natural de algunas especies.
Este estudio es de inters particular
ya que la mayor diversidad de
mariposas diurnas en el mundo se
ha documentado en las parte altas
del ro Amazonas en Colombia,
Ecuador, Per, y Brasil. Conforme
otros lugares en reas tropicales
sean estudiados, es muy posible que
se encuentren especies nuevas de
este diverso grupo.
En los sitios de los pozos,
muestreamos 176 especies de
mariposas diurnas, incluyendo ms
del 15% de las especies que

esperamos encontrar. Para tener un
muestreo completo de la diversidad
de mariposas del rea y para poder
detectar fluctuaciones en los
patrones naturales, se deben
estudiar las mariposas durante
varias estaciones y a lo largo de
varios aos.
Las mariposas nocturnas (i.e.
polillas) han sido menos estudiadas
que las mariposas diurnas, an
cuando sabemos que son ms
diversas. No hay muestras
significativas de mariposas
nocturnas para la regin baja del ro
Urubamba. Sin embargo, los
investigadores registrron 87 de las
103 especies de la subfamilia
Ctenuchinae reportadas para
Tambopata, en el sureste del Per,
en apenas 18 das de trabajo de
campo, indicando que la regin baja
del ro Urubamba podra ser una de
las regiones ms ricas en especies
para el Per de mariposas
nocturnas.
Hormigas
Las hormigas son
particularmente adecuadas para
programs de inventario y
monitoreo. Son bioindicadores
ideales debido a que son muy
diversas (aproximadamente 9500
especies descritas en el mundo), se
encuentran en abundancia en casi
todos los habitats del globo. Las
hormigas son numericamente
dominantes en estudios del dosel
del bosque tropical, comprendiendo
entre el 19% y el 50% de todos los
individuos de artrpodos. En un
estudio en la amazona brasilea se
encontr que las hormigas
contribuyeron con el 80% de la
biomasa de insectos y ms del 30%
de la biomasa considerando a todos
los animales. Estos organismos son
muy importantes ecologicamente

debido a que funcionan a muchos
niveles en el ecosistema - como
depredadores, como presas, y como
detritvoros. Las hormigas tambin
tienen asociaciones diversas con
plantas, ya que sirven como
dispersadores de semillas, como
consumidores de semillas, y como
herbvoros.
Dado a que las hormigas son
comunes y aparentes en casi todos
los habitats, ellas son facilmente
muestreadas. Se pueden usar varias
tcnicas, dependiendo del propsito
del proyecto, del habitat y de los
recursos disponibles. Muchas
especies de hormigas son muy
dependientes del microclima y de la
estructura del habitat, por lo que
responden rapidamente a cambios
ambientales. Dado que son insectos
sociales, las hormigas viven en
colonias que van desde unas
cuantas hormigas por colonia hasta
millones; todas las obreras son
estriles y sirven a una varias
reinas. La estructura social de las
colonias de hormigas permite el
muestreo de varias hormigas
obreras sin afectar
significativamente la unidad
reproductiva de la colonia.
Los investigadores estudiaron
30 cuadrantes en San Martn-3. Los
mtodos de muestreo usados fueron
cuantitativos y pueden ser repetidos
a traves del tiempo, lo cual los hace
apropiados para el monitoreo.
Futuros muestreos de hormigas en
los mismos sitios usando esos
mtodos pueden ayudar a detectar
y monitorear cualquier cambio en la
comunidad de hormigas de las
parcelas de biodiversidad y de las
reas circundantes.
Nidos de 55 especies fueron
muestreados, con un promedio de
2.2 especies por metro cuadrado
(m 2). Unas cuantas especies fueron

comunes, pero 27 (49%) de las 55
especies fueron encontradas en uno
solo de los cuadrantes de 1m2. Esto
indica que estas especies no son
comunes en el bosque cerca de San
Martn-3. Muestreos adicionales en
el rea provern mayor informacin.
Se encontr un promedio de 4.3
nidos de hormigas por m 2, lo que
hace un total de 120 nidos en 30
m2. Mientras 13 (43%) de los
cuadros solo tuvieron dos o tres
nidos, un cuadro tuvo nueve nidos y
otro tuvo 11. Con una media de 4.3
nidos por m2, uno puede estimar
43,000 nidos por hectrea. El
nmero de hormigas obreras por
nido vara grandemente entre
especies, algunos nidos tuvieron 10
hormigas, mientras otros tuvieron
varios cientos. Por tanto, aunque es
difcil estimar el nmero de
hormigas por unidad de rea, es
muy posible que su nmero y
biomsa exceda significativamente
el de todas las otras especies
animales. Dada su importancia en el
ecosistema, las hormigas son
ciertamente un elemento esencial en
el funcionamiento del bosque en
San Martn-3.
Los 30 cuadrantes no fueron
suficientes para muestrear todas las
especies de hormigas en el suelo y
la hojarasca. Con cada nuevo
cuadrante muestreado, ms especies
se aadieron, excepto de los
cuadros 27 al 30. Esta nivelacin de
la curva de acumulacin de las
especies no significa que no se
aadirn especies nuevas al
muestrear cuadros adicionales. Ms
an, las especies que fueron
encontradas una sola vez fueron
incrementndose a medida que se
muestrearon ms cuadros,
indicando que muy posiblemente se
muestrearn ms especies raras con
Resumen
Ejecutivo
En un
estudio en la
amazona
brasilea se
encontr que
las hormigas
contribuyeron
con el 80% de
la biomasa de
insectos y con
ms del 30%
de la biomasa
considerando
a todos los
animales.
liii
Resumen
Ejecutivo
Los
investigadores
reportaron
grupos
taxonmicos
supragenricos
nuevos para
el Per: los
Sclerogibbidae
y
Scolebithydae.
liv
cada cuadro adicional. Este tipo de

curva de acumulacin de especies,
que no se nivela, indica que hay
ms especies en el rea que las que
fueron muestreadas y que se
requieren muestreos adicionales.
Abejas y Avispas
Las abejas y las avispas son
extremadamente importantes en el
funcionamiento del ecosistema ya
que son los polinizadores primarios
de cientos de especies de plantas
con flores. Muchas especies de
avispas parasitan otras especies de
insectos, sirviendo como
reguladores de especies de plagas
potenciales, y varias especies de
abejas han evolucionado atributos
sociales que las hacen muy
competitivas. La miel que las abejas
producen es usado por la gente
nativa en reas remotas como una
importante fuente de azcares y
protenas.
Muestreamos 54 especies de
abejas y avispas, sin duda solo una
pequea fraccin de la diversidad
de abejas y avispas que viven en
San Martn-3 y Cashiriari-2. Veinte
de las especies de avispas y cuatro
de las de abejas fueron encontradas
en el estrato bajo cerca del suelo y
en la hojarasca, mientras que 24
especies de avispas y 6 de abejas lo
fueron en los estratos medio y alto.
Los investigadores reportaron
grupos taxonmicos supragenricos
nuevos para el Per: los
Sclerogibbidae y Scolebithydae.
Estos hallazgos aumentan nuestro
entendimiento de la biogeografa del
rea dado que los Scolebithydae
tambin estn reportados para el
Africa tropical y Brasil, y confirma
cuan poco se conoce de la fauna de
hymenoptera en esta rea - uno de
los lugares biolgicamente ms ricos
en el mundo. Tambin se observ
que abejas Trigona spp. estaban

tomando minerales de las tuberas
en las cercanias del pozo en
Cashiriari-2. El hacer un estudio de
sta observacin podra ayudar a
determinar si estas especies seran
tiles como indicadores de niveles
de contaminacin, y estudios
genticos podran darnos una idea
de los efectos que los qumicos
puedan tener en ste grupo.
Liblulas
Las liblulas son organismos
excelentes para estudios de
biodiversidad. Comparados con
otros grupos de insectos tropicales
son relativamente bien conocidos,
facilmente muestreados y
estudiados, y son importantes en
comunidades bnticas las cuales son
usadas en forma rutinaria para
evaluar la calidad del agua superficial. Los adultos son insectos
grandes, coloridos, que vuelan
durante el da, y que pueden ser
generalmente identificados en vuelo
a nivel de gnero. Las larvas son
acuticas y con frecuencia se
encuentran en estudios bnticos.
Tanto los adultos como las larvas
son grandes depredadores cerca de
la cspide de sus respectivas
cadenas trficas. El muestreo es
generalmente no disruptivo debido
a que las poblaciones locales de
estos insectos tienen nmeros altos
de individuos, y slo se toma una
pequea muestra de ellos.
En San Martn-3, muestreamos
las larvas de stas especies
cortando internodos de bamb y
luego examinando su contenido en
una bandeja blanca. Se confirm
que posiblemente dos especies de
libelulas de alas de helicptero son
depredadores en el sistema, y que
un nmero de mosquitos, otras
moscas, y escarabajos tambin
fueron encontrados. La cantidad de

agua disponible en los internodos
del bamb fue estimada
aproximadamente a 10,000 l/ha.
Una tabulacin cruzada de los datos
originales di un listado de 40
morfoespecies en 9 tipos de habitat.
Araas
Las araas se encuentran en
todo el mundo y en todos los
ambientes terrestres. Los miembros
de ste grupo de invertebrados
tienen una gran abilidad para
encontrar presas y para dispersarse.
Los individuos son depredadores
nocturnos, principalmente de otros
invertebrados. Ellas han
desarrollado una variedad de forms
de capturar su alimento: algunas
hacen redes orbiculares, otras hacen
redes planas, otras son cazadoras
ambulantes, y an otras son
expertas en emboscar a sus presas.
Todas juegan un papel muy
importante en controlar las
poblaciones de sus presas. Las
araas son particularmente
numerosas en los tropicos, donde la
vegetacin es abundante, es rica en
especies, y soporta un gran nmero
de sus presas. La complejidad del
habitat y los cambios en la
temperatura y precipitacin (i.e.,
estacionalidad) tienen una gran
influencia en la distribucin y
abundancia de las comunidades de
araas en los ecosistems. Las
prcticas del uso de la tierra
tambin afectan la composicin de
las especies de araas.
Las araas son muy diversas, ya
que es evidente del hecho que se
han reportado alrededor de 34,000
especies en el mundo. Al menos la
mitad de las especies se encuentran
en Amrica Neotropical. Cerca de
1120 gneros de araas se han
reconocido para Amrica tropical, y
al menos la mitad de stos gneros

se encuentran en el Per. En la
primera evaluacin de la diversidad
de araas en el bajo Urubamba se
han identificado un total de 69
morfoespecies de araas.
Caracoles Terrestres
Los moluscos terrestres
incluyen caracoles, babosas y
pseudo-babosas (Veroniclidos). El
nmero estimado de stas especies
en el mundo est entre 30,000 a
35,000, esto es, ms que el total
combinado de especies de anfibios,
reptiles, aves, y mamferos. La
mayor proporcin de especies de
moluscos que se conocen se
encuentra en los tropicos, y sta
podra ser mayor puesto que mucho
de las zonas tropicales no han sido
exploradas.
Los moluscos terrestres de San
Martn-3 se clasificaron en 34
especies - 33 de caracoles
(moluscos con concha) y un
veroniclido (pseudobabosa sin
concha). Treinta y dos especies se
encontraron en la parcela de
biodiversidad #1. La mayora de las
especies estuvieron en estado
reproductivo tal como lo indica la
presencia de huevos. La variacin
en la abundancia de especies fu
alta, desde especies abundantes con
207 individuos hasta cuatro
especies de las cuales slo se
encontraron un individuo. Ms de la
mitad (19) de las especies
estuvieron representadas por menos
de 10 individuos. Ms del 70% de
las especies registradas en San
Martn-3 se encontraron en la
hojarasca y el humus. Solo se
encontraron 3 especies de caracoles
arbreos.
Resumen
Ejecutivo
En la primera
evaluacin de
la diversidad
de araas en
el bajo
Urubamba se
han
identificado
un total de 69
morfoespecies.
lv
Resumen
Ejecutivo
lvi
Escarabajos
Los escarabajos tienen gran
potencial para el monitoreo de
biodiversidad debido a los mltiples
roles que juegan en los ecosistems
y debido a su sensibilidad a cambios
del medio ambiente. Los
escarabajos de estircol, principal
foco de este estudio, estan bien
adaptados para la excavacin, ya
que sus patas son muy fuertes y les
ayudan a mover grandes cantidades
de suelo y comida. Son muy
importantes en el reciclado de
nutrientes, lo que ha generado un
gran inters por estudiarlos. Su
habilidad para reciclar el estircol
animal ha sido usada para mejorar el
cultivo de pastos, aunque se les ha
asociado con la transmisin de
enfermedades, debido a que
mueven los huevos de parsitos que
afectan la salud humana hacia la
superficie del suelo, aumentando as
la transmisin del parsito a los
hospederos. Los escarabajos son un
componente importante de la dieta
de muchas aves insectvoras. Por
ejemplo, las aves consumieron hasta
el 85% de stos animales en la terra
firme de la amazona central y 49%
en el Parque Nacional Manu.
Fluctuaciones en las poblaciones de
escarabajos se han correlacionado
con cambios de comportamiento en
varias especies de mamferos. En el
rea de estudio se muestrearon 47
morfoespecies de escarabajos de
estircol distribudos en 13 gneros
y 3 subfamilias.
F. Evaluacin de la
Biodiversidad de
Anfibios y Reptiles
en la Regin Baja
del Ro Urubamba
Los anfibios y reptiles

constituyen un componente
importante de la fauna de
vertebrados en bosques tropicales.
Tienen gran diversidad y
especializacin ecolgica con
preferencia de ciertos hbitats, y
son relativamente importantes en
muestreos cuantitativos. El objetivo
primario de las dos evaluaciones de
anfibios y reptiles llevados a cabo
en la zona de estudio, fue iniciar el
establecimiento de una informacin
de lnea de base de anfibios y
reptiles que se encontraron en las
cercanias de las dos plataforms de
perforacin. Esta informacin es
necesaria para ser incorporada en el
programa de monitoreo multi-taxa
diseado para detectar cambios en
el ecosistema y en el manejo de los
recursos naturales del rea.
Algunos especmenes de
anfibios y reptiles encontrados
durante las dos evaluaciones fueron
muestreados y preservados como
material de referencia, para
documentar la presencia de
especies, y para empezar a examinar
y clarificar la distribucin y
taxonoma de la herpetofauna en la
regin. En la primera evaluacin, un
total de 504 especmenes fueron
muestreados, documentando ms de
80 especies de anfibios y reptiles en
San Martn-3 y Cashiriari-2. Se
encontraron ms individuos por
especie de anfibios, y ms especies
de diferentes familias y gneros de
reptiles; los reptiles slo
constituyeron el 18% del total de
especmenes muestreados. Adems
Tabla 1. Comparacin de la fauna conocidas para reas tropicales

del Per (El nmero de especies de anfibios encontrados en la regin
baja del ro Urubamba en San Martn-3 y Cashiriari-2 [datos de
Reynolds et al. e Icochea y Mitchell este volumen] estn comparados
con Manu [datos de Morales y McDiarmid 1996] e Iquitos [datos de
Rodrguez y Duellman 1994] )
Familia
Bufonidae
Dendrobatidae
Centrolenidae
Hylidae
Leptodactylidae
Microhylidae
Pipidae
Ranidae
Urubamba
3
4
11
30
1
de los especmenes muestreados, los

investigadores registraron cuatro
grupos adicionales en San Martn-3
que no fueron muestreados.
Tambin se captur y se solt un
caimn en una quebrada al este del
campamento y se identific
positivamente el canto de una rana
arborcola; la vocalizacin
constituy el nico registro para esa
especie. Los botnicos reportaron el
encuentro de una tortuga mientras
mapeaban los rboles en la parcela
de biodiversidad #1. Varias
especies de lagartijas fueron
capturadas con las trampas de
goma.
Varias de las especies de
anfibios reportadas en este estudio
han sido encontradas en otras
tierras bajas tropicales del sureste
del Per91% en Cosha Cashu y
78% en Pakitza. La Tabla 1
compara la fauna de anfibios del
rea de estudio y el rea cercana de
Parque Nacional Manu y la de
Iquitos. La fauna de reptiles
tambin se sobrepone con sas de
Cosha Cashu y Pakitza72% y
67% respectivamente.
Comparaciones equitativas de los
Manu
3
6
1
27
26
4
Resumen
Ejecutivo
Iquitos
7
8
50
37
7
2
1
resultados de San Martn-3 y

Cashiriari-2 no son posibles por
ahora ya que el esfuerzo de
muestreo fue ms intensivo en San
Martn-3.
Los investigadores muestrearon
tambin 84 especmenes de anfibios
y reptiles para determinar la
diversidad gentica de las especies
del rea. Fueron tomadas cincuenta
muestras de sangre (para la prueba
de microncleos) de 21 especies,
3000 ncleos por individuo estn
siendo procesados, y 51 muestras de
18 especies fueron tomadas para
efectuar las pruebas de alteracin
cromosmicas.
G. Evaluacin de la
Biodiversidad de
Aves en la Regin
Baja del Ro
Urubamba
Las aves son probablemente las

ms ampliamente estudiadas dentro
del grupo de los vertebrados,
principalmente por su gran
diversidad, la influencia tan considerable que tienen en el ambiente, y
lvii
Resumen
Ejecutivo
Al parecer,
la tasa de
acumulacin
de especies
en San
Martn-3
excede la de
cualquier
otro hbitat
muestreado,
incluyendo
los bosques
del Parque
Nacional
del Manu.
lviii
el lugar que ocupan en la historia

humana y cultural. En todas las
comunidades, incluyendo las
dominadas por el hombre, las aves
son importantes consumidores,
depredadores y presa de otros
organismos. Su rol en el control de
las poblaciones de insectos,
incluyendo plagas agrcolas, es bien
conocida. En muchos ambientes,
las aves son los principales
dispersores de semillas de plantas,
generalmente esparcindolas
despus de haberse alimentado de
una variedad de frutas y semillas.
Varios grupos importantes de aves
han evolucionado para alimentarse
de nctar y polen, y como resultado
son polinizadores exclusivos de
plantas. En su rol como dispersores
de semillas y polinizadores, las aves
sirven como agentes de
movilizacin que vinculan varios
componentes de la cadena
alimenticia. En esta capacidad, y
como depredadores y consumidores,
algunas aves son especies claves
que determinan grandemente la
estructura y funcin de las
comunidades naturales. Las aves
tambin facilitan la formacin de
microhbitats para otros
organismos, incluyendo otras aves.
Adems, el nido de las aves sirve
como hbitat para una variedad
muy amplia de organismos, y las
aves misms son hospederos de una
gran variedad de parsitos y
comensales.
Las comunidades de aves de las
tierras bajas del Per estn entre las
ms diversas del mundo. Sin embargo, relativamente pocas reas
dentro de esta vasta regin han sido
evaluadas por ornitlogos. En el
sureste del Per, existe
relativamente buena informacin de
varios lugares en la regin de la
provincia del Madre de Dios,
incluyendo El Parque Nacional del

Manu, y las reservas de Tambopata,
y Cusco Amaznico. En contraste,
pocos estudios se han llevado a
cabo en las tierras bajas del
Urubamba de la Provincia del
Cusco. En cambio, la parte alta del
ro Urubamba ha sido bastante
estudiada por ornitlogos.
En este estudio se muestrearon
un total de 198 especies en San
Martn-3 y Cashiriari-2, 65 de las
cuales fueron observadas en ambos
lugares, 100 en San Martn-3, y 33
en Cashiriari-2. El bajo nmero de
especies observadas en Cashiriari-2
es probablemente el resultado del
limitado tiempo de muestreo en ese
sitio. Examinando el nmero de
especies registradas dentro de un
mismo hbitat, se observa, que los
dos lugares muestreados del bajo
Urubamba son bastante similares a
otros en el sureste del Per. Datos
obtenidos con redes de neblina en
muestran que en San Martn-3,
ochenta y siete especies fueron
determinadas de un total de 440
capturas (representando 360
individuos). En Cashiriari-2,
sesenta y dos epecies fueron
determinadas de un total de 248
capturas (representando 224
individuos).
Como en otros lugares
tropicales, slo se encontraron en
las muestras unas pocas especies
comunes y muchas raras. En San
Martn-3, 83% de las especies
capturadas representan el 2% o
menos de las capturas, mientras que
en Cashiriari-2 fue del 77%. Al
parecer, la tasa de acumulacin de
especies en San Martn-3 excede la
de cualquier otro hbitat
muestreado, incluyendo los bosques
de Parque Nacional del Manu (Fig.
9).
Hubo poca evidencia de

actividad reproductiva en los dos
lugares. En las muestras de las redes
de neblina, ninguna ave fue
observada con polluelos. En
contraste, muchos de los individuos
se encontraron mudando las plums
en los dos sitios. En San Martn-3,
64% de todas las aves examinadas
durante los muestreos con redes
(316) mostraron signos de muda. El
porcentaje en Cashiriari, 43%, fue
ms bajo.
Aves de caza de tamao mayor
como las gallinetas, ponchas, pavas
y trompeteros son las primeras
especies en desaparecer como
respuesta a la actividad humana,
principalmente como resultado de la
sobrecacera. En San Martn-3,
grupos pequeos de pava uquira
fueron observados regularmente, y
la pava rajadora fue observada en
tres oportunidades. El paujil culo
colorado y la grulla fueron

reportados por otros investigadores.
Tres especies de gallinetas, grande,
cinereous, y con capa negra, fueron
tambin observadas. Las gallinetas
tienen por lo general un
comportamiento bastante discreto y
son principalmente reportadas por
el canto. Las actividades de canto
son bastante limitadas durante la
ltima parte del perodo de lluvias,
y es posible que evaluaciones
durante otras estaciones del ao
registren especies adicionales de
gallinetas. La frecuencia de
encuentros de esas especies de caza
ms su relativa calma cuando se
observaron, indica que la presin de
cacera en el rea inmediata a San
Martn-3 es bastante reducida.
Loros grandes, guacamayas, y
tucanes tienden a desaparecer
relativamente rpido debido a la
actividad humana; ellas son cazadas
Resumen
Ejecutivo
100
90
# of Species
80
70
60
50
40
30
20
10
San Martin
Cashiriari
Manu
Panama
Brazil
Costa Rica
0
0
61
87
100
109
171
200
248
283
300
323
359
400
440
550
Captures
Figura 9. Curvas de acumulacin de especies de aves registradas en San

Martn-3 y Cashiriari-2 (diamantes = San Martn-3, cuadrados =
Cashiriari-2). La lnea con tringulos indica la curva de acumulacin de
especies para el Parque Nacional del Manu, construida de Karr et al. 1990.
La lnea con x representa Panam, los asteriscos representan datos de
Brasil, y los crculos a Costa Rica.
lix
20
18
Resumen
Ejecutivo
16
Todo esfuerzo
debe hacerse
para que esta
avifauna
excepcional
no sufra
degradacin
como
resultado del
desarrollo del
gas natural en
el rea.
lx
# of Species
14
12
10
8
6
4
San Martin - 3
C ashiriari - 2
0
0
10
11
12
13
Sampling Days
Figura 10. Curvas de acumulacin de especies para los mamferos

pequeos no voladores en San Martn-3 y Cashiriari-2.
no solo como alimento sino tambin
para su venta. Como se mencion
anteriormente, esas especies fueron
raras y poco comunes en San
Martn-3. Sin embargo es posible
que las bajas densidades en este
lugar sean resultado de la estructura
del bosque ms que de la presin de
caza, la cual es casi inexistente.
La avifauna de San Martn-3 y
Cashiriari-2 es bastante diversa, y la
riqueza de especies en esas reas
puede ser igual o sobrepasar la de
hbitats equivalentes al Parque
Nacional Manu, uno de los lugares
ms ricos en diversidad de especies
en el mundo. Por ahora, la
comunidad de aves parece
encontrarse en una condicion
pristina, con poca evidencia hasta la
fecha de que la actividad humana
haya tenido un impacto
significativo. Todo esfuerzo debe
hacerse para que esta avifauna
excepcional no sufra degradacin
como resultado del desarrollo del
gas natural en el rea.
H. Evaluacin de la
Biodiversidad de
Mamferos en la
Regin Baja del Ro
Urubamba
Mamferos Pequeos no
Voladores
Los mamferos pequeos no
voladores, se estima constituyen
entre el 15% y 25% de la fauna de
mamferos del bosque hmedo
tropical. Estos animales abundantes
y ampliamente distribuidos ocupan
una variedad de hbitatsarbreos,
terrestres, y semiacuticos
dependiendo de sus diferentes
adaptaciones morfolgicas, y as son
candidatos excelentes para estudios
de evaluacin y monitoreo.
En San Martn-3 y Cashiriari-2,
este grupo fue muestreado
alrededor de las trochas, quebradas,
y cerca de la plataforma de
perforacin usando trampas con
cebos. Seis especies de marsupiales
y 13 especies de pequeos roedores
fueron encontrados (Fig. 10). A
pesar del corto tiempo invertido
muestreando en esos dos lugares

(15 das en cada sitio), los
resultados fueron suficientes para
permitir una comparacin con los
datos de Manu y del Cusco
Amaznico. En San Martn-3, la
tasa de muestreo fue del 5%; en
Cashiriari-2 del 2%. La tasa de
muestreo para Manu y Cusco
Amaznico fue del 1% y 5%
respectivamente.
El mayor nmero de especies
en San Martn-3 que en Cashiriari-2
puede deberse a la presencia de
bamb en San Martn-3. Adems, la
estructura del bosque en San
Martn-3 puede proporcionar un
mayor nmero de lugares para
refugio, fuentes de alimento, y
forms de reducir la depredacin.
Murcilagos
Los murcilagos son el nico
grupo de mamferos voladores.
Ellos constituyen el segundo orden
ms grande de mamferos con 177
gneros y 925 especies. Se
distribuyen a travs de las regiones
tropicales y templadas del mundo,
donde ocupan una gran variedad de
nichos ecolgicos. Los murcilagos

son ms activos durante la noche, y
buscan refugio durante el da en
cuevas, cavidades en los rboles, y
otros lugares. En las ciudades, las
colonias pueden ser encontradas en
edificios, debajo de puentes, y en
tneles. Los murcilagos son
conocidos porque se orientan por
ecolocalizacion (e.g., vuelan y
encuentran su comida a travs de
un sistema de sonidos de alta
frecuencia y sus ecos). Ellos pueden
capturar insectos mientras vuelan
utilizando la ecolocalizacin, pero
tambin se alimentan de la sangre
de mamferos, peces, pequeos
vertebrados, y productos vegetales
como polen, nctar, y frutas.
Los murcilagos juegan un
papel muy importante en las
funciones del ecosistema. Ellos
consumen grandes cantidades de
insectos, ayudan en la polinizacin
de cientos de especies de plantas
con flores las cuales han
evolucionado la apertura de sus
flores durante la noche, y sirven
como dispersores de semillas
alimentndose de frutas. Los
Resumen
Ejecutivo
50
45
# of Species
40
35
30
25
20
15
10
San Martin - 3
Cashiriari - 2
0
0
10
11
12
13
14
Sampling Days
Figura 11. Curva de acumulacin de especies de murcilagos muestreados

en San Martn-3 y Cashiriari-2.
lxi
Resumen
Ejecutivo
Huellas,
lugares de
descanso y
otras seales
de otorongo,
pecary, y
shacavaca
fueron
registradas
muy cerca
(100 m) de las
plataformas
de
perforacin.
agricultores utilizan el guano rico en

nutrientes (heces) producido por los
murcilagos para fertilizar los suelos
agrcolas.
Un total de 51 especies de 31
gneros y cinco familias de
murcilagos (379 individuos) fueron
muestreados en San Martn-3 y
Cashiriari-2. Treinta especies fueron
localizadas en San Martn-3, y 43 en
Cashiriari-2. De sas, 22 fueron
encontradas en ambas localidades, 8
fueron encontradas solo en San
Martn-3, y 21 slo en Cashiriari-2.
La curva de acumulacin de
especies para los datos obtenidos en
cada sitio se muestra en la figura
11, lo que sugiere patrones
diferentes de diversidad para cada
localidad. Como una comparacin,
55 especies de murcilagos fueron
registrados en un perodo de cinco
aos en el Parque Nacional del
Manu, Per, y 57 en la Guayana
Francesa despus de diez aos de
muestreos.
Mamferos Medianos y
Grandes
Debido a la creciente
preocupacin acerca del manejo de
los sistems biolgicos
posiblemente impactados por el
desarrollo de gas y petroleo,
evaluaciones de la biodiversidad
han sido sugeridas como mtodos
para el desarrollo de lneas de base
para esos sistems. Procesos de
monitoreo para el seguimiento a
partir de la informacin de lnea de
base estn siendo desarrollados y
probados. El objetivo de esta
evaluacin fue llevar a cabo un
reconocimiento preliminar de los
mamferos de tamao mediano y
grande en preparacin para una fase
de monitoreo en los alrededores de
las plataforms de perforacin del
bajo Urubamba.
lxii
Una lista de especies que se

esperara ocurran en los dos lugares
fue compilada con base a
descripciones y mapas de
distribucin. La lista de especies
reflej la suma de cuatro fuentes,
los cuales fueron confirmados como
presentes o no por la evidencia
acumulada en cada sitio. Cincuenta
y ocho especies de mamferos
fueron considerados potencialmente
presentes en los dos lugares de
perforacin. En San Martn-3 se
registraron 25 especies, y 26
especies en Cashiriari-2 para un
total de 33 especies no duplicadas.
El nmero total de especies
confirmadas como presentes fue
57% de la lista potencial.
Venticuatro 41% de las especies
posiblesfueron no confirmadas
como presentes, pero pueden estar
presentes, y quizs ser comunes.
Algunas especies de monos,
habitantes del dosel del bosque, con
territorios grandes y/o poca
movilidad, y animales nocturnos
requieren un esfuerzo mayor para su
evaluacin. Todas las especies
listadas tienen una probabilidad
razonable de estar presentes y
ocasionalmente moverse a travs
del rea de estudio. Ms an, las
pacas, gatos pequeos, y especies
de armadillo tienen huellas y
tamaos muy similares, y son
bastante difciles de distinguir sin la
coleccin de especmenes. Esas
especies son generalmente
agrupadas en el mismo bloque.
Encontramos ms dificultades
en distinguir las huellas de los gatos
pequeos, particularmente los
tigrillos, ocelotes y jaguarundis.
Durante este estudio, se tomaron
medidas de cuatro grupos de
huellas, todas identificadas como
huellas de ocelote. Estas fueron
encontradas dentro de un perodo
de dos horas y sobre tres km, y

fueron probablemente de individuos
diferentes. Todas las huellas fueron
frescas en arena o arcilla y de una
calidad excelente. Tambin fueron
observadas huellas de gato con
diferente configuracin en la
almohadilla del taln y que variaron
en tamao entre 2.5 y 5.0 cm. Se
asumi que por lo menos dos
especie de gatos pequeos
estuvieron presentes, probablemente, el margay y el ocelote; sin
embargo son necesarios ms
estudios para verificar su presencia.
Observaciones generales
indicaron que las especies esperadas
en los lugares de estudio estuvieron
presentes, y que los nmeros y
densidades de esas poblaciones se
encontraban en los rangos
esperados. La actividad de los
mamferos grandes en los
alrededores del rea de perforacin
de San Martn-3 y Cashiriari-2
continua y cercana; el ruido y las
operaciones no pareci tener ningn
impacto severo a excepcin de la
plataforma. Huellas de venado de
selva y del conejo brasileiro
estuvieron presentes en ambos
lugares. Huellas, lugares de
descanso y otras seales de
otorongo, pecary, y shacavaca
fueron registradas muy cerca (100
m) de las plataformas de
perforacin. Las especies de felinos
(ocelotes, pums y otorongos) que
con frecuencia son los primeros en
escaparse del desarrollo humano
tambin estuvieron activos dentro
de 100 m de la plataforma de San
Martn-3. Se encontr un rea
importante usada como letrina por
otorongos y pums a
aproximadamente 2.5 km de la
plataforma de perforacin de
Cashiriari-2, donde los motores
podan escucharse claramente. Las
observaciones y signos de
mamferos grandes no parecieron
ser menos numerosas cerca de las
plataforms que a 2.5 km. Las
observaciones y signos de los dos
lugares fueron suficientes para
desarrollar un estimado de
densidades para 28 especies. San
Martn-3 tuvo claramente un ndice
de abundancia mayor (1643) para la
mayora de los animales que
Cashiriari-2 (1028).
Parsitos de Mamferos
De las muestras de
murcilagos, 207 especmenes
fueron revisados para el muestreo
de ectoparsitos. Se encontr una
tasa de infeccin de 68.5 %.
Aproximadamente 93% de los
mamferos pequeos analizados
estuvieron tambin infectados.
Ciento cincuenta y cinco
especmenes de ectoparsitos
fueron muestreados de pequeos
mamferos en los dos pozos. Esos
especmenes fueron clasificados en
8 familias, y 22 gneros. Cuarenta y
cinco especies se reportan por
primera vez en el bajo Urubamba.
Resumen
Ejecutivo
Ciento
cincuenta y
cinco
especmenes
de ectoparsitos
fueron
muestreados
de pequeos
mamferos en
los dos
pozos.
I. Plan de Monitoreo
de Biodiversidad
para Los Grupos
Taxonmicos
Estudiados en la
Regin Baja del Ro
Urubamba
Como se describe en este
documento, existe la necesidad de
establecer un programa de
monitoreo cientfico en los sitios de
perforacin de SPDP y a lo largo de
la ruta propuesta para el
gaseoducto. El programa de
monitoreo ayudar a evaluar
lxiii
Resumen
Ejecutivo
patrones de cambio en los grupos

taxonmicos que se estn
estudiando en la regin del bajo
Urubamba (vegetacin,
invertebrados, anfibios y reptiles,
aves, y mamferos). Esto permitir
asistir en la determinacin de
cambios en la composicin y
funcionamiento del bosque que
pueden ocurrir como resultado de la
construccin y operacin de los
pozos de perforacin. La siguiente
discusin se basa en las necesidades
y usos del monitoreo en cada uno
de los cinco grupos y proporciona
recomendaciones especficas para
incluirlas en el diseo del programa
de monitoreo a largo plazo.
Vegetacin
Los sistems biolgicos, como
son los ecosistems forestales,
tienen interacciones biticas y
abiticas extremadamente
complejas. Mientras todos los
componentes de un sistema
complejo son esenciales, las plantas
comprenden quiz el componente
estructural y funcional ms
importante de un ecosistema forestal. Las plantas del bosque como
productores primarios juegan un
papel crucial para el monitoreo de
todos los grupos taxonmicos.
Algunas rboles gigantes, como
los higos, se han considerado como
piezas claves del bosque porque
desempean un papel muy
importante para la sobrevivencia de
muchas especies, por ejemplo las
aves y los murcilagos dependen de
los rboles de higo durante las
estaciones cuando otras frutas no
son abundantes. Sin estas especies
claves, otras poblaciones de muchas
especies seran afectadas, y como
resultado, esto podra afectar otras
poblaciones de plantas que
dependen de esos animales para la
lxiv
dispersin de semillas y la
polinizacin de flores.
Las implicaciones cientficas de
la biodiversidad del bosque y el
papel crtico que las plantas del
bosque juegan con respecto a la
poblacin humana no puede
subestimarse. Existen muchos
beneficios econmicos, mdicos y
agrcolas de las plantas del bosque.
Adems del valor que tienen las
plantas como posibles remedios
para curar enfermedades que
afectan a los humanos, las plantas
tropicales proporcionan un
reservorio gentico natural que
ayuda a mantener muchas especies
agrcolas domesticadas.
Hasta el momento, los estudios
han proporcionado informacin
preliminar de la distribucin de
especies de rboles y una
descripcin de los hbitats de dos
sitios. El monitoreo de las parcelas
permanentes permitir un mejor
entendimiento de los impactos
causados por las perturbaciones
naturales y humanas en la
composicin de especies y en las
estructuras de las comunidades. Los
datos del programa de monitoreo
pueden ser tambin utilizados para
predecir posibles cambios en el
futuro, con base al conocimiento
actual de los procesos, estructuras y
funciones de los ecosistems. El
programa de monitoreo incluye los
siguientes objetivos:
* Continuar la toma de datos y
de informacin para la lnea de base
en los sitios de perforacin y en
general en la zona del bloque #75
donde SPDP se encuentra operando.
Este objetivo, incluye la
continuacin de las evaluaciones e
inventarios, facilitando la
identificacin de diferentes tipos de
bosques, un estimado de la
diversidad de especies, y
descripciones de las caractersticas

de la vegetacin. La cantidad de
informacin sobre la dinmica de
las especies se incrementa a medida
que ms lugares de muestreo son
estudiados.
* Lograr un mejor
entendimiento de los tipos de
hbitats, tipos de vegetacin y
distribucin espacial de las especies
en los lugares de perforacin, a lo
largo del gaseoducto, y en el rea de
influencia del proyecto de
desarrollo de gas natural. Es
importante que las implicaciones de
la actividad humana en los patrones
de vegetacin y la regeneracin del
bosque sean estudiadas
cuidadosamente. Cambios en la
estructura del bosque causados por
la actividad humana en los sitios de
perforacin y a lo largo del
gaseoducto inducirn la
regeneracin de la vegetacin en las
reas perturbadas. Las especies que
dominan esos lugares, sern
especies secundarias pioneras que
formarn un bosque bastante
diferente del bosque circundante no
perturbado. La influencia de esta
sucesion tendr impactos
significativos en los animales que
actualmente habitan el rea. Por lo
que recomendamos estudiar y
analizar la vegetacin del rea
boscosa no perturbada y el bosque
en regeneracin, tanto cerca de las
comunidades, como en los sitios de
operacin de SPDP.
* Obtener informacin
cuantitativa de la representatividad
de las parcelas de monitoreo de
biodiversidad con las parcelas
multi-escala de vegetacin, as
como en el rea no muestreada.
Estudios de la vegetacin requieren
datos de numerosos lugares en el
bosque para obtener muestras
representativas. Ya que el bosque de
Camisea es muy extenso y que slo

es posible muestrear algunas
hectreas, es muy importante que
los lugares sean representativos del
rea en general.
Artrpodos
Los artrpodos (insectos,
araas, caros, entre otros) son el
grupo ms diverso de organismos en
los ecosistems terrestres. Clculos
recientes estiman la fauna de
artrpodos entre 10 y 30 millones
de especies. Los artrpodos proveen
informacin en casi todos los micro
y macro hbitats dentro del
ecosistema. Tienen miembros de
muchos tamaos; pueden ser desde
muy especializados hasta muy
generalistas en cuanto a
preferencias dentro del ecosistema.
Poseen grandes habilidades para
dispersarse, interesantes ciclos de
vida, con tiempos cortos o largos de
desarrollo; estn involucrados en
casi todas las funciones dentro de
los ecosistems tales como la
descomposicin del material
orgnico manteniendo la estructura
del suelo y su fertilidad; regulan las
poblaciones de otros organismos
(incluyendo a los artrpodos
mismos, vertebrados y plantas);
responden rpidamente a cambios
en el ambiente; y actan como
conectores esenciales para la
reproduccin de muchas plantas
con flores. La informacin que se
deriva de las comunidades de
artrpodos puede ser usada para
caracterizar adecuadamente casi
todos los aspectos de un
ecosistema.
Un aspecto importante en el
monitoreo y evaluacin de los
cambios en los ecosistems, es la
habilidad de tener referencia
espacial-temporal para los cambios
biticos. Para analizar cambios
Resumen
Ejecutivo
Un aspecto
importante en
el monitoreo y
evaluacin de
los cambios en
los ecosistems, es la
habilidad de
tener
referencia
espacialtemporal para
los cambios
biticos.
lxv
Resumen
Ejecutivo
lxvi
biticos asociados con el clima o

con la perturbacin humana, los
datos biolgicos deben ser
comparables en espacio y tiempo y
puestos dentro del contexto de los
sistems naturales (i.e., los datos de
sistems perturbados deben
compararse con los datos de sitios
no perturbados). Ambos protocolos
de muestreo y los mtodos para
registrar los datos abiticos
asociados con los protocolos deben
ser estandarizados. La visin para
llevar a cabo el monitoreo tambin
debe ser estandarizada para todo el
ecosistema.
La estructura de la vegetacin
clasificada en varios estratos,
permite una visin que puede ser
utilizada para organizar el
monitoreo de los artrpodos. Los
datos obtenidos usando los mtodos
apropiados dentro de esta visin a
varias escalas, puede ser entonces
comparada con las comunidades de
artrpodos a travs de gradientes y
entre varios ecosistems o con
comunidades anlogas en diferentes
ecosistems. El reconocimiento de
que los ecosistems tienen varios
estratos tiene implicaciones tanto
en el diseo del programa de
muestreo como en los protocolos
utilizados para hacerlo. Por ejemplo,
los protocolos de muestreo del
dosel deben ser aplicados
verticalmente en un gradiente de
humedad a travs de varios puntos
de muestreo (i.e., un dosel bajo,
mediano, y alto de plantas
herbceas, arbustos, y rboles), y
horizontalmente a travs de un
gradiente hidrolgico representado
por la transicin de una zona
riberea a puntos elevados de
muestreo. La fauna transiente
(aqulla que est en movimiento
localmente), la ms diversa en la
comunidad de artrpodos, puede
entonces ser caracterizada

utilizando protocolos especficos y
no podr ser confundida con la
fauna de artrpodos endmicos al
suelo, a la hojarasca, al dosel de los
rboles, ni a las comunidades
acuticas. Estos grupos tambin
pueden caracterizarse con
protocolos apropiados asociados
con cada estrato de vegetacin.
Dada la gran diversidad de
artrpodos, no es posible
monitorear todas las especies o
evaluar cambios en algunas especies
sin tener los datos base de
referencia para los ecosistems
naturales apropiados. La adquisicin
de la base de referencia en los
estudios de biodiversidad de sitios
no perturbados es un componente
extremadamente importante dentro
de la estrategia de monitoreo. Los
datos de referencia sirven para
comparar, usualmente con un
tamao de muestra ms pequeo,
otros muestreos para reflejar la
realidad en el ecosistema. La base
de datos de referencia es el primer
paso para el desarrollo del
monitoreo, mientras se adquiere la
base de datos que funcionar como
la herramienta ms poderosa para
analizar los cambios en el
ecosistema. La ejecucin de los
programs de monitoreo se debe
basar en el conocimiento amplio y
bsico de las especies.
Estrategia para el Monitoreo
de Artrpodos
Para el proyecto del bajo
Urubamba, los investigadores han
recomendado que la base de datos
para artrpodos este restringida a
aquellos grupos que sean
taxonmicamente adecuados (i.e.,
su taxonoma se conoce y los
taxnomos estn disponibles para
determinar los especmenes). La
base de referencia para los

artrpodos debe consistir de cuando
menos dos parcelas de
biodiversidad para cada sitio: una
situada en el bosque ms tpico del
rea que vaya a ser afectada por el
pozo de exploracin; la otra, el
control, situada en el mismo tipo de
bosque que el primero, pero en un
rea en donde no vaya a ser
afectada por el pozo o por la
construccin del gaseoducto. Los
programs subsecuentes de
monitoreo debern usar las misms
parcelas de biodiversidad en donde
se obtuvieron los datos base. Cada
base de referencia deber estar
asociada a la toma de datos
abiticos que estructuran el
ecosistema (i.e., clima, suelo,
calidad de agua, entre otros), de tal
forma que se puedan comparar con
los datos de otros expertos de la
comunidad internacional.
La evaluacin de la diversidad
de artrpodos empez siguiendo los
preceptos bsicos descritos en los
prrafos anteriores. Los objetivos
son:
* adquirir la base de datos de
referencia para los artrpodos, al
menos en dos parcelas de
biodiversidad, incluyendo un control y un tratamiento, en cada sitio
de estudio;
* en los aos subsecuentes,
monitorear componentes selectos
de la fauna de artrpodos
representativos de diferentes tallas,
habilidades para dispersarse, niveles
trficos en el suelo, hojarasca, y
dosel que existen a travs de los
gradientes fsicos y qumicos en las
parcelas de biodiversidad;
* cuantificar la taza de cambio
en la biodiversidad de las
comunidades de especies de
artrpodos que puedan ser
asociadas a la extraccin del gas

natural;
* evaluar el impacto del cambio
en la composicin de especies de
artrpodos en las zonas de estudio;
y
* desarrollar un inventario de la
fauna de artrpodos en la regin
usando los datos obtenidos para la
base de referencia y de los
programs de monitoreo.
Se ha logrado un avance
substancial al adquirir los datos
iniciales para la base de datos de
referencia. Durante las siguientes
fases del proyecto, el trabajo
adicional necesario para la toma de
datos de referencia y el desarrollo
del programa de monitoreo se
desarrollar en tres partes
capacitacin de personal,
adquirir los datos de base de
referencia, y monitoreo.
Se espera que la estrategia para
la evaluacin y el monitoreo que se
ha presentado en este plan,
produzca informacin en todos los
estratos del ecosistema que est
alrededor de la zona de estudio. Se
generar informacin sobre la
biodiversidad de las comunidades
de artrpodos que habitan el suelo,
la hojarasca, la vegetacin, y el
dosel. Se estima que ms de 15,000
especies de artrpodos sern
incluidas en el anlisis de
biodiversidad considerando el
costo-beneficio ms efectivo.
Resumen
Ejecutivo
Uno de los
objetivos del
monitoreo es
evaluar el
impacto del
cambio en la
composicin
de especies
de
artrpodos
en las zonas
de estudio.
Anfibios y Reptiles
Los anfibios son un grupo
diverso de animales depredadores
que se encuentran en todos los
continentes y alcanzan su mayor
abundancia en las zonas tropicales.
La mayora posee ciclos de vida
complejos, un estadio larval
acutico, y un estadio adulto
terrestre que es nico entre los
lxvii
Resumen
Ejecutivo
vertebrados. Cada individuo, por lo

tanto, se expone a ambientes
acuticos, terrestres y atmosfricos.
Este tipo de vida, combinado con
una piel hmeda, sensible, que
utilizan para respirar, y la falta de
una gran movilidad, hace que las
poblaciones de anfibios sean muy
susceptibles a cambios en el
ambiente. Estos organismos son
generalmente los primeros en
responder a cambios en el ambiente.
En el grupo de los anfibios, las
perturbaciones a gran escala han
ocasionado la reduccin de
poblaciones y de su rango de
distribucin, as como la extincin
de varias especies. La causas de la

reduccin incluyen la prdida y
degradacin de habitats, la lluvia
cida, la radiacin UV-B, las
enfermedades, la introduccin de
especies exticas, la cosecha por los
humanos, as como las
fluctuaciones naturales de las
poblaciones. Las causas principales
en la reduccin del rango de
distribucin y de poblaciones y la
alteracin en la estructura
poblacional de las comunidades se
deben principalmente a la prdida y
degradacin del hbitat.
Las perturbaciones iniciales que
probablemente ocurrirn como
Figura 12. Un ejemplo de micro-habitats nocturnos de varias especies de ranas en la

regin baja del Urubamba.
lxviii
consecuencia de las actividades en

los pozos de exploracin que
afectarn a los anfibios sern: 1)
prdida directa del hbitat en cada
sitio de construccin, 2) cambios en
la calidad del hbitat adyacente a
cada sitio (i.e., efecto de borde), y
3) sedimentacin en los arroyos y en
los cuerpos de agua si es que el
suelo se erosiona en los sitios de
construccin. Debido a que la
mayor parte de las especies estn
adaptadas a las condiciones dentro
del bosque, ya ha habido un efecto
en los sitios que han sido abiertos o
alterados. Sin embargo, hay dos
especies que sobreviven en lugares
abiertos - los sapos Bufo marinus y
una especie de rana leptodactila.
Una vez que el bosque se deforesta
para la creacin de un pozo, los
rboles y otro tipo de vegetacin
son expuestos a una cantidad mayor
de luz, temperatura, humedad, y
viento. La composicin de la
vegetacin en los bordes del bosque
cambia como respuesta a las nuevas
condiciones ambientales. Ya que
muchas especies de anfibios de los
bosques tropicales estn adaptadas
a las condiciones del interior del
bosque, muy pocas especies toleran
las zonas abiertas o los bosques de
borde (Fig. 12). Son pocas las
especies que pueden vivir en el
suelo y reproducirse en los charcos
de agua y que pueden alimentarse
de los insectos atraidos a las luces.
La sedimentacin no parece ser un
problema dentro de las operaciones
hoy da en los pozos San Martn-3 y
Cashiriari-2 (basado en la presencia
de renacuajos), aunque las
operaciones en el futuro pudieran
ocasionar daos en la estructura de
la comunidad de anfibios.
Un programa de monitoreo para
los anfibios de la parte baja del
Urubamba puede ayudar a
determinar si los cambios en la

distribucin de especies se debe a
que el hbitat ha sido alterado a
otro fenomeno. Si es conducido por
varios aos, tambin se puede
determinar qu especies, si existen,
han sido impactadas por las
operaciones realizadas y si los
cambios en la fauna que ocurrieron
en el tiempo se deben a alteraciones
ambientales a gran escala. Los
cambios en la fauna debidos a
condiciones climticas o
perturbaciones atmosfricas no se
esperan en un futuro cercano. Sin
embargo, el anlisis de los cambios
en la distribucin y abundancia de
anfibios debe de tomar en cuenta
cambios posibles en las condiciones
abiticas.
Las metas principales del
programa de monitoreo de anfibios
son:
* evaluar los efectos de la
construccin y operacin de cada
pozo de exploracin, del gaseoducto
y de la planta de gas en las
comunidades de anfibios,
* evaluar los patrones a largo
plazo en la estructura de la
comunidad y en la distribucin
relativa de las especies de anfibios
con respecto a cambios en el hbitat
y en el clima en esta regin del
Urubamba, y
* incrementar los inventarios
faunsticos y de historia natural de
las especies que se encuentren en
cada sitio usando datos del
programa de monitoreo. Estas
metas tendrn buenos resultados
siempre y cuando se utilicen
protocolos estandarizados para el
monitoreo.
Resumen
Ejecutivo
El anlisis de
los cambios
en la
distribucin y
abundancia
de anfibios
debe de
tomar en
cuenta
cambios
posibles en
las condiciones
abiticas.
Aves
Aunque las comunidades de
aves tropicales estan tipificadas por
una alta riqueza de especies,
lxix
Resumen
Ejecutivo
lxx
muchas de stas son raras

localmente, lo que las hace
vulnerables a extinciones locales
causadas por la actividad humana.
Parte de la gran diversidad que se
encuentra en estas comunidades se
debe al grado de especializacin de
hbiat que muchas especies tienen
encontrndose en un solo hbitat o
varios micro-hbitats y que son
incapaces de sobrevivir si el hbitat
es alterado, degradado o eliminado.
Hasta el momento, no es
posible monitorear todos los
impactos que el desarrollo pueda
causar en las poblaciones del rea
de estudio. Por tanto, esta discusin
se limita a los impactos esperados
del futuro desarrollo del rea si se
inicia la produccin a gran escala de
gas natural (i.e., instalacin
permanente de pozos de
produccin, su operacin a largo
plazo, y la construccin del
gaseoducto subterraneo con la
reforestacin a lo largo de la ruta).
Se asume que no habr carreteras de
acceso permanente hacia el interior
de la regin, que SPDP u otras
autoridades continuarn
controlando el acceso, por lo que la
colonizacin descontrolada no
debera ocurrir. Si estas condiciones
no se cumplen, programs
adicionales de monitoreo sern
necesarias para examinar los efectos
por la deforestacin y la
fragmentacin del bosque.
Tomando en cuenta estas
observaciones, los impactos de
desarrollo ms importantes sern
aquellos causados por la presencia
de los pozos, por la planta de gas y
por los gaseoductos. El rea total de
bosque que ser aclareada para cada
pozo y para la tubera del
gaseoducto ser pequea comparada
con el rea total de bosque, por lo
que el efecto directo de la
deforestacin en las poblaciones de

aves ser poco. Sin embargo, los
efectos potenciales indirectos de
hacer un aclareamiento en el bosque
podra afectar la poblacin de aves
en una rea ms grande, debido al
efecto de borde y a la colonizacin
de especies.
SPDP prohibe la caza cerca de
los pozos, ya sea con arms de
fuego u otro tipo de instrumento.
Una vez que el gaseoducto sea
construido, el control de acceso al
rea del proyecto ser ms dificil,
an si la ruta del gaseoducto se
reforesta. Por tanto, el monitoreo de
aves de caza a lo largo de la ruta del
gaseoducto es imperativo. Adems,
el establecer estaciones de control
en colaboracin con las
comunidades indgenas ayudar a
prevenir el acceso de personas no
autorizadas. La eficiencia de este
mtodo puede evaluarse por medio
del programa de monitoreo.
Cazadores indgenas y algunos
grupos ambientalistas han
especulado que el ruido y las
actividades humanas en los pozos
podra afectar negativamente la
fauna del rea, posiblemente
espantando a los animales de los
sitios. Con base a observaciones
obtenidas en este estudio, los
investigadores han hipotetizado que
ni el ruido (incluyendo el generado
por los helicopteros), ni la actividad
humana sin caza de animales, ni
otro tipo de persecucin, tendrn
efectos a largo plazo en la fauna del
bosque siempre y cuando no se
altere el hbitat. Esto puede ser
comprobado al estudiar las especies
de caza en los pozos, y en el
gaseoducto.
Las siguientes prioridades se
recomiendan para el programa de
monitoreo:
Los Pozos
debern ser incluidos en el
programa de monitoreo. Sitios de
estudios adicionales sern incluidos
conforme nuevos pozos sean
establecidos en hbitats diferentes.
Las prioridades son:
* Establecer un sistema para
evaluar las reas cercanas a los
pozos para incluir muestreos en
sitios afectados y en sitios no
afectados.
* Iniciar una evaluacin con
redes de niebla en estas reas para
evaluar el efecto de borde.
puntos de muestro para
complementar la evaluacin del
efecto de borde.
* Establecer transectos e iniciar
evaluaciones de las densidades de
las aves de caza y otras especies
que son explotadas.
El Gaseoducto
El monitoreo deber ser
conducido como mnimo en dos
sitios, uno a unos cuantos
kilmetros del punto en donde el
gaseoducto empieza y otro dentro
del bosque no alterado.
Las prioridades son:
* Establecer un sistema con
tres reas de muestreo en cada sitio
de estudio que incluya una rea
control y una cerca del campamento
base.
redes de niebla en estas reas para
evaluar el efecto de borde.
* Establecer transectos e iniciar
evaluaciones de las densidades de
las aves de caza y otras especies
que son explotadas.
puntos de muestro para
complementar la evaluacin del
efecto de borde.
Capacitacin
Un curso de entrenamiento y
capacitacin de tres semanas
conducido por expertos en aves
tanto Peruanos como extranjeros,
producir asistentes de campo
capacitados para conducir los
programs de monitoreo para la
regin baja del Urubamba.
Resumen
Ejecutivo
Mamferos
Pequeos Mamferos Novoladores
El programa de monitoreo
debe de utilizar protocolos
estandarizados para poder comparar
los datos obtenidos tanto en los
pozos como en el gaseoducto. El
objetivo principal es determinar la
distribucin y abundancia de las
poblaciones de pequeos mamferos
no-voladores. Esto va a requerir
procedimientos efectivos para
optimizar la captura de especmenes
mientras se facilita la comparacin
de las variables dentro del estudio.
El programa requiere que se
seleccione un adecuado nmero y
tipo de hbitats directamente
relacionados con el uso de una
variedad de mtodos y trampas
(audio y visuales) usados para
identificar mamferos.
Murcilagos
En la regin baja del
Urubamba, como en otras partes del
mundo, no hay un metodo nico
para monitorear a todos los
murcilagos. Una combinacin de
tcnicas de observacin y mtodos
de muestreo debern ser usados
para evaluar varias especies de
murcilagos. Estas incluyen el
conteo directo de sitios de
descanso, conteos en las
guarniciones cuando se les perturba,
conteos nocturnos de dispersin,
conteos directos visuales, conteos
lxxi
Resumen
Ejecutivo
con detectores de movimiento,

detectores de murcilagos con
ultrasonido, y por captura directa.
Mamferos Medianos y
Grandes
Las actividades que se han
realizado hasta ahora estn
enfocadas a determinar qu
especies hay en el rea cerca de los
pozos de exploracin. Para poder
determinar los efectos que
resultarn por la exploracin de gas
natural, es muy importante obtener
una base de datos de referencia.
Los investigadores enfatizan
que los retos que posiblemente
tenga el proyecto sern el resultado
de la construccin del gaseoducto si
no se toman las medidas necesarias
para evitar su uso por personas no
autorizadas. El monitoreo de la
biodiversidad proveer informacin
necesaria para que por medio del
manejo adaptativo se pueda influir
positivamente en la toma de
decisiones para evitar problems
ecolgicos.
Basados en este razonamiento,
las siguientes cuatro prioridades se
han propuesto para el estudio de los
mamferos medianos y grandes,
durante la siguientes fases del
programa de evaluacin y
monitoreo en el bajo Urubamba.
Aunque el foco se haga en
lxxii
mamferos medianos y grandes,

estas prioridades se pueden aplicar
a otros grupos taxonmicos.
Los siguientes prioridades se
recomiendan para el programa de
monitoreo:
* Conducir el monitoreo de la
diversidad de mamferos medianos y
grandes en San Martn-3, Cashiriari2, y las instalaciones asociadas con
la exploracin de gas natural.
* Conducir el monitoreo de la
biodiversidad a lo largo del
gaseoducto de la regin baja del
Urubamba a Lima, as como de las
instalaciones asociadas para su
transporte.
* Desarrollar manuales para la
capacitacin y materiales de apoyo
para asistir a los investigadores y a
sus asistentes durante las
evaluaciones de biodiversidad; e.g.,
producir manuales que contengan
huellas, muestras y seas de
animales del bosque para los
mamferos de America del Sur.
* Capacitar a los bilogos
peruanos en tcnicas para
monitorear mamferos medianos y
grandes.
Biodiversity
Assessment
and Long-term
Monitoring
of the Lower
Urubamba
Region, Per:
San Martin-3
and Cashiriari-2
Well Sites
Introduction
Alfonso Alonso and
Francisco Dallmeier
Smithsonian Institution/Monitoring
and Assessment of Biodiversity
Program (SI/MAB)
The biologically rich Amazonian lowlands of Per represent an
exciting opportunity to integrate
science, conservation, and development through careful planning,
assessment, monitoring, and decision-making. In the Lower
Urubamba region, the Smithsonian
Institutions Institute of Conservation Biology has joined Shell Prospecting and Development (Per)
B.V. (SPDP) in a unique venture

aimed at achieving environmentally
sensitive development of natural
gas and condensate resources.
Collaborators include Peruvian
scientists from the Museo de
Historia Natural of the Universidad
Nacional Mayor de San Marcos,
Cusco, and Universidad Nacional de
la Libertad (Trujillo).
SPDP is currently appraising
gas reservoirs (at the San Martin-3
and Cashiriari-2 well sites) that
were discovered during exploration
in the mid-1980s. This work consists of drilling four wells and then
completing a logistics base. Plans
for additional development of the
gas reservoirs call for flow lines, a
gas plant located in the study area,
and a pipeline across the Andes to
the coast where coastal facilities
will also be built.
Two phases of the project have
been completed. First, a relatively
quick assessment of the biological
and cultural diversity in the Lower
Urubamba region was completed.
This was followed by a workshop
about the area in September of
1996 to set the parameters for a

biodiversity assessment and monitoring program. Participants included representatives of government institutions, non-governmental organizations, the scientific
community, and local Native
groups. A report on Phase I was
published in early 1997 (Udvardy
and Sandoval 1996).
Phase II of the project included
initial assessments of biodiversity
in the study region and establishment of multi-scale plots and
permanent biodiversity monitoring
plots. SI/MAB Series #1 is the final
Phase II report (January-July 1997).
It describes the results of work to
date and explains the steps that
have been taken to begin Phase III
of the project (September-December 1997).
The report begins with a short
overview of various aspects of
Phase II, followed by a chapter
describing the study region and a
chapter discussing the adaptive
management process and the advantages of incorporating its principles
into the project. The next chapter
addresses assessments of biological
diversity and the long-term monitoring plan. The final chapters
present the results from the initial
surveys and assessments of vegetation, invertebrates, amphibians and
reptiles, birds, and mammals, as
well as a description of the proposed monitoring programs for the
selected taxa.
Overview of Phase II
The overall goals of
Smithsonians work are to link
vegetation information to data
about other taxa that are being
assessed and monitored and apply
the results to management decisions
at the well sites and, later, in the
2
larger area. Multi-disciplinary teams

of researchers are carrying out the
project under the research and
training framework devised by SI/
MAB (Smithsonian Institution/
Biodiversity Program) for longterm, multi-taxa forest biodiversity
monitoring at permanent research
sites (Appendices 2 and 3).
Objectives
* Conduct a baseline
biodiversity assessment of target
groups at well sites San Martin-3
and Cashiriari-2,
* Select and establish multiscale vegetation plots to estimate
species/area relationships and
determine the number and size of
long-term biodiversity research
plots needed to monitor vegetation
and other selected groups, and
* Study the area and gather
data from trails and survey sites.
Tasks
* Identify professional gaps in
the work force and implement
training of technical personnel and
para-taxonomists (individuals that
separate and group invertebrate
specimens with the same
morphology).
* Assess needs for infrastructure building, including personnel
and scientific equipment.
* Identify partner organizations
and establish an operating framework and protocols for field work,
sampling, preparation, and deposition of specimens.
* Conduct biodiversity inventories and assessments at the time
of clearing of well sites, trails, and
the areas adjacent to the wells.
* Survey and establish multiscale and biodiversity monitoring
plots.
Mammals
16%
Multi-scale and
biodiversity plots
16%
Birds
7%
Plants
16%
Amphibians and
Reptiles
19%
Invertebrates
26%
Figure 1. Percentage of researchers by specialty (%) for the assessment

of biodiversity of each taxanomic group.
* Complete field work in small
groups.
* Complete field and laboratory
preparation of voucher specimens.
* Set up species database
management and compile all gathered information.
* Prepare final Phase II report.
Schedule
December 1996-January 1997
* Develop an operational
framework that will include establishing protocols with local institutions for field work, sampling,
preparation, and deposition of
specimens (Appendix 2).
February-June 1997
* Conduct biodiversity assessments at the time of clearing of
well sites and in the areas adjacent
to the wells.
* Design monitoring plan.
* Survey and establish multiscale and biodiversity monitoring
plots.
* Complete interim Phase II
report.
June-August 1997
* Prepare final Phase II report.
Phase III of the project (September-December 1997) will
include:
* Survey the area along the
Camisea River between the well
sites and the Urubamba River for
assessment and multi-scale vegetation plots and biodiversity monitoring plots establishment.
* Biodiversity assessment of
Cashiriari-3 well site.
Personnel
The project is being conducted
by multi-disciplinary teams of
researchers from many national and
international institutions (Appendix
3, Appendix 1). Our assessment
was targeted to five biologically
diverse groups that included vegetation (trees, shrubs, ferns), invertebrates (insects, snails, spiders),
amphibians and reptiles (frogs,
salamanders, snakes, lizards), birds
and mammals. Fifty-two individuals
collaborated in Phase II. Thirtynine researchers from several
For each of
the taxa
studied, the
biological
community
appeared to
be in nearly
pristine
condition.
Peruvian and foreign institutions

participated in the field assessments, seven concentrated on data
management and interpretation, and
six provided technical support.
Research at the field site was
assisted by six guides from the
Machiguenga and Yaminahua
communities. Resources were
allocated in relation to the taxa
sampled (Fig. 1).
Logistical support
SPDP provided financial and
logistical support. We arrived to
the logistics support base camp at
Nuevo Mundo by charter plane
directly from Lima. We flew in
helicopters that SPDP is providing
as the way of transportation from
Nuevo Mundo to the well sites, and
thus avoiding the impacts of road
building. The Peruvian construction
company COSAPI, provided logistical support within the camps and
facilitated our work by providing
tools and personnel. Concessions
and Catering S. A. provided highquality food.
The Well Sites
San Martin-3
San Martin-3 is found approximately five kilometers (km) north
of the Camisea River at an altitude
of 474 m (geographical reference
coordinates are 11 47' 09.8"S, 72
42' 05.3"W). The area contains
steep hills (slopes range from 20 to
70) crossed by numerous rivers and
streams and an abundance of dense,
spiny, bamboo stands (Guadua
sarcocarpa) known locally as
"pacales." This terrain makes
walking difficult. For example, it
took researchers an entire afternoon
to obtain one undescribed species
of Pleurothyrium, a plant of the
avocado family (Lauraceae). The
site can be described as a canopy

emerging grassland with very few
large trees, a relatively abundant
understory layer, and numerous
epiphytes and lianas. The Cashiriari
15 km south-west of the drilling
site. We conducted the biodiversity
assessment at this site from January
through April of 1997, at the same
time that SPDP was constructing a
3.5-ha platform for natural gas
exploration.
Cashiriari-2
Cashiriari-2 is located approximately four km south of the
Camisea River and three km northeast of the Cashiriari River at an
altitude of 579 m (geographical
position is 11 51' 51.3"S, 72 46'
45.6"W). This site is characterized
by a hilly terrain with rather gentle
slopes (no rocky outcrops) dissected
by small creeks. Soils are, for the
most part, deep and clayey. The
vegetation surrounding the drilling
site is covered by primary, wet,
closed forest with numerous representatives of different botanical life
forms such as trees, herbs, epiphytes, and lianas. Large trees are
common, the understory vegetation
is moderately abundant, and there
are no bamboo stems. Within the
area of the platform, adult pioneer
species were observed. They apparently colonized the site after it was
cleared in the 1980s. The Cashiriari
six km northwest of the drilling
site. We carried out the biodiversity
assessment at Cashiriari-2 from May
through June of 1997, while SPDP
was conducting drilling operations.
Findings
Overall, we found that the

biological diversity at each of the
two sites was extraordinarily high.
In addition, for each of the taxa
studied, the biological community
appeared to be in nearly pristine
condition. There was virtually no
evidence that human activities have
had significant impacts. The main
perturbations that may affect
biological communities are 1) direct
habitat losses, 2) changes in the
quality of the habitat, and 3) siltation. Direct habitat losses are the
obvious result of constructing
operational facilities. To date, the
areas occupied by those facilities
are small islands in a sea of green
vegetation. Changes in the quality
of habitat occur in the areas that
are opened and along their edges
where pioneer forest species are
exposed to more sunlight and higher
levels of temperature, humidity,
and wind. These factors may change
the species composition of the
disturbed areas or the surrounding
forest, and changes in habitat will
affect the structure of biological
communities. Soil washing into
streams from the construction sites
may cause abnormal levels of
siltation, although this does not
now appear to be a problem in the
small streams near the well sites,
based on the presence of tadpoles
and invertebrate aquatic fauna.
More specifically, our findings
indicate the following:
* Tree species densities proved
to be among the highest known to
date anywhere in the world. At our
1-hectare (ha) permanent
biodiversity monitoring plots, we
found, on average, 251 species of
trees represented by 1500 individuals with diameters equal to or
greater than 4 centimeters (cm).
Many species of herbs, lianas, and

epiphytes were also abundant in the
forest.
* The Lower Urubamba may be
one of the most diverse areas on
earth for nocturnal butterflies. The
region contains a high diversity of
invertebrates, including many in the
groups we are studying (wasps,
bees, butterflies, dragonflies, ants,
spiders, and snails, among others).
We found 87 species of nocturnal
butterflies (subfamily Ctenuchinae)
in just 18 days of field work, or
approximately 85% of the 103
species found for Tambopata, Per,
during a much more intensive study.
We also discovered new supra
generic taxaSclerogibbidae and
Scolebithydaefor Peruvian entomofauna.
* Amphibians (e.g., frogs and
toads) and reptiles (e.g., lizards and
snakes) are very important components of all tropical forests. We
documented more than 80 species
of amphibians and reptiles at the
sites, which included 43 amphibians
and 44 reptiles.
* Bird species richness at the
well sites may equal or exceed that
of equivalent habitats in Manu
National Park, one of the most
diverse areas for birds known in the
world. We recorded 198 bird species
at San Martin-3 and Cashiriari-2. In
addition, the bird communities
(including game birds) at the well
sites appear to be largely
undisturbed.
* We sampled 51 bat species at
the well sites during just one month
of field work. This total is very high
when compared to the 55 bat
species recorded at Manu National
Park over five years and the 57
found in French Guyana after 10
years.
Helicopter
transport,
key to
SPDPs
commitment to
minimize
environmental
impacts,
should
continue.
* Neither the well operations

nor the noise they generate seemed
to limit large mammal activity,
except directly on the drilling pad.
We found that 58 species of
medium- and large-sized mammals
were potentially present at the well
sites. Jaguar, ocelot, and tapir
activity was evident close to the
sites. Big cats are often the first
animals to flee human development,
but these animals were active to
within 100 meters (m) of the San
Martin-3 well site.
Key
Recommendations
Based on our findings, we
recommend the following.
Current Operations
* Helicopter transport, key to
SPDPs commitment to minimize
environmental impacts, should
continue. Our findings show that
noise produced by vehicles and
machinery at the well sites is having
few, if any, effects on game animals,
which exhibit a distinct lack of
wariness upon encountering humans. (Part of this behavior is likely
due to low hunting pressure in the
vicinity of the well sites.)
* Botanical studies should be
continued. Research on the plants
used by local Native communities
should be added to the results on
current growth rate and distribution
to increase understanding of the
natural cycles of species abundance
and scarcity. This activity will also
provide an opportunity for more
participation by local people in the
biological assessments.
* Fisheries studies should also
be continued. Fish are particularly
important because they are a major
source of protein for Natives. A
biological assessment of fish fauna

along the Urubamba and Camisea
rivers will provide baseline data for
work on fish migration cycles,
reproductive patterns, and growth
rates, as well as understanding of
natural cycles of fish species.
The Longer Term
The outline below lists important recommendations for the next
phase of the project. The final
section of this report presents
details of the second
recommendation.
* Complete baseline
biodiversity assessments at the well
sites.
* Establish a long-term monitoring program for target groups
vegetation, invertebrates, amphibians and reptiles, birds, and
mammalsat all of the well sites.
* Conduct a biodiversity assessment of the pipeline area and
design a monitoring program for
selected points along the pipeline.
* Assess selected points in six
geological regions identified on
satellite images.
* Use the study process as an
opportunity to train Natives in
natural resource management.
* Develop educational materials and films along with a training
package for use with local communities and the public in the region.
* Convene a symposium in
1998 to present project findings;
publish the proceedings of the
symposium.
References
Udvardy, S., and A. Sandoval. 1996.

Proceedings from the Workshop on
Biological and Cultural Diversity
of the Lower Urubamba River.
Smithsonian Institution,
Washington, DC.
Appendix 1. Urubamba program participants.

Overall Direction
Francisco Dallmeier
Director, Monitoring and Assessing
Biodiversity Program (SI/MAB)
National Museum of Natural History
10th & Constitution Ave, NW
Washington, D. C. 20560-0180
Tel: (202) 786-3114
Fax: (202) 633-8918
Email: fdallmeier@ic.si.edu
Tom Lovejoy
Director
Institute for Conservation Biology
1000 Jefferson Drive, S.W., Suite 320
Washington, D. C. 20560
Tel: (202) 786-2263
Fax: (202) 786-2304
Email: lovejoy.tom@si.edu
Email: tlovejoy@worldbank.org
Don Wilson
Director
Office of Biodiversity Programs
Tel: (202) 786-2944
Fax: (202) 786-2934
Email: wilson.don@nmnh.si.edu
Coordination and Administration
Alfonso Alonso
Scientific Coordinator
Urubamba Project
SI/MAB Program
Tel: (202) 357-4793
Fax: (202) 786-2557
Email: ic.aalonso@ic.si.edu
Tatiana Pacheco
SI/MAB Program Administrator
Tel: (202) 786-3114
Fax: (202) 633-8918
Email: ic.tpacheco@ic.si.edu
Jose Santisteban
Research Associate, Urubamba Project
Departamento de Entomologia
Universidad Nacional de la Libertad
(Trujillo)
Bartolome de las Casas #1599-D
La Esperanza, Trujillo
Tel: 0-44-27-1658
Email: jlsac@chanchan.unitru.edu.pe
Abelardo Sandoval
Field Manager, Biodiversity Programs
Washington, D.C. 20560-0180
Tel: (202) 633-9805
Fax: (202) 786-2934
Email: sandoval.abelardo@nmnh.si.edu
Marsha Sitnik
Program Administrator
Biodiversity Programs
Tel: (202) 786-2821
Fax: (202) 786-2934
Email: sitnik.marsha@nmnh.si.edu
Shana Udvardy
Program Manager, Urubamba Project
SI/MAB Program
Tel: (202) 786-3114
Fax: (202) 633-8918
Email: sudvardy@simab.si.edu
Appendix 1. Urubamba program participants (Cont.).

Authors and Contributors
Pedro Acevedo
Department of Botany
Tel: (202) 786-2692
Fax: (202) 786-2563
Email: Acevedo.Pedro@nmnh.si.edu
Cesar Aguilar
Museo de Historia Natural
Universidad Nacional
Mayor de San Marcos
Avenida Arenales 1256
Jess Mara, Apartado 140434, Lima 14
Tel: 511-471-0117
Email: jhcordova@musm.edu.pe
Jorge Aliaga
Asociacion Perana para la
Conservacion de la Naturaleza
Parque Jose de Acosta 187
Magdalena - Lima 17, Per
Tel: (51) 1-264-0970
Fax: (51) 1-264-3027
Email: apeco@amauta.rcp.net.pe
Leeanne Alonso
Conservation International
2501 M street, NW, Suite 200
Washington, D.C. 20037
Tel: (202) 973-2282
Fax: (202) 887-0193
Email: lalonso@conservation.org
Cathy Anderson
Department of Entomology
Tel: (202) 357-1856
Email: anderson.cathy@nmnh.si.edu
George Angehr
Smithsonian Tropical Research Institute
Unit 0948
APO AA 34002-0948
Tel: (507)227-6022 ext 2343
Fax: (507) 232-5978
Email: stri.tivoli.angehrg@ic.si.edu
Constantino Aucca
Urbanizacion Ttio Q-1-13
Pasaje Uriel Garcia Wanchac
Cusco, Per
Tel: 511-0-84-23-5850
Email: manuexpe+@amauta.rcp.net.pe
Robert Baker
Department of Biology
Texas Tech University
Lubbock, Texas 79409
Tel: (806) 742-2702
Fax: (806) 742-2963
Email: rjbaker@ttu.edu
Severo Balden
Mayor de San Marcos
Tel: 471-0117
Deborah Bell
Tel: (202) 357-2795
Fax: (202) 786-2563
Email: bell.deborah@nmnh.si.edu
Hamilton Beltrn
Mayor de San Marcos
Tel: 511-471-0117
Email: hamilton@musm.edu.pe

Major Boddicker
Rocky Mountain Wildlife Enterprises
P.O. Box 999
La Porte, CO 80535
Tel: (970) 484-2768
Fax: (970) 484-0807
Email: dlindahl@lamar.colostate.edu
John Chavez
Mayor de San Marcos
Tel: 511-471-0117
Geneva Chong
Midcontinent Ecological Science Center
Biological Resources Division
U.S. Geological Survey
Natural Resource Ecology Laboratory
Colorado State University
Fort Collins, Colorado 80523-1499
Tel: (970) 491-1980
Fax: (970) 491-1965
Email: geneva@nrel.colostate.edu
James Comiskey
Research Ecologist, SI/MAB Program
Tel: (202) 786-3114
Fax: (202) 633-8918
Email: ic.jac@ic.si.edu
Jesus Crdova
Mayor de San Marcos
Tel: 511-471-0117
Email: jhcordova@musm.edu.pe
Saida Crdova
Mayor de San Marcos
Tel: 511-471-0117
Email: aracno@musm.edu.pe
Timoteo Driskell
12 Old Steele Rd.
Ashburnham, MA 01430
Tel. (508) 827-3350
Email: Tdriskell@net1plus.com
Janine Duarez
Mayor de San Marcos
Tel: 511-471-0117
Email: aracno@musm.edu.pe
Albert Finnamore
Provincial Museum of Alberta
12845 - 102 Avenue
Edmonton, Alberta, Canada T5N 0M6
Tel: (403) 453-9177
Fax: (403) 454-6629
Email: Afinnamore@mcd.gov.ab.ca
Thomas Fritts
US Geological Survey
Division of Amphibians and Reptiles
10th & Constitution Ave., NW
Tel: (202) 357-1930
Fax: (202) 357-1932
Email: fritts.thomas@nmnh.si.edu

Steve Gotte
US Geological Survey
Division of Amphibians and Reptiles
Tel.: (202) 357-4805
Fax: (202) 357-1932
Email: Gotte.Steve@nmnh.si.edu
Juan Grados
Mayor de San Marcos
Tel: 511-471-0117
Email: grados@musm.edu.pe
Ricardo Guerrero
Instituto de Zoologia Tropical
Facultad de Ciencias
Universidad Central de Venezuela
PO Box 47058
Caracas 1041A
Tel: (58) 1-605-2207
Fax: (58) 1-605-2204
Email: rguerrer@strix.cienc.ucv.ve
Javier Icochea
Mayor de San Marcos
Tel: 511-471-0117
Email: javier@musm.edu.pe
Manuel Laime
Mayor de San Marcos
Tel: 511-471-0117
Email: plozada@musm.edu.pe
10
Jerry Louton
Department of Entomology
Tel. (202) 357-1867
Email: louton.jerry@nmnh.si.edu
Shahroukh Mistry
Department of Ecology Evolution and
Natural Resources, Rutgers University
14 College Farm Road
New Brunswick, New Jersey
08901-8551
Tel: 732-932-4520
Fax: 732-932-4517
Email: smistry@rci.rutgers.edu
Joseph Mitchell
Department of Biology
Univerisity of Richmond
Richmond VA 23173
Tel/Fax: (804) 740-7086
Email: jmitchel@richmond.edu
Percy Nuez
Facultad de Ciencias Biologicas
Universidad San Antonio
Abad del Cusco
Cusco, Per
Email: posacus+@qenqo.rcp.net.pe
Rina Ramirez
Mayor de San Marcos
Tel: 511-471-0117
Email: rina@musm.edu.pe
Catherine Rankin
Tel: (202) 357-4369
Fax: (202) 786-2563
Email: rankin.catherine@nmnh.si.edu

Robert Reynolds
Division of Amphibians and Reptiles,
Tel: (202) 357-4803
Fax: (202) 357-1932
Email: reynolds.robert@nmnh.si.edu
Juan Jos Rodrguez
Mayor de San Marcos
Tel: 511- 471-0117
Email: jrodrigu@li.urp.edu.pe
Christopher Ros
SI/MAB Education Coordinator
Tel: (202) 786-3116
Fax: (202) 633-8918
Email: ic.cjr@ic.si.edu
James Siegel
US Fish and Wildlife Service
Shepherdstown, WV 25443
Tel: (304) 876-7482
Fax: (304) 876-7225
Email: Jim_Siegel@fws.gov
Sergio Solari
Mayor de San Marcos
Tel: 471-0117
Email: ssolari@musm.edu.pe
Thomas Stohlgren
Midcontinent Ecological Science Center
Biological Resources Division,
Natural Resource Ecology Laboratory
Fort Collins, Colorado 80523-1499
Tel: (970) 491-1980
Fax: (970) 491-1965
Email: toms@nrel.colostate.edu
Guillermo Tello
Justo Vigil 469
Lima 17
Tel: 511-461-1884
Fax: 511-461-1962
Email: 9012035@li.urp.edu.pe
Gorky Valencia
Universidad San Antonio Abad
Urbanizacion Mateo Pumacahua
Segunda Etapa H11
Wanchaq, Cusco, Per
Tel: 511- 0-84-22-6782
Email: gork@quenqo.unsaac.edu.pe
Steve Smith
Tel: (202) 357-2541
Fax: (202) 786-2563
Email: smith.steve@nmnh.si.edu
11
Appendix 1. Urubamba program participants (Cont.)

Field Assistants
Antonio Gomez
(Yaminahua Community)
Sepahua
Distrito Sepahua
Provincia Atalaya
Departamento Ucayali, Per
Justino Martn
(Machiguenga Community)
Kiriguetti
Distrito Echarate
Provincia La Convencion
Departamento Cusco, Per
Jacobo Pacaya
Nuevo Mundo
Distrito Echarate
Ismael Pascal
Kiriguetti
Distrito Echarate
Federico Ramrez
(Yaminahua Community)
Calle Barrio Julio C. Tello
Sepahua
Distrito Sepahua
Provincia Atalaya
Departamento Ucayali, Per
Jos Tenteyo
Apartado 17 Satipo Junin
Communidad Nativa Timpia
Distrito Echarate
12
Appendix 2. Program of activities for Phase II of the Lower Urubamba project.

1
January
2 3 4
February
2 3 4
March
2 3 4
April
2 3
May
2 3
June
2 3
July
2 3
August
2 3 4
Site Selection for Biodiversity Plots

San Martin-3
Cashiriari-2
Multi-scale Biodiversity Plots
San Martin-3
Cashiriari-2
Topography/mapping
San Martin-3
Cashiriari-2
Data Processing and Analysis
San Martin-3
Cashiriari-2
Vegetation Assessment
San Martin-3
Cashiriari-2
Invertebrate Assessment
San Martin-3
Cashiriari-2
Amphibian/Reptile Assessment
San Martin-3
Cashiriari-2
Bird Assessment
San Martin-3
Cashiriari-2
Mammal Assessment
San Martin-3
Cashiriari-2
Report Preparation
Development of Monitoring Plan
13
Appendix 3. Program of biodiversity assessment activities of personnel at the well sites.

January
February
10 13 15 17 20 22 24 27 29 31 14 17 19 21 24 26 28 3
March
5
7 10 12 14 17 19 21 24 26 28 31
COORDINATORS
Alfonso Alonso
Abelardo Sandoval
Francisco Dallmeier
BOTANIST
Pedro Acevedo
Deborah Bell
Steve Smith
Catherine Rankin
Percy Nunez
Severo Baldeon
Hamilton Beltran
INVERTEBRATES
Gorky Valencia
Jose Santisteban
Jerry Louton
Cathy Anderson
Leeanne Alonso
Juan Grados
Rina Ramirez
Saida Cordova
Manuel Laime
Albert Finnamore
AMPHIBIANS
Javier Icochea
Guillermo Tello
Bob Reynolds
Steve Gotte
Tom Fritts
Joe Mitchell
Cesar Aguilar
Jesus Cordova
BIRDS
Constantino Aucca
George Angehr
Jim Siegel
MAMMALS
Ricardo Guerrero
Major Boddicker
Sergio Solari
Don Wilson
Robert Baker
Juan Jose Rodriguez
John Chavez
TOPOGRAPHERS
Benjamin Guerrero
Tim Driskell
FIELD GUIDES
Federico Ramirez
Jacobo Pacaya
Ismael Pascal
Justino Martin
Jose Tenteyo
Antonio Gomez
14
Appendix 3. Program of biodiversity assessment activities of personnel at the well sites (Cont.).
April
2
9 11 14 16 18 21 23 25 28 30 2
May
5
9 12 14 16 19 21 23 26 28 30 2
June
4
9 11 13 16 18 20
COORDINATORS
Alfonso Alonso
Abelardo Sandoval
Francisco Dallmeier
BOTANIST
Pedro Acevedo
Deborah Bell
Steve Smith
Catherine Rankin
Percy Nunez
Severo Baldeon
Hamilton Beltran
INVERTEBRATES
Gorky Valencia
Jose Santisteban
Jerry Louton
Cathy Anderson
Leeanne Alonso
Juan Grados
Rina Ramirez
Saida Cordova
Manuel Laime
Albert Finnamore
AMPHIBIANS
Javier Icochea
Guillermo Tello
Bob Reynolds
Steve Gotte
Tom Fritts
Joe Mitchell
Cesar Aguilar
Jesus Cordova
BIRDS
Constantino Aucca
George Angehr
Jim Siegel
MAMMALS
Ricardo Guerrero
Major Boddicker
Sergio Solari
Don Wilson
Robert Baker
Juan Jose Rodriguez
John Chavez
TOPOGRAPHERS
Benjamin Guerrero
Tim Driskell
Federico Ramirez
Jacobo Pacaya
Ismael Pascal
Justino Martin
Jose Tenteyo
Antonio Gomez
15
16
The Lower
Urubamba
Region
Alfonso Alonso and
Francisco Dallmeier
Program (SI/MAB)
Description
The study area for this project,

approximately 170,000 hectares
(ha) in size, is defined as the land
that encompasses and surrounds the
four proposed well sites and the
centers for logistical activity at
Nuevo Mundo and Camisea (see
map, page 28). Located east of the
Andes Mountains in the Urubamba
River Valley of the Ucayali watershed, the area is bordered by the
northern spurs of the Vilcabamba
Mountains to the west and
Urubamba Mountains to the east
between 10 and 13 latitude and 72
and 74 longitude. The areas lowlands, hills, and mountains vary in
elevation, from less than 500 meters
(m) (lowland rain forest) and 500 to
1,000 m (highland rain forest) to
1,000 m or more(montane cloud
forest). Slopes range from 25% to
70%.
The remainder of this section
summarizes the extensive description of the study area reported in
the Environmental Impact Assessment (ERM 1996).

Geomorphology
The study area exhibits a
diverse geomorphology with three
distinct landscapes, or physiographic units. They include:
* the great alluvial landscape, a
flat depression originating from the
deposition of sediments or the
erosive actions of the Camisea and
Cashiriari rivers and their tributaries;
* the great hill landscape,
which was formed from sandy
sediments, luthite, and clay from
the Tertiary; and
* the great mountainous landscape, typical of areas with high
elevations.
Climate and Temperature
The dominant climate zone is
lowland rain forest (below 500 m)
with a wet season (October-April)
and a dry season (May-September).
Annual rainfall averages between
3,000 and 3,500 millimeters (mm).
Temperatures vary across the
study region, from about 23.5o C to
17
35.0
120
Temperature at 8am
Precipitation(mm)
100
25.0
80
20.0
60
15.0
40
10.0
20
5.0
pr
pr
pr
12
-A
pr
15
-A
pr
18
-A
pr
21
-A
pr
24
-A
pr
27
-A
pr
30
-A
pr
3M
ay
6M
ay
9M
ay
12
-M
ay
9A
6A
ar
3A
ar
31
-M
ar
28
-M
ar
25
-M
ar
22
-M
19
-M
ar
16
-M
ar
0.0
13
-M
Precipitation (mm)
Temperature at 8 am
30.0
Date
Figure 1. Temperature and precipitation registered at San Martin-3 well site, Lower
Urubamba region.
24.5o C (Fig. 1). As in many other
tropical regions, the temperature
remains fairly constant. So does the
average relative humidity, which is
high (more than 80%). In Phase II
of the study, our work study overlapped the transition period between the wet and the dry seasons.
The seasonal picture will be completed by incorporating data gathered during future phases of the
project.
Geology
The Lower Urubamba gas
reservoirs are located within a
Cretaceous, gas-bearing vertical
sequence. The sequence is folded in
compressed, elongated, steep
anticlines; the steepness is associated with major thrust faults. San
Martin-3 and Cashiriari-2 are both
located in anticlines separated by
the Camisea syncline. From seismic
data, it is apparent that substantial
horizontal movements are associated with the main thrust fronts; for
example, the Pica that borders the
18
San Martin anticline. There is also

evidence of minor thrust faulting in
a parallel direction to the main
thrust trends. There is no conclusive evidence of cross-faulting.
Soil
Two soil typesthe Entisols
(recent) and Ultisols (ultimate)
dominate the study area. Entisols
occur only on alluvial benches;
hence, they are the minor group.
They are typically young, sandy, and
yellowish with minimal epipedon
development. Ultisols are usually
deep red or yellow soils, with low
native fertility and poor physical
properties. Accumulations of
humified organic matter in the
surface layer are always present, but
minimally so in the study area
(Table 1).
Water
Rivers in the study area are of
the white-water variety. They are
high in suspended sediments but
low in nutrients and organic matter
Table 1. Lithostratigraphy of the Study Area

Formation
Lower red bed
Upper and Lower Vivian
Upper Chonta
Lower Chonta
Basal Lower Chonta
Upper and Lower Agua Caliente
Esperanza
Upper and Lower Cushabatay
Ene and Copacabana
Constituent rock
Cap rock/claystone/shale
Sandstone (gas bearing; Cashiriari only)
Claystone/shale/some limestone
Sandstone/intercalcated claystone/shale (gas bearing)
Mainly sandstone
Sandstone (gas bearing)
Claystone/shale/some limestone
Sandstone (gas bearing)
Limestone (gas bearing)
and with a neutral pH. These rivers

form depositional areas and may be
direct, meandering, or braided
depending on the topography. The
main river of the area, the
Urubamba, has a distinct flood
plain and forms meanders downstream from the Camisea area.
Upstream, the Urubamba is more of
a montane river with rapids and
waterfalls. Other rivers have less
well-developed flood plains.
The water table is tapped for
household use by villagers at
Camisea and Segakiato and is highly
vulnerable to pollution. Water
quality was sampled at various sites
in the study area during work on the
Environmental Impact Assessment
(ERM 1996) for the gas wells. The
results showed that river temperatures averaged 21.2 C (standard
deviation of 0.3 C) at depths
ranging from about 0.3 to 2.0 m.
The pH in the creeks, wells, and
rivers averaged 7.7. The sampled
waters were free of petroleum
hydrocarbon and detectable levels
of the trace minerals, such as
barium, cadmium, chromium,
mercury, and lead. Overall, the
sampling indicated excellent waterquality conditions in the study area,
based on national and international
water-quality standards.
Flora and Fauna

Prior to this study, little was
known about most of the plant and
animal species that inhabit the
Lower Urubamba region. To date, it
appears that the region contains
previously unidentified species,
some of which may be found
nowhere else. In relation to potential endemism, an important factor
to consider is that the study area
may be a Pleistocene refuge. It is
thought that during the Pleistocene,
climatic conditions in the Amazon
Basin changed so that the forest
retreated into isolated pockets to be
replaced by savannas. The forest
refuges served as incubators for
new species of rainforest organisms,
accounting for some of the observed biodiversity in the Amazon.
The Urubamba valley (including the
study area) has been proposed as
one such site.
In addition, a 1991 survey of
protected areas and parks in Per
catalogued rare and endangered
plant species. Manu National Park,
the protected unit nearest to the
study area, is home to numerous
plant species ranked in critical
danger because of rarity or limited
range, in danger for the same
reasons, and rare in the region. It
is possible that some of these
19
The well
sites are in
an area
peopled by
indigenous
groups in
44 Native
communities, with
some
colonist
settlements
along the
Urubamba
River.
20
species or other endangered plants

may be present in the study area.
Because many animals are quite
mobile and because several zones in
the study area intersect, substantial
intergradation in the distribution
and occurrence of animal species
can be expected. The fauna of the
area has value based on ecological
importance and economic and
subsistence importance to local
human populations. Native communities depend on aquatic and terrestrial wildlife for food, clothing, and
ornamentation. There is also a
substantial economic cash value for
the pelts of many animals. As a
consequence, several species have
been hunted close to extinction in
some parts of the Peruvian Amazon, but not in the study area to the
best of current knowledge.
Settlement and Culture
The well sites are in an area
peopled by indigenous groups in 44
Native communities, with some
colonist settlements along the
Urubamba River. Most of the
Natives are Machiguenga. The
communities of Nuevo Mundo,
Camisea, Kiriguetti, Shivankoreni,
Segakiato, and Cashiriari are all
within the study area; the four
closest to the well sites are
Cashiriari, Segakiato, Shivankoreni,
and Camisea. They are recent
settlements, established since 1958,
with a combined population of just
under 1,000 inhabitants.
As with most other Native
peoples in Amazonia, the
Machiguenga rely on the forest for
their livelihood and subsistence. In
the deep Amazon, subsistence and
commerce employment revolves
around a combination of agriculture, gathering, hunting, fishing, and
extraction of forest products. In the
study area, such activity is primarily

limited to subsistence; the people
there have not yet developed
regular commerce. Family income
stems from the sale of agricultural
products to traders who may visit
(such visits have been very limited
in number), the periodic sale of
wood (a communal activity in
which the proceeds are normally
divided among the community on
an equal basis), and minimally by
providing temporary labor to estates
and petroleum companies.
The Machiguenga communities
in the study area have legal rights to
their lands, except for the
Montetoni located in the NahuaKugapakori Reserve. The land titles
specify information on different
land uses and capabilities (crop
cultivation, cattle raising, forestry,
forest protective functions).
Communal ownership is only
applicable to lands suitable for
agriculture and cattle-raising. It is
nontransferable except that forest
lands and forest edges are subject to
the following: 1) free transit via
existing bridges, railways, or forest
paths and those that will be constructed in the future and 2) passage of oil pipelines, gas pipelines,
and other installations for the
exploration and exploitation of
minerals and petroleum.
References
ERM (Environmental Resource

Management). 1996. Camisea
Appraisal Drilling Campaign,
Environmental Impact Assessment. ERM Per S.A., Lima.
Photo Gallery
21
22
23
24
25
26
27
Printer:
Place map and caption from page 49 of SI/MAB Series #2 here.
28
Biodiversity
Assessment
and
Monitoring
for Adaptive
Management
Francisco Dallmeier
Program (SI/MAB)
In the Lower Urubamba
project, we have incorporated
aspects of the adaptive management processsetting objectives,
carrying out an assessment and
monitoring plan for forest
biodiversity, evaluation, and decision-making. This process can be
represented as a cycle (Fig. 1),
which is periodically calibrated to
ensure that appropriate information
from each component feeds the
next level (Holling 1978, Walters
1986, Hilborn 1992). Adjustments
to the components are made as
needed to achieve the objectives.
The cyclical nature of the process is
maintained through refining the
objectives and management decisions based on the ongoing results.
Thus, adaptive management gives
managers the flexibility to adjust

on-the-ground practices.
Adaptive management is well
suited to the type of biodiversity
project underway in Urubamba. The
process serves as a guide in planning and carrying out a sound forest
biodiversity monitoring program. As
hydrocarbon exploration continues
in Urubamba, pressures to provide
adequate development of the
identified gas resource in conjunction with conservation of the
Urubamba region will likely increase, resulting from greater public
and private interest and involvement in use of the local resources.
An assessment and monitoring
process can aid in evaluating existing management approaches and
their impacts on the Urubamba
forest ecosystems and define new
approaches, if needed. Monitoring
provides data needed to ensure that
the effects of management are
within a previously desired range. If
data indicate otherwise, management decisions and practices can be
re-calibrated and adapted to achieve
the desired outcome.
29
Figure 1. Adaptive management cycle for developing biodiversity assessment

and monitoring plan frameworks.
Appropriate management of
forest ecosystems requires the
ability to understand changes in
those systems that are brought
about by both natural influences
and human actions. Natural influences include events such as hurricanes and typhoons, extended outof-season droughts, and volcanic
eruptions. Human-induced effects
comprise a large and varied spectrum from acid rain or dust deposition and the introduction of exotic
species to rapid landscape alterations caused by development. It is
becoming increasingly clear that
species are less able to adapt to
natural changes at local and regional scales when those changes
are compounded by human-caused
alterations (UNEP 1995).
It is no simple task to acquire
the information necessary to understand such changes, especially
30
considering that forest biodiversity

monitoring is a science in the
process of development. For complex tropical forest ecosystems,
researchers are still defining the
most appropriate methodologies to
sample and monitor biodiversity.
Yet, tools such as the guidelines
offered by the adaptive management process are improving monitoring programs and their capacities
for making relevant and reliable
information available to managers.
In this chapter, we describe the
evolving framework initiated for the
sites within the context of adaptive
management.
Forest Monitoring for
Adaptive Management
Our program represents a
coordinated approach to gathering,
analyzing, and disseminating infor-
SPDP & Smithsonian/MAB Biodiversity Program, Lower Urubamba Per
mation about the Lower Urubamba

region and the areas affected by the
hydrocarbon exploration. A series
of protocols and sampling methodologies were tested at the two well
sites by a multi-disciplinary team of
researchers. The type of vegetation
plots established at the study site
have been adopted at more than
200 sites world wide, facilitating
the transfer of comparable data and
providing a global framework for
data analysis and dissemination
(Dallmeier 1992, Dallmeier and
Comiskey 1996). Other protocols
are being refined and tested for the
specific requirements of work at the
well sites and will be implemented
during monitoring. The protocols
will enable integration of scientific
research methods and analysis with
strategies for getting the resulting
information to SPDP managers.
Setting the Objectives
At the San Martin-3 and
Cashiriari-2 well sites, the following
site-specific objectives were emphasized:
* gather baseline information
about forest structure, composition,
and diversity within the established
forest,
* inventory plots and the
adjacent area of the well sites,
* conduct a baseline
biodiversity assessment of five taxa
in the surrounding area of the well
sites and design a monitoring
program, and
* analyze the cartographic
information and aerial and satellite
pictures to identify key habitats for
biodiversity assessment of the
general area.
In planning the site-specific
assessment and monitoring at the
wells, we considered realistic
objectives for sampling the selected
taxa. Monitoring protocols were

also selected to achieve the sitespecific objectives within a reasonable time line. We are emphasizing
the need to get information into the
hands of managers and defining an
appropriate scale for the sitespecific and broader biodiversity
assessments and monitoring.
Assessment and Monitoring
The forest biodiversity plots
and multi-taxa forest assessment
and monitoring provide an essential
tool for constructing the first
biological picture of the well sites.
Subsequent monitoring over time
will provide a biological "movie" of
natural and man-made changes
(Fig.2). These long-term data are
helpful in detecting the magnitude
and duration of changes, how
related taxa are changing, and early
warning of forest health. They also
serve as the basis for formulating
additional research and management hypotheses, and, perhaps most
important, the data can be used to
guide management decisions related
to biodiversity conservation.
Assessments
Most traditional approaches to
biodiversity assessments have been
oriented to improving the prospects
of intense logging in temperate
forests (Hilborn 1992). More recent
biodiversity assessments have also
included habitats and the population status of rare or endangered
species to determine which areas
are important for conservation.
Assessments should include field
surveys, inventories, and literature
reviews to gather information that
is relevant to the site-specific
monitoring program. In all such
cases, research objectives should be
31
Space
Mammal
Community
Bird
Community
Amphibian
Community
Arthropod
Community
Weather Info.
Acid Rain
Soil/
Topography
Vegetation
T2=Second Census
Tn=Nth Census
Time
Figure 2. SI/MAB Program framework for forest biodiversity monitoring.

geared to provide a framework for
decision-makers to evaluate the
consequences of action (or
inaction) regarding the use of forest
resources (Spellerberg 1992).
Biodiversity assessments at
appropriate scales afford decisionmakers the opportunity to consider
higher quality, cost-effective
choices. Generally, the scale is
based on geographical boundaries
of the protected area or conservation unit or on subsets of such
areas. In addition, participation of
all interested parties in the assessment is important during the early
stages to create a consensus as
assessment moves into the planning
and monitoring phases. This also
aids in addressing research needs
and avoiding costly duplication.
These guidelines were followed in
the initial assessment at the study
sites.
32
Monitoring
Understanding how an ecosystem works requires careful study of
the properties and scales of space
and time. Recent research reveals
complex ecological interactions that
determine how organisms are
distributed and their abundance on
local scales (Diamond and Case
1986; Gentry 1988, 1992; Ashton
1992; Hubbell and Foster 1992).
The evidence also indicates that
ecological communities experience
internal and external long-term
changes with sometimes drastic
turnover (Dallmeier et al. 1998).
These long-term changes may have
radical effects on the composition,
structure, and diversity of the
communities, resulting in imminent
local extinctions and colonizations
(Dallmeier and Comiskey 1998).
Such effects can only be perceived through long-term monitoring of ecological communities and
environmental changes (Clinebell et

al. 1995). The impacts of environmental change on communities
operate at various time scales,
ranging from days and weeks (for
example, gypsy moth defoliation in
eastern North America) to decades
(hurricanes and typhoons, extended
droughts), or centuries or millennia
(volcano activity) (Davis 1976,
1986; Anonymous 1994). Such
scales are often difficult to measure
(Fig. 3).
Further difficulty is encountered in relating ecological processes operating at local scales to
those in effect at regional or global
scales (Stohlgren et al. In press). It
is important to consider scale
because localized or small-scale
events can contribute to large-scale
impacts. As examples, in the Caribbean the most severe drought on
record (1994-95) produced a dry
forest mortality of more than 13%
and eliminated most seedlings and

saplings (Dallmeier 1998). The
droughts impact on the forest was
accentuated by the effects of
hurricanes Marilyn in 1995 and
Bertha in 1996. In Bolivias Beni
region (and elsewhere around the
world), annual fires intentionally set
as a management tool are transforming biologically rich forested
areas into less productive
grasslands.
The monitoring program must
measure the effects of such events
in space and time and determine
their consequences for biodiversity.
Reliable scientific information on
habitat composition, structure, and
dynamics is needed. The greater the
knowledge about a particular site,
the more options are available for
promoting sustainable use and
conservation of resources and a
more stable local economic base.
Increasing public awareness of and
Figure 3. Environmental changes and their impact on biodiversity over space

and time.
33
In c r e a sin g
K n o w le d g e
of
B io d iv e r sity
S I/M A B
B io d iv e r sit y
M o n ito r in g
P lo ts
CTFS
50ha
P lo ts
F rug ivor e- ins ect ivor e
10 %
Nect ar ivor es
10 %
Hem at ophag ous

4%
Car nivo res

4%
Inse cti vo res

23 %
A ll T a x a
B io lo g ic a l
In v e n to r y
D e ta ile d
In v e n to r y
T a rg e t
G roups
F rug ivor es
49 %
Sp e ci es Ar e a C ur v e fo r Ba t s
45
40
35
75%
30
25
San M ar t in - 3
Ca sh ir i ar i - 2
20
15
10
5
0
0
10
11
12
13
14
Ti m e ( D a ys )
Cumu lat ive Speci es vs. Captures

90
S an Ma r t i n
80
C a sh i r i ar i
70
Ma n u
Pa na m a
60
B r az il
50
C o st a Ri c
40
30
7
0
20
10
C AP TUR E S
50%
An t Fr e q ue n cie s
30
25%
R e m o te
S e n sin g
20
15
10
25
V a r ia b le
T ra n se c ts
5
0
1
10
N UM B ER O F P LOT S
G .I.S .
In c r e a sin g tim e , te c h n o lo g y , a n d r e so u r c e s
Figure 4. Time, technology, and resources required to increase our knowledge of biodiversity.
appreciation for the benefits provided by that ecosystem will assist
in its protection. In addition, there
will be more possibilities that the
information will prove useful at farremoved sites with similar
characteristics.
Biodiversity monitoring consists of measuring and sampling
species over time and comparing
the results to a predetermined
standard or noting their deviation
from an expected norm (Hellawell
1991). Different inventories over
time may measure the same variables but with different objectives
in mind and without specified
standards for comparing the results
over temporal scales. Only when
standard protocols allow the results
to be compared to a baseline standard does the repetitive inventory
become a monitoring program.
Significantly, the standards used to
compare biodiversity monitoring
34
data provide the basis of calibration

toward or away from an objective or
baseline norm (see evaluation
section).
In essence, biodiversity monitoring provides information about
the status of species and assemblages of species as well as trends
in both. Monitoring can also identify potential cause-and-effect
relationships that can be addressed
through research with an eye toward
use in the decision-making process.
Sampling and
Data-gathering Designs
Credible sampling designs
should provide guidelines for the
most cost-efficient and effective
way to gather and analyze data
while maintaining high qualitycontrol standards. The design of the
biodiversity assessment and monitoring program for the two well
sites rests on the overall objectives
outlined above in conjunction with

site-specific scientific and management objectives. Both types of
objectives help to identify the
information required, but the sitespecific objectives are usually most
important in determining the sampling intensity to allow comparisons
among habitat types and in assessing long-term changes within
habitats and the effects of disturbance. Time, technology and resources are the three key elements
that determine our knowledge of
biodiversity (Fig. 4).
The sampling design of the
project addressed the following
questions:
* What do researchers need to
know about the site (e.g., data on
forest composition, structure, and
diversity as well as site-specific
information such as the effects of
small-scale deforestation, noise,
rain cycles, and land-use patterns)?
* How will data from the
monitoring program will be used
(scientific and management uses)?
* Are the sites representative
of the selected habitat?
* What degree of sampling
resolution is sensitive enough to
detect changes?
* What limits of change are
expected and are important to
detect (for example, changes in
densities of mammals and birds in
the areas surrounding the well sites,
mortality rates of forest stands
because of natural and humaninduced changes)?
* What is the degree of confidence expected from the results?
* Could these results be compared with other rainforest areas?
Indicators
For well site-specific monitoring, we are identifying some indica-
tors groups for use in assessing

environmental quality. For example,
one might select the bamboo forest
at San Martin-3 to estimate damage
to the forest as well as recruitment
and mortality caused by breaking of
the bamboo. We are using the
following criteria to choose indicator taxa for monitoring: 1) the
species within a taxa are good
measures of one or more questions
that the monitoring program was
designed to answer, 2) the indicators should be able to detect a
condition far enough in advance to
assist in addressing the undesired
outcome, and 3) it should be possible to monitor the indicators
within the constraints of a realistic
budget and other such considerations.
In tropical forests where indicator species are not well known,
SI/MAB selects taxonomic groups
as indicators. Birds and vascular
plants are the most frequently used
taxonomic indicators of environmental health (Spellerberg 1992).
Key habitats may also be important
indicators (e.g., flooded forest,
marshes, and bamboo forest in the
area).
Baseline Information
Monitoring requires a base or
expected norm against which all
future changes may be compared.
Baseline information is normally
gathered from the literature and/or
initial surveys of the areas under
study. In the case of San Martin-3
and Cashiriari-2, minimum biological information was available before
this study (ONERN 1987, 1990;
ERM 1996). Our initial assessments
provided representative data of the
biodiversity for selected taxa at the
two well sites. For example, in
setting up the first three biodivers-
35
ity test plots in San Martin-3 and

Cashiriari-2, our first census
resulted in a list of species and a
description of how they are assembled and distributed at the sites.
The
sampling
frequency
is determined
based on
the taxa
to be
monitored.
36
Site-specific Sampling
In designing a scientifically
sound monitoring system, researchers define the population(s) to be
monitored. The population may be
representative of habitats that are
considered important within a
region. For example, in the Beni
Biosphere Reserve of Bolivia,
SI/MAB has defined 14 forested
habitats for monitoring.
At times, certain population
values such as the number of
individuals of a selected species
within a given area may only be
estimated (Tyrrell et al. 1996). A
measure of the standard error of
the mean, along with the reliability
of the calculated value in the form
of a 95% confidence limit, should
be provided to assess monitoring
intensity (Usher 1991). If the
sampling is repeated, the true mean,
which is of particular interest to the
monitoring program, should fall
within these limits. The confidence
limits decrease as a function of
increasing sample size. This is
important because small confidence
intervals provide more accurate
estimates for informed management
decisions. But obtaining these data
is often constrained by logistic and
cost factors because of the large
sample sizes required.
Within a selected habitat,
sampling may be carried out in a
systematic, random, systematic/
random, or stratified fashion
(Cochran 1977, Stohlgren 1994,
Stohlgren et al. 1995, Tyrrell et al.
1996). Systematic sampling requires
establishment of a grid within
which a series of points are

sampled. For random sampling, all
points have an equal probability of
being sampled. Systematic random
sampling requires that the habitat
be subdivided based on topography;
samples are then chosen at random
(Goldsmith 1991). The area may
also be stratified according to the
needs of the monitoring program.
Under the protocol for assessment and long-term monitoring,
sampling sites are being selected
randomly for each well site to
reduce bias. Because so little is
known about the landscape-scale
spatial patterns of biodiversity in
the region, determining the number
of sampling sites will be a calibrated process based on the species
composition and early stages found
during the assessment. The next
chapter discusses these issues and
plot design at the well sites in more
detail.
The sampling frequency is
determined based on the taxa to be
monitored. For trees, this may be on
a scale of years (Dallmeier 1992),
while for arthropods the scale may
only be weeks (Stork 1995). In any
case, the scale should be sufficient
to enable detection of change.
Frequency of sampling at the
Urubamba plots will vary with the
taxa selected for monitoring.
Data Gathering
Data gathering entails the
measurement and assimilation of
information in the field. A central
issue in the data gathering process
is the need to obtain data under
consistent standards. Data collectors must be trained in the standard
protocols that can be used to
facilitate cross-site comparisons and
evaluation in multi-taxa monitoring.
For example, in past efforts related
to forest management, dead and

dying trees were viewed as less
important than live trees because
they had little value for wood. More
recent efforts related to biodiversity
recognize the importance of dead
trees as elements of the community
that support other taxa. The approach promoted in this project
develops and uses standards across
disciplines to ensure that data can
be transformed into information
that is comparable and compatible.
Data Management
and Analysis
Of primary importance to data
management is to ensure that data
are accurately transferred from field
data sheets to a secure place for
storage and making them available
for future analysis. As defined by
Tessler (1995), managing monitoring data sets should include data
entry, verification, validation,
archiving, and documentation. The
protocols for monitoring species
diversity include such data management procedures. Biodiversity plot
data from Urubamba are being
stored and processed in the
biodiversity monitoring database
BioMon, a computer based system
to assist data management process.
Through the BioMon program, data
entry, verification, and validation
are accomplished.
Because monitoring may take
place over long periods of time,
data must be safely archived. Data
documentation incorporates a
number of descriptors such as data
set title, files and their structures,
location, investigator, and dates of
monitoring along with any other
information pertinent to the data.
Under BioMons protocols, a data
dictionary is stored with the
archived data.
Data analysis is essential in

relating data to the established
goals and objectives of the monitoring program. The results from data
analysis in the form of syntheses or
reports need to be available to other
researchers and to managers and
decision-makers. The National
Academy of Sciences (1990) provides the following guidelines for
analysis:
* focus the analysis to ensure
that the priority question or questions regarding the aspect of biodiversity being monitored are effectively answered,
* emphasize the importance of
early detection of critical changes
to allow lower cost solutions,
* recognize that results from
the analysis should contribute to
knowledge about the biodiversity
component being monitored, and
* emphasize that the results
should provide biodiversity managers with a scientific rationale for
setting appropriate standards.
While data analysis should
synthesize large amounts of data to
make the data meaningful to the
user, it is important to maintain the
focus of the monitoring program by
returning to the specific questions
that need to be answered. The
results from the data analysis
should be presented in a format that
enables others to make their own
interpretations without difficulty.
This may involve the preparation of
reports detailing how the findings
affect management of the study
area along with recommendations.
Evaluation
Evaluation elicits answers to
the questions implicit in the
projects objectives, thus allowing
for generation of management
recommendations and calibration of
37
One of
the most
crucial and
challenging
issues in
ensuring
the success
of a monitoring program is the
need to
gain longterm
support.
the monitoring program. In addition, it allows an assessment of the

monitoring program. The National
Academy of Sciences (1990) recommends the following questions for
use in evaluation:
* Are the results of each specific monitoring technique well
integrated with the overall monitoring program?
* Do methods used ensure
reliable, timely, and powerful data
analysis?
* Are the sampled data subject
to the appropriate techniques for
data management and analysis?
* Could the data gathered be
coupled with new technologies for
analysis and management?
* What mechanisms exist or
can be developed to allow timely
transfer of data and information to
managers and decision-makers?
Management approaches should
be viewed as means to test hypotheses to reach operational goals
(Ecological Society of America
1996). Through evaluation of the
monitoring data, managers receive
timely feedback as these hypothesis
are tested. Thus, evaluations are the
tools for improving management by
checking on management actions
and providing guidelines for improvement. For example, when a
predetermined degree of change is
detected, appropriate action is
taken and the results must be
evaluated.
Decision-making
In adaptive management, the
preceding steps lead to decisionmaking. If the findings determine
that biodiversity trends are within
the expected values, monitoring can
continue without substantial alterations. If significant changes in the
trends are observed, managers or
38
decision-makers need to design the

most appropriate response, thus
restarting the cycle of setting
objectives, conducting assessment
and monitoring, and so forth. The
reasons for monitoring are evaluated at this stage. Is the monitoring
still required, and, if so, do the
objectives remain the same?
Inconclusive results require
adjustment of the objectives and
sampling approaches to increase the
degree of precision. However,
careful planning and design of the
monitoring program can reduce the
risk of inconclusive information.
Long-term Commitment
One of the most crucial and
challenging issues in ensuring the
success of a monitoring program is
the need to gain long-term support.
Project personnel must elicit the
endorsement of high-level managers
and the commitment of resource
experts and technicians in the field.
One method of achieving this goal
is to incorporate monitoring as a
routine duty of appropriate personnel. Another is to implement a
reward system for personnel who
detect a problem and take steps to
devise a management-oriented
solution.
A common practice by SI/MAB
is to develop partnerships among
several organizations to assist in
fund raising, standardizing protocols, providing site-specific documentation, and supporting training
for high-quality research and monitoring. This has encouraged reliable
data and information management
and reduced duplication.
Such partnerships have also
aided in disseminating information
about a monitoring program to
inform the public of its importance
and benefits. Greater public under-
standing may lead to increased

acceptance and support for the
program.
References
Anonymous. 1994. Scientists link

CO 2 to changes in tropical forests.
Global Environmental Change Report
6: 1-3.
Ashton, P. S. 1992. The structure
and dynamics of tropical rain
forest in relation to tree species
richness. Pages 53-64 in The
Ecology and Silviculture of
Mixed-Species Forests. (M. J.
Kelty, B. C. Larson, and C. D.
Oliver, eds.) Kluwer, Boston.
Clinebell, R., O. L. Phillips, A. H.
Gentry, N. Start, and H. Zuuring.
1995. Prediction of neotropical
woody plant diversity from climatic and soil data. Biodiversity and
Conservation 4: 56-90.
Cochran, W. G. 1977. Sampling
Techniques. John Wiley and Sons,
New York.
Dallmeier, F. 1992. Long-term
monitoring of biological diversity
in tropical forest areas: methods
for establishment and inventory of
permanent plots. MAB Digest 11.
UNESCO, Paris.
Dallmeier, F., and J. A. Comiskey.
1996. From the forest to the user:
a methodology update. Pages 4156 in The Biodiversity of Southeastern Per: Manu. (D. Wilson
and A. Sandoval, eds.) Editorial
Horizonte, Lima.
Dallmeier, F., J. A. Comiskey and
F. Scatena.1998. Five years of
forest dynamics following Hurricane Hugo in Puerto Rico's
Luquillo Experimental Forest.
Pages 231-248 in Forest
Biodiversity in North, Central and
South America and the Caribbean:
Research and Monitoring. Man and
the Biosphere Series, Vol. 21.

(F. Dallmeier and J. A. Comiskey,
eds.) Parthenon Press, Carnforth,
Lancashire, UK.
Dallmeier, F., and J. A. Comiskey
(eds). 1998. Forest Biodiversity
Research, Monitoring and Modelling: Conceptual Background and
Old World Case Studies. Man and
the Biosphere Series, Vol. 20.
Parthenon Press, Carnforth,
Lancashire, UK..
Davis, M. B. 1976. Pleistocene
biogeography of temperate deciduous forests. Geosciences and Man
13:13-26.
Davis, M. B. 1986. Climatic instability, time lags, and community
disequilibrium. Pages 262-284 in
Community Ecology. (J. M. Diamond and T. C. Case, eds.) Harper
and Row, New York.
Diamond, J. M., and T. J. Case, eds.
1986. Community Ecology. Harper
and Row, New York.
Ecological Society of America.
1996. The report of the Ecological
Society of America committee on
the scientific basis for ecosystem
management. Ecological Applications
6(3): 665-691.
ERM. 1996. Camisea Appraisal
Drilling Campaign, Environmental
Impact Assessment. ERM Per S.
A., Lima.
Gentry, A. H. 1988. Changes in
plant community diversity and
floristic composition on environmental and geographical gradients.
Annals of the Missouri Botanical
Garden 75: 1-34.
Gentry, A. H. 1992. Tropical forest
biodiversity: distributional patterns
and their conservational significance, Oikos 63: 19-28.
Goldsmith, F. B. 1991. Vegetation
monitoring. Pages 77-86 in Monitoring for Conservation and Ecol-
39
ogy. (F. B. Goldsmith, ed.). Conservation Biology Series 3,

Chapman and Hall, London.
Hellawell, J. M. 1991. Development
of a rationale for monitoring.
Pages 1-14 in Monitoring for
Conservation and Ecology. (F. B.
Goldsmith, ed.) Conservation
Biology Series 3, Chapman and
Hall, London.
Hilborn, R. 1992. Can fisheries
learn from experience? Fisheries
17(4): 6-14.
Holling, C. S. 1978. Adaptive
Environmental Assessment and
Management. John Wiley and Sons,
New York.
Hubbell, S. P., and R. Foster. 1992.
Short-term population dynamics of
neotropical forest: why ecological
research matters to tropical conservation and management. Oikos 63:
48-61.
National Academy of Sciences.
1990. Managing Troubled Waters:
The Role of Marine Environmental Monitoring. Natural Research
Council Committee on a Systems
Assessment of Marine Environmental Pollution Monitoring,
National Academy Press, Washington, DC.
ONERN. 1987. Inventario y
Evaluacin de los Recursos Naturales del Medio y Bajo Urubamba,
Departamento del Cusco
(reconocimiento). Oficina
Nacional de Evaluacin de
Recursos Naturales, San Isidro,
Lima.
ONERN. 1990. Estudio
Semidetallado de Suelos y
Forestales del Curso Medio y Bajo
Urubamba, Departamento del
Cusco. Oficina Nacional de
Evaluacin de Recursos Naturales,
San Isidro, Lima.
Spellerberg, I. F. 1992. Evaluation
and Assessment for Conservation.
40
Conservation Biology Series 4,

Chapman and Hall, London.
Stohlgren, T. J. 1994. Planning longterm vegetation studies at landscape scales. Pages 209-241 in
Ecological Time Series. (J. H.
Steele and T. M. Powell, eds.)
Chapman and Hall, New York.
Stohlgren, T. J., D. Binkley, T. T.
Veblen, and W. L. Baker. 1995.
Attributes of reliable long-term
landscape-scale studies: malpractice insurance for landscape ecologists. Environmental Monitoring and
Assessment 36: 1-25.
Stohlgren, T. J., G. W. Chong, M. A.
Kalkhan, and L. D. Schell. In press.
Rapid assessment of plant diversity patterns: a methodology for
landscape. Environmental Monitoring.
Stork, N. E. 1995. Measuring and
inventorying arthropod diversity in
temperate and tropical forests.
Pages 257-270 in Measuring and
Monitoring Biodiversity in Tropical
and Temperate Forests. (T. J. B.
Boyle and B. Boontawee, eds.)
Center for International Forestry
Research, Bogor.
Tessler, S. 1995. Inventory and
Monitoring Program Instructions
for Data Handling. Shenandoah
National Park, U.S. National Park
Service, Washington, DC.
UNEP. 1995. Global Biodiversity
Assessment. United Nations
Environment Programme, Cambridge University Press, Cambridge.
Usher, M. B. 1991. Scientific requirements of a monitoring
programme. Pages 15-32 in Monitoring for Conservation and Ecology. (F. B. Goldsmith, ed.) Conservation Biology Series 3, Chapman
and Hall, London.
Walters, C. J. 1986. Adaptive
Management of Renewable Resources. McGraw Hill, New York.
Assessment
of Biologicial
Diversity and
Long-term
Monitoring
Plan for
the Lower
Urubamba
Region
Thomas Stohlgren and
Geneva Chong
Midcontinent Ecological Science
Center, Biological Resources
Division, U.S. Geological Survey,
Introduction
The Lower Urubamba project
involves an assessment of
biodiversity in a large, pristine
tropical forest and monitoring over
time to detect potential changes in
biological communities resulting
from local disturbances caused by
constructing and operating wells for
natural gas and condensate evaluation operated by Shell Prospecting
and Development (Per) B.V.
Normally, only a small portion of
any given landscape can be monitored because of cost constraints,

but scientist and project managers
can make valid inferences about
larger, unsampled landscapes. It is
important to identify areas of high
biodiversity (i.e. hot spots) with
distinct species composition for
protection. It is equally important to
identify areas of relatively low
diversity in large areas for potential
development.
The location of long-term
monitoring sites should be based on:
1) a better understanding of the
spatial distributions of species,
vegetation types, and habitat types;
2) spatially explicit models of biotic
and abiotic components and processes; and 3) quantitative information to determine how representative the long-term plots are compared with other plots and the
larger, unsampled landscape. Monitoring programs normally benefit
from a preceding systematic assessment of biotic and abiotic resources
(Stohlgren et al. In press a,b).
Revised protocols for vegetation
sampling, renewed economic incen-
41
We present
a conceptual design
for an
assessment
of biological diversity
and a
long-term
monitoring
plan for the
Lower
Urubamba.
42
tives to improve monitoring programs, and recent advances in the

design of ecological assessments
have made it easier to improve the
design of integrated monitoring
programs.
We present a conceptual design
for an assessment of biological
diversity and a long-term monitoring plan for the Lower Urubamba. It
takes advantage of the latest multiscale sampling designs and technologies proven in the recent, peerreviewed scientific literature. The
proposed design has three phases:
landscape-scale assessment, establishment of monitoring sites, and
long-term monitoring.
Landscape-scale Assessment
Landscape-scale assessments
begin by using remotely sensed data
to stratify a site into relatively
homogeneous ecological land units
(Kalkhan et al. 1995). Randomly
selected sampling sites in each
stratum allow for an unbiased
assessment of biodiversity hot
spots and identify habitats that are
less sensitive to land-use change
because of high redundancy (i.e.,
where the same habitat is abundant
nearby). Similar to the findings by
the Oficina Nacional de Evaluacin
de Recursos Naturales of Per
(ONERN 1990), eight ecological
land unit types were identified at
the Lower Urubamba site. These
appeared to differ in topography
and drainage patterns (based on
elevation contour maps) and color
and texture (based on satellite
imagery). These sites are likely
different in vegetation composition,
successional stage, soils, and site
history.
Sampling sites were randomly
selected at San Martin-3 and
Cashiriari-2. Unbiased site selection
adds tremendous power in extrapolating to the larger, unsampled

landscape (Stohlgren et al. In press
a,c) (Ferreira and Stohlgren In
press). Because so little is known
about the landscape-scale spatial
patterns of biodiversity in the study
region, determining the number of
sample sites needed will be an
iterative process. We anticipate that
5 to 10 sites will be needed per
stratum, but this depends on the
variation in species composition
and other factors found in the early
stages of the assessment. If withinstrata variation is low, 5 to 10 sites/
stratum may be enough. If withinstratum variation is high, more sites
will be needed. This iterative
process will result in a powerful,
well-replicated survey of diversity
patterns (Stohlgren et al. In press
a,c).
At each of the sampling sites, a
team of flora ecologists assessed
the biodiversity. A 20 m x 50 m
(1000 m 2) vegetation plot was
established at each sampling site.
Nested plots of .5 m x 2 m (1 m 2;
subplot a) and 2 m x 5 m (10 m2;
subplot b) and 5 m x 20 m (100m 2,
subplot c) within the 1000 m2 plot
(d) allow the development of
species-area curves to estimate the
number of species in a larger,
unsampled area and a means to
evaluate biodiversity patterns
(Stohlgren et al. 1995, Stohlgren et
al. In press a; Fig. 1). In the three
smaller subplots, each tree was
measured (diameter at breast height
and height) by species, and in the
larger plot, the number of new
species encountered was also
recorded and tallied. The larger plot
was also used by animal ecologists
to assess the biodiversity of selected groups of species (e.g., birds,
some insect groups).
N
2
20 m
20
.5
5
5
2
W
50 m
Figure 1. Multi-scale vegetation sampling design for tropical forests (modified from Stohlgren et al. 1995).
The data are then analyzed at
three spatial scales: the plot scale,
the strata scale, and the landscape
scale (entire study area). Spatial
analyses (trend surface maps,
kriging maps, and co-kriging maps)
of plant and animal species richness
are then used to reveal the patterns
in diversity. Species composition
overlap within and among strata
show where unique species and
common assemblages occur in the
landscape. Species-area relationships are used to estimate the total
plant species richness on the landscape (Stohlgren et al. In press c).
Information on plant and animal
species distributions provides a
sound basis for selecting representative long-term monitoring sites.
Monitoring Site
Establishment
To extrapolate information
from plots to landscapes with
known levels of accuracy and
precision, it is vitally important to
know how representative long-term
monitoring sites are compared to
the broader landscape. Long-term
monitoring sites are selected to
represent the landscape of concern.
In the Lower Urubamba, the

long-term monitoring plots are
being established near the drilling
sites to study local changes in
biodiversity Here, the internationally accepted SI/MAB design (oneha plot) is followed for monitoring
site-specific changes in tree demography and changes in plant and
animal species diversity (Dallmeier
1992, Dallmeier et al. 1992). Additional long-term monitoring plots
will be needed in other strata to
evaluate changes in biodiversity at
landscape scales. Decisions concerning the number of plots needed
are based on information gained
from the biological assessment.
The long-term monitoring
phase includes periodic remeasurement of the monitoring plots. The
time between measurements may
vary. A few plots may be assessed
yearly at first to evaluate annual
variability. Overall, however, 3 and
10 year intervals may be adequate
for remeasurement. Periodically, it
may be important to return to a few
of the assessment plots to assess
landscape-level changes for some
43
indicator species. For example, a

rare species of tree may die out in a
long-term monitoring plot, but the
same species might be abundant in
another assessment plot. It is
important to be able to factor out
local extinction, which is a very
common phenomenon, from
landscape-scale extinction, which is
a very rare event.
References
Dallmeier, F. (ed.). 1992. Long-term

Monitoring of Biological Diversity
in Tropical Forest Areas: Methods
For Establishment and Inventory
of Permanent Plots. MAB Digest
11. United Nations Educational,
Scientific, and Cultural Organization (UNESCO), Paris.
Dallmeier, F., C. M. Taylor, J. C.
Mayne, M. Kabel, and R. Rice.
1992. Effects of Hurricane Hugo
on the Bisley biodiversity plot,
Luquillo Biosphere Reserve,
Puerto Rico. Pages 47-72 in Longterm Monitoring of Biological
Diversity in Tropical Forest Areas:
Methods For Establishment and
Inventory of Permanent Plots.
MAB Digest 11. (F. Dallmeier ed.).
United Nations Educational,
Ferreira, L.V., and T. J. Stohlgren. In
press. Effects of river level fluctuation in species richness, diversity, and plant distribution in a
floodplain in Central Amazonia.
Oecologia.
Kalkhan, M. A., T. J. Stohlgren, and
M. Coughenour. 1995. An investigation of biodiversity and landscape-scale gap patterns using
double sampling: a GIS approach.
Pages 708-712 in Proceedings of
the Ninth Conference on Geo-
44
graphic Information Systems.

Vancouver.
ONERN. 1990. Estudio
Semidetallado de Suelos y
Forestales del Curso Medio y Bajo
Urubamba, Departamento del
Cusco. Oficina Nacional de
Evaluacin de Recursos Naturales,
San Isidro, Lima.
Stohlgren, T. J., M. B. Falkner, and
L. D. Schell. 1995. A modifiedWhittaker nested vegetation
sampling method. Vegetatio 117:
113-121.
Kalkhan, and L. D. Schell. In press
a. Rapid assessment of plant
diversity patterns: A methodology
for landscapes. Ecological Monitoring
and Assessment.
Stohlgren, T. J., M. B. Coughenour,
G. W. Chong, D. Binkley, M.
Kalkhan, L. D. Schell, D. Buckley,
and J. Berry. In press b. Landscape
analysis of plant diversity. Landscape Ecology.
c. Multi-scale sampling of plant
diversity: effects of the minimum
mapping unit. Ecological Applications.
Flora I:
Biodiversity
Assessment
in the Lower
Urubamba
Region
Pedro Acevedo, Deborah Bell,
Katherine Rankin, and
Stephen Smith
Department of Botany,
History, Smithsonian Institution
Introduction
The authors comprised a team

of four botanists from the U.S.
National Herbarium at Smithsonian
Institution, who participated in a
four-week expedition to the Lower
Urubamba and the gas well sites of
(Per) B.V. We were helped by two
local guides, who assisted in searching for and sampling plants, opening
trails, and transporting equipment
and samples. They also helped build
rudimentary laboratory facilities.
The purpose of our trip was to
survey the vascular plant flora in
the vicinity of the well sites. This
entailed sampling trees, epiphytes,
lianas, herbs, and ferns that were
found in a fertile condition (i.e.,

flowering or fruiting).
Methods
To document the floristic

diversity of the area, we sampled
specimens of every species that
appeared to be different in each
distinct habitat. We explored the
area using existing trails and by
opening new trails. We began
sampling activities as early as
possible each day and usually
returned to camp by mid-afternoon
to process specimens until the
electrical generators were turned
off, usually by 10 o'clock at night.
This schedule varied according to
weather conditions.
Our sampling gear included a
set of 36-foot telescoping pole
clippers, hand-clippers, plastic bags,
a plant press, and tree climbing
irons (the irons were rarely used).
We walked along and off the trails
in search of fertile plant specimens.
The team members worked together
to locate and sample specimens and
record data. Trees, lianas, and
epiphytes were sampled by climbing
or with telescoping pruning poles.
45
We also surveyed the trees being

cut down by the construction crew
at the well sites and sampled specimens using the same procedure
described above. Delicate plants
were pressed at the site of sampling; the more sturdy material was
bagged for later pressing at the
camp. Notes describing the plants
were made in the field. Later in the
evening, we transcribed them into a
data-base file using a laptop
computer.
The samples were pressed
between newspaper, using standard
methods, and assigned a number
according to the field notes. Each
sample (a distinctive number)
contained from one to 10 duplicates. The samples were assembled
in bundles, placed in plastic bags,
saturated with 55% ethanol, and
sealed for transit and later drying.
The materials were marked and
bagged separately at camp for each
participating institution. Plastic
bags containing the prepared specimens were placed in rice bags for
added protection, tagged, and sent
by helicopter to the base camp at
Nuevo Mundo. Bulky fruits were
prepared separately. They were
wrapped in newspaper and saturated with a 70% ethanol solution.
Relatively small, bulky fruits were
left whole, while large fruits were
sliced. A few very fleshy fruits were
injected by syringe with 70% alcohol.
At Nuevo Mundo, arrangements were made to send the
specimens by plane to Lima. The
specimens were picked up at the
Lima airport and taken to the
Museum of Natural History. The
samples will be distributed to the
participating institutions as follows:
one set is designated for the
46
Marcos, Lima, including any

unicates; another set will be housed
at the Smithsonian Institution' s
National Herbarium; and the third
set will be deposited at the
Cusco.
Sampling Sites
1. San Martin-3.
We sampled this site from
January 11-26 and January 29February 4, 1997 (sampling days =
17 for a total of 542 samples; P.
Acevedo et al. #8581-9064, 90829133, and 9287-9292). The topography of San Martin-3 is hilly with
steep to gentle slopes, dissected
with numerous small creeks (GPS
11 4709.8"S, 72 4205.3"W;
elevation of 487 m). For the most
part, the soil is deep and clayish,
with very few rock outcrops. The
area was extremely difficult to walk
because of the steep hills and the
abundance of dense, spiny bamboo
stands (Guadua sarcocarpa, known
locally as "pacales"). In addition to
the bamboo, the area has an open
forest cover, which can be characterized as non-flooded and wet.
Physiognomically, it is difficult to
characterize the vegetation under
different types. It is clear, however,
that the main difference is the
relative abundance of bamboo.
Bamboo is dominant on the gentle
slopes and less abundant on very
steep areas such as riverbeds and
creek drainages. The area contains
very few large trees, a relatively
abundant understory layer, and
numerous epiphytes and lianas.
The first few days, we concentrated our sampling efforts along
the trail that leads from the eastern
side of the camp to the stream.
During the following days, we
explored the area west of the camp
in anticipation of the drilling

platform construction. There, we
followed all the existing trails and
established new ones to gain access
to different trees and forested areas.
After a few days, we explored the
southern limits of the workers camp
by opening new trails. Similar
procedures were followed along the
northern limits of the camp.
We thoroughly explored the
area and took representative
samples. Still, we could not sample
a few species of trees and lianas
because of their inaccessibility and
the rough terrain. Sampling during a
different season of the year will add
more species to the checklist.
2. Malvinas.
Dr. Acevedo took a few
samples during a two-hour layover
at Malvinas on his way to Nuevo
Mundo in January 24, 1997 (17
collections; P. Acevedo #90659081). Although the area is highly
disturbed and mostly covered with
pasture or agriculture fields, there
were a few remnant vegetation
spots with plant species that had
not been sampled on previous trips
to the region.
3. Cashiriari-2.
We sampled this site January
27-28, 1997 (153 collections; P.
Acevedo et al. #9134-9286). The
topography of the Cashiriari-2 is
hilly with rather gentle slopes,
dissected with a few small creeks
(GPS 11 5151.3"S, and 72
4645.6"W, elevation 469 m). The
soil for the most part is deep, and
we did not observe any rocky
outcrops. The area surrounding the
workers camp was covered with a
primary, wet, closed forest (no
bamboo) with numerous representatives of different life forms such as
trees, herbs, epiphytes, and lianas.

We sampled in the surrounding
areas and along an old service road
that leads to water pumps.
The flora of this site was
somewhat different from that at San
Martin-3. In particular, the frequency of the species seems to be
different (i.e., species found to be
common at San Martin-3 were
rather rare at this site). In addition,
we discovered numerous species
that we had not sampled on previous trips to the region. We did not
anticipate spending more than one
night at Cashiriari-2 because we
believed that this site was not
significantly different from San
Martin-3. Therefore, we took no
alcohol for processing samples on
site and had to fly back to San
Martin-3 the next day to process the
samples properly.
Results
The checklist below was produced based on specimens sampled.

Families and generic determinations
were done in the field by Pedro
Acevedo and Stephen Smith. Further determinations to species were
done by Stephen Smith or by different plant specialists, as indicated on
the checklist.
Not all samples have been
identified to date. A few species
appear to be new to science, including one species in the Acanthaceae
family and two terrestrial ferns.
Other records of interest are a
possibly new color form of Episcia
fimbriata and a species of Phinaea,
relatively rare to Per, with few
samples. In the Heliconiaceae
family, two of the Heliconia species
sampled seem to occur rarely in
Per.
47
Appendix 1. List of the plant species of the Lower Urubamba region,

Department of Cusco, Per. Numbers represent collection numbers; numbers in parentheses represent number of duplicates; names in parentheses
after family name indicate persons responsible for identifying specimens.
DIVISION BRYOPHYTA
(by H. Robinson)
Lepidopilum polytrichoides (Hedw.) Brid.
8932(2)
Leucodontopsis geniculata (Mitt.) Crum
8975(2)
Sematophyllum subpinnatum (Brid.)
Britton 8814(2)
DIVISION PTERYDOPHYTA
(by D. Lellinger)
9083(1);8643(3); 9027(1);
8782(12); 8894(1); 9006(2);
8600(2); 8772(2); 8783(1);
8802(?);
Adiantum 8994(1)
Adiantum 9264(1)
Adiantum platyphyllum (Sw.) 8924(3)
Adiantum tetraphyllum (Willd.)8690(6)
Alsophila cuspidata (Kunze) Conant
8615(10)
Asplenium 8673 (?)
Asplenium 8768(2)
Asplenium 9029(1)
Asplenium auriculatum (Sw.) 8651(2),
8906(3)
Asplenium auritum Sw. 8748(2)
Asplenium pearcei (Baker) 8707(2),
8771(2)
Asplenium serratum L. 8631(6)
Campyloneurum asplundii (C.Chr.) Ching
8769(8)
Cyclopeltis semicordata (Sw.) J. Sm. 8735(6)
Diplazium bombonasea (Rosenth.)
8904(2), 8989(8), 8992(3),
9030(2)
Diplazium grandifolium 8671(4), 9266(2)
Diplazium grandifolium var. andicola
(Stolze)8830(5), 8612(5)
Diplazium pinnatifidum 8840(2)
Elaphoglossum blepharoglottis (Miekel)
9008(2)
Lastreopsis effusa ssp. Divergens (Willd.)
Findale 8732(4)
Lomariopsis fendleri (DC.) Eaton 9108(3)
Lomariopsis japurensis (Mart.) J. Sm.
8710(5)
48
Microgramma fuscopunctata (Hook.)

Vareschi 8652(4), 8597(2)
Pleopeltis percussa (Cav.) Hook. & Grev.
8896(2); 9124 (13)
Polypodium 8987(1)
Polypodium decamanam (Willd.) J. Sm.
8653(4)
Polypodium caeseresii (Sod.) 8691(4)
Pteris haenkeana 8733(4)
Selaginella chrysoleuca Spring 8991(9)
Selaginella erythropus (Mart.) Hieron.
8997(2)
Selaginella exaltata (Kunze) Spring
8780(9)
Selaginella haematodes (Kunze) Spring
8877(2); 8996 (5)
Tectaria incisa (Cav.) 8695(4), 9031(2)
Tectaria 9032(1)
Tectaria pilosa (Fee) Moran 8711(3)
Tectaria plantaginea (Jacq.) Maxon
9046(9)
Thelypteris 8895(1)
Thelypteris 9076(1)
Thelypteris angustifolia (Willd.) Proctor
8990(2)
Thelypteris biformata (Rosenst.) Tryon
8763(2)
Thelylpteris glandulosa var. brachyodus
(Kinze) A.R.Sm. 9118(3),
9235(4)
Thelypteris jamesonii (Hook.) Tryon
8988(9), 8993(4)
Thelypteris rufa (Poir) A.R.Sm. 9172(3),
8998(2)
Thelypteris tristis (Kunze) Tryon 8862(4),
8980(2)
DIVISION ANGYOSPERMAE
ACANTHACEAE
(by D. Wasshausen)
Aphelandra aurantiaca (Scheidew.) Lindl.
9010(7)
Aphelandra maculata (Nees) A. Voss.
8675(5)
Justicia 8880(1)
Justicia 9103(9)
Mendoncia 8897(2)
Appendix 1. Plant species list of the Lower Urubamba region(Cont.).

Mendoncia glabra (RuRz & Pav\n) Nees
8965(6), 8784(7), 9140 (3)
Stenostephanos 9130(2)
Stenostephanos longistaminus (RuRz &
Pav\n) V.M. 9071(3)
AMARANTHACEAE
Chamissoa altissima (Jacq.) Kunth var.
rubella Suess. 8667(6); 8787(8)
AMARYLLIDIACEAE
Bomarea 8614(1)
Bomarea 9065(1)
ANACARDIACEAE
Tapirira myriantha Triana & Planch.
8951(9); 8972(16)
ANNONACEAE
9227(5); 9132(1); 8635(10);
9101(4); 8973(10); 9082(5);
8849(2)
Annona 8852(3)
Guatteria 8705(3)
Guatteria 8887(1)
Guatteria 8888(2)
APOCYNACEAE
8959(9); 8941(3); 8778(6)
Mandevilla 9256(5)
Prestonia 9254(8)
Prestonia 9258(12)
Tabernaemontana 8610(6)
ARACEAE
(by T. Croat) 8692(4); 8801(1)
Anthurium 8647(1)
Anthurium 9026(1)
Anthurium 9170(1)
Anthurium breviscapum Kunth 8694(5)
Anthurium clavigerum Poepp. 8659(4)
Anthurium croatii Madison 8749(3);
8716(2)
Anthurium gracile (Rudge) Schott8817(4)
Chlorospatha 8836(5)
Chlorospatha 8890(4)
Dieffenbachia 8902(4)
Dieffenbachia 8969(3)
Filarum manserichense Nicolson 8717(9)

Monstera lechleriana Schott 8781(2)
Philodendron 8825(11)
Philodendron campii Croat 8914(5)
Syngonium podophyllum Schott 9243(2)
Syngonium yurimaguense Engler 8706(2)
Xanthosoma purpurata K. Krause8737(7)
ARALIACEAE
Dendropanax williamsii (Harms) Harms
9251(13)
Schefflera sprucei (Seem.) Harms 9057(6)
ARECACEAE
8665(7); 8672(1); 9289(?);
8601(1); 8898(1); 8756(1);
8773(1)
Bactris 8660(1)
Chamaedorea ? 8921(1)
Iriartea 9088(10)
ASCLEPIADACEAE
8811(5)
ASTERACEAE
(by H. Robinson & J. Pruski)
9135(1); 9078(1); 9186(1)
Dasyphyllum cf. brasiliense (Spreng.)
Cabrera8658(3)
Eclipta alba (L.) L. 9239(3)
Eirmocephala megaphylla (Hieron) H.Rob.
9159(4)
Mikania hookeriana DC. 8878(4);
8967(8)
Mikania simpsonii Holmes & McDaniel
9147(5)
Piptocarpha lechleri 9123(3)
Wulffia baccata (L.) Kuntze 9075(7);
9143(6)
BALANOPHORACEAE
Helosis cayennensis Spreng. 9000(1)
BEGONIACEAE
Begonia wollnyi? 8682(5)
Begonia glabra Aubl. 8633(7)
Begonia maynensis A.DC. 9050(7)
BIGNONIACEAE
8948(10)
49
Appendix 1. Plant species list of the Lower Urubamba region (Cont.).

BOMBACACEAE
Matisia cordata 8920(7)
BORAGINACEAE
9012(2)
Cordia lomatoloba I.M. Johnston 8856(8)
Cordia nodosa Lam. 9192(3)
Cordia ucayaliensis I.M. Johnst. 8625(8)
Heliotropium indicum L. 9233(3)
BROMELIACEAE 8834(1); 9081(2);
9007(1)
Aechmea angustifolia Poepp. & Endl.
8953(2); 8650(4)
BURSERACEAE
Protium 8882(8); 9044(6)
CACTACEAE
Epiphyllum phyllanthus 8818(1)
CAMPALUNACEAE
(by B. Stein)
Centropogon 9137(1)
Centropogon capitatus Drake 8613a(4)
Centropogon granulosus Presl 9134(5);
9056(2)
Centropogon roseus Rusby 8613b (4)
CAPPARIDACEAE
Capparis prisca Macbr. 8688(4)
Cleome spinosa Jacq. 9230(5)
CARICACEAE
Carica 8808(2)
Carica glandulosa (A.DC.) Solms-Laub.
8981(2)
Carica microcarpa Jacq. Ssp. heterophylla
(Poepp. & Endlch.) Badillo
8944(10)
Jacaratia digitata (Poepp. & Endl.) SolmsLaub. 8889(9)
CLUSIACEAE
(by J. Pipoly)
Chrysochlamys weberbaueri Engler
8727(6); 8939(7); 9247(4)
Clusia flavida (Benth.) Pipoly 9187(5)
Clusia hammeliana Pipoly 8835(10)
50
Tovomita brasiliensis Walp. 9201(9)

Vismia baccifera (L.) Triana & Planch.
9156(5)
Vismia cayennensis (Jacq.) Pers. 9174(5)
COMBRATACEAE
Terminalia oblonga (RuRz & Pav\n)
Steudl. 9085(9)
COMMELINACEAE
8799(3); 8798(3); 8947(3);
9279(1); 8930(2); 8758(?);
8755(2)
Dichorisandra 8893(8)
Floscopa 8754(4)
CONVOLVULACEAE
Dicranostyles 8837 (11)
COSTACEAE
Costus 8581(3)
Costus 8622(2)
Costus 8699(4)
Costus 9182(2)
Costus 9190(1)
Costus 9218(3)
CUCURBITACEAE
(by M. Nee)
Calycophysum pedunculatum Karst. &
Triana 8918(10)
Cayaponia cruegeri (Naud.) Cogn. 8907(5)
Cayaponia glandulosa (Poepp. & Endl.)
Cogn. 8964(10)
Cucumis 9113(1)
Fevillea 8812(5)
Gurania eriantha (Poepp. & Endl.) Cogn.
8679(5); 8931(2)
Gurania spinulosa (Poepp. & Endl.)
Cogn. 8654(9); 8704(6)
CYCLANTHACEAE
9036(3)
Dicranopigium 8775(6)
Carludovica palmata RuRz & Pav\n
9021(2); 9145(2)
Cyclanthus bipartitus Poit. 8719(2);
9146(2)

Evodianthus funifer (Poit.) Lindm.
8731(5); 9196(2)
CYPERACEAE (by M.T. Strong)
Cyperus laxus L. 8999(3)
Cyperus odoratus L. 9241(3)
Fimbristylis dichotoma (L.) Vahl 9240(3)
Pleurostachys 8846(5)
Rhynchospora umbraticola Poepp. & Kunth
8670(6); 9183(2)
Scleria huberi C.B. Clarke 9005(6)
DICHAPETALACEAE
Stephanopodium peruvianum Poepp. &
Endlch. 8766(9); 8934(10)
DILLENIACEAE
9217(1)
DIOSCOREACEAE
Dioscorea piperifolia H. & B. ex Willd.
8986(3)
EBANACEAE
Diospyros tessmanii Mildbr. 8606(9)
ELAEOCARPACEAE
Sloanea fragrans Rusby 8839(8)
Sloanea spathulata C.E. Smith 8854(12)
ERYTHROXYLACEAE
Erythroxylum macrophyllum Cav. 9288(3)
EUPHORBIACEAE
9270(5)
Acalypha 8599(4)
Acalypha 9189(1)
Acalypha cf. benensis Britton 9068(10);
9073(5); 9077(4)
Croton sampatik Muel. Arg. 8911 (11)
Drypetes 8793(4)
Drypetes 8797(10)
Manihot leptophylla Pax & K. Hoffm.
8816(11); 9177(4)
Omphalea diandra L. 8883(4); 9249(3)
Plukenetia volubilis L. 9139(2)
Sapium marmieri Huber 8958 12
FABACEAE
Acacia loretensis Macbr. 8617(11)
Andira multistipula Ducke 8655(3)

Crotalaria incana L. 9242(3)
Desmodium axillare (Sw.) DC. 9136(3)
Dioclea 9157 (6)
Erythrina edulis Triana 8955 (3)
Inga densiflora Benth. 9086(9)
Inga edulis Mart. 9272(5)
Inga tenuistipula Ducke 9261(3)
Pithecellobium 9250(1)
Senna herzogi (Harms) I. & Barneby
8677(4); 9055(2); 9162(2)
Senna ruiziana (G.Don) I. & Barneby
9141(10)
Swartzia arborescens (Aubl.) Pittier
8794a(2)
Swartzia simplex (Sw.) Spreng. var.
ochnacea Cowan 8794b(2)
Zapoteca amazonica (Benth.) H. Herndz.
9151(3)
Zygia juruana (Harms.) L. Rico 8730(6)
Zygia obolingoides L. Rico 8790 (6)
FLACOURTIACEAE
Banara nitida Spruce ex Benth. 9212
(10)
Casearia 9198(6)
Casearia decandra Jacq. 8915(10);
8630(6)
Casearia pitumba Sleumer 9199(10);
8979(4); 8892(9)
Lunania parviflora Spruce ex Benth.
9232(8); 8742(5); 8666(10)
Mayna grandifolia (Krst.) Warb. 8590(7)
Mayna odorata Aubl. 8709(5)
Tetrathylacium macrophyllum Poepp.
9286(8)
GENTIACEAE
Voyria 8638(2)
GESNERACEAE (L. Skog)
Besleria 8584(3)
Besleria 8712(5)
Besleria 8744(4)
Besleria 9273(3)
Codonanthe uleana Fritsch. 8605(3)
Diastema 8683(3)
Diastema racemiferum Benth. 9051(2)
Drymonia 9052(6)
Drymonia coccinea? 8783b (5)
51

Drymonia pendula (Poepp.) Wiehl
8596(6)
Drymonia warzewicziana Hanst. 9033(2);
9100(2)
Kohleria? 8616(2)
Monopyle 8685
Nautilocalyx 8821(3)
Nautilocalyx 9290(1)
HAEMADORACEAE
Xiphidium caeruleum Aubl. 8823(9)
HELICONIACEAE (by J. Kress)
Heliconia 8734(1)
Heliconia 9049(1)
Heliconia 9114(1)
Heliconia 9185(2)
Heliconia aemygdiana Burle-Max
8586(12); 9259(6)
Heliconia episcopalis Vell. 8585(7)
Heliconia rostrata RuRz & Pav\n
8587(2); 9219(6)
Heliconia spathocircinata Aristeg. 8795(5);
9195(6)
Heliconia subulata RuRz & Pav\n
9142(1)
HERNANDIACEAE
Sparattanthelium 8962(3)
HIPPOCRATEACEAE
9094(3)
Salacia 8850(3)
Salacia 9084(11)
Salacia 8645(2)
ICACINACEAE
8646(10)
Citronella melliodora (Sleumer) Howard
8627(10); 8696(10); 9202(6)
Leretia cordata Vell. 8982(3); 9215(8)
LAURACEAE
8917(9); 8977(9); 8804(5)
Aneuria 8874(6)
Nectandra 8807(10)
Nectandra 9025(9)
Ocotea 8656(5)
Ocotea 8657(7)
Ocotea 8697(2)
52
Persea 8806(8)
LECYTHIDACEAE
Grias peruviana Miers 9213(5)
LILIACEAE
Smilax 9283(1)
LOGONIACEAE
Potalia amara Aubl. 9064(2)
Strychnos panurensis Sprague & Sandw.
8736(10); 8868(3); 9110(10);
9148(3)
Strychnos tarapotensis Sprague & Sandw.
8786(5); 9188(2)
LORANTHACEAE
(J. Kuijt) 8604
Oryctanthus alveolatus (Kunth) Kuijt
8884(6)
Phoradendron woodsonii Trelease 8910(5)
Phthirusa pyrifolia (Kunth) Eichl.
8626(8)
MALPHIGIACEAE
9039(9); 8847(10); 9268(2)
Bunchosia 9226(4); 8796(10)
Mascagnia 9116 12
MALVACEAE
8689(1)
Pavonia fruticosa (Mill.) Fawcett &
Rendle 8978(5)
Pavonia leucantha Garcke 8698(10)
MARANTACEAE
9092(1); 9171(2); 9133(1)
Calathea 8623(2)
Calathea 8676(6)
Calathea 8708(5)
Calathea 8785(5)
Calathea 8819(2)
Calathea 9035(3)
Calathea 9138(1)
Calathea 9208(2)
Calathea 9209(2)
Ctenanthe ericae 8624(4)
MARCGRAVIACEAE
Souroubea 8843(10)

MELASTOMATACEAE
(by J.J. Wurdack)9150(3);
9154(1); 8903(5)
Clidemia dentata D.Don 9158(6)
Clidemia heterophylla (Desv.) Gleason
8743(5); 9015(3)
Clidemia septuplinervia Cogn. 9155(3)
Clidemia serpens (Triana) Cogn. 9109(5)
Clidemia sprucei Gleason 8636 (4)
Leandra 8591(5)
Leandra 8740(8)
Leandra dichotoma (Don) Cogn. 8620(4);
9281(?); 9181(3); 9168(2)
Maieta guianensis Aubl. 9009(8)
Miconia 8777(5)
Miconia 8885(6)
Miconia 9165(5)
Miconia 9175(5)
Miconia aurea (D.Don) Naudin
8983(12); 8632(5)
Miconia cf. filamentosa Gleason 8668(6);
9163(6)
Miconia cf. poeppigii Triana 9169(6)
Miconia dimorphica Macbride 9011(10)
Miconia muricata (D.Don) Triana 8984(6)
Miconia triplinervis RuRz & Pav\n ssp.
exalluvia Wurdack 9269(6);
9062(9); 8841(5); 8923(7)
Tococa caquetana Sprage 9107(3); 9047(3)
MELIACEAE
Cabralea canjerana (Vell.) Mart. ssp.
canjerana 9117(11)
Guarea 8909(1)
Guarea 9131(2)
Guarea kunthiana A. Juss. 9096(5)
Guarea macrophylla Vahl ssp. pachycarpa
(C.DC.) Pennington 8640(5)
Guarea macrophylla Vahl ssp. tuberculata
(Vell.) Pennington 8913(2)
Guarea pterorachis Harms 9265(2)
Trichilia 8687(1)
Trichilia elegans A. Juss. ssp. elegans
8582(6)
Trichilia elsae Harms 8942(3)
MENISPERMACEAE 8961(1)
Abuta 8848(3)
Abuta 9037(1)
Chondodendron tomentosum RuRz &

Pav\n 8872(4)
Disciphania convolvulacea Diels 9001(4);
9234(4)
Odontocarya tripetala Diels 9080(2);
9102(3); 9105(3)
MONIMIACEAE
Mollinedia 9098(1)
Siparuna cervicornis Perkins 8663(9)
Siparuna heteropoda Perkins 8753(7)
Siparuna thecaphora (Poepp. & Endl.) A.
DC. 8602 (6?); 9204(6)
MORACEAE
(by Berg)
Cecropia ficifolia Snethlag 9285(3)
Clavisia racemosa RuRz & Pav\n 8949(2)
Ficus maxima Miller 9041(2)
Ficus paraensis (Miq.) Miq. 8869(7);
8928(4)
Naucleopsis pseudonaga (Mildbr.) C.C.
9248(8)
Perebea guianensis Aubl. 8583(9);
9038(3); 9014(7)
Perebea humilis C.C. Berg 8940(3)
Trophis caucana (Pittier) C.C. 8833(10)
MYRISTICACEAE
Virola calophylla Warb. 9126 (20)
MYRSINACEAE
Ardisia nigrovirens Macbr. 9287 (9)
Cybianthuspoeppigii Mez 9099(3)
Parathesisadenanthera (Miq.) Hook.f. ex
Mez 8759(7)
Stylogyne cauliflora (Mart. & Miq.) Mez
8916(3); 9104(5)
MYRTACEAE
9024(3); 8648(10); 8765(1)
Calyptranthes longifolia Berg. 8715(8);
Eugenia 8608(10)
Eugenia macrocalyx (Rusby) McVaugh
9244(5)
Myrcia 8871(2)
Myrcia splendens (Sw.) DC. 8779(10)
NYCTAGINACEAE
Neea 8792(1)
53

Neea laxa Poepp. & Endl. 8876(2);
9040(3)
Neea longipedunculata Britt. 8870(3);
9277(2)
Neea oppositifolia RuRz & Pav\n9072(3)
OCHNACEAE
Ouratea pendula Engler 8815(8)
OLACACEAE
Heisteria acuminata (Humb & Bonpl.)
Engle 8886(7)
ONAGRACEAE
Ludwigia 9253(3)
Ludwigia 9252(3)
ORCHIDACEAE
8681(1); 9034(1); 8746(1);
8956(1)
Epidendron 8822(1)
Masdavalia? 8680(1)
OXALIDACEAE
Biophytum 9002(2)
Biophytum soukupii Lourt. 9275(3);
8684(8)
PASSIFLORACEAE
(by C. Feuillet)
Dilkea 8925(1)
Dilkea 9214(1)
Passiflora 9069(1)
Passiflora 9203(1)
Passiflora menispermifolia Kunth 8751(3)
PIPERACEAE
Peperomia 9017(1)
Peperomia 9053(1)
Peperomia alata RuRz & Pav\n 8813(6)
Peperomia ellsworthii Trelease & Yunker
9054(3)
Peperomia filiformis RuRz & Pav\n
8724(5)
Peperomia macrostachya (Vahl) A. Dietr.
9121(3)
Peperomia mercedana C.DC. 9087(4)
Peperomia rotundifolia (L.) Kunth 8943(3)
Peperomia serpens (Sw.) Lou. 8594(6);
8899(3); 8974(3)
54
Piper
Piper
Piper
Piper
acreanum C.DC. 8875(4)

aduncum L. 9144(5); 9160(5)
aleyreanum C.DC. 8832(7)
angustum Rudge 8855(3); 8935(4);
9045(4); 9210(4)
Piper anonifolium (Kunth) C.DC. 8938(5)
Piper costatum C.DC. 8933(2)
Piper dumosum Rudge 8726(5); 8714 (5);
8927(7); 8929(2)
Piper hermannii Trealese & Yunker
9016(5); 9128(2)
Piper hispidum Sw. 8957(5); 9149(5)
Piper longestylosum C.DC. 8721(3)
Piper reticulatum L. 8701(4)
Piper secundum RuRz & Pav\n 8713(8)
Piper subsilvestre C.DC. 8728(10)
Piper udisilvestre C.DC. 8589(2);
9060(4); 9063(6); 9211 (10)
POACEAE
8995(9); 9228(3); 8946(?)
Guadua sarcocarpa LondoZo & eterson
8609(9)
Olyra 8595(4)
Olyra 8858(3)
Pariana 9280(2)
Pariana 9028(2)
Pariana 8644(1)
Pharus lappulaceus Aubl. 8853(6)
Setaria 9267(4)
RANUNCULACEAE
Clematis guadeloupae Pers. 9070(6)
RUBIACEAE
9229(1); 9095(1)
Cephaelis flaviflora 8800(7)
Chimarrhis glabriflora Ducke 8954(21)
Coussarea sessilifolia Standl. 8598(13)
Coussarea? 8926(8)
Duroia longifolia (Poepp. & Endl.) K.
Schum. 8621(4)
Faramea anisocalyx Poepp. & Endl.
8729(5); 9106(5)
Faramea capillipes Muell. Arg. 9043(2);
9257(3)
Faramea glandulosa Poepp. & Endl.
9023(6)

Faramea multiflora A.Rich. var. maynensis
(Spruce ex Rusby) Steyerm.
8619(5); 9216(4); 8641(3);
8678(2)
Gonzalagunia cornifolia (Kunth) Standl.
9074(3)
Hamelia axillaris Sw. 8851(7); 8905(8)
Isertia laevis (Triana) Boon 9167(3)
Ixora 9197(1)
Ixora killipii Standl. 9125(3)
Macrocnemum roseum (RuRz & Pav\n)
Wedd. 9194(8)
Manettia divaricataWernh 8879(10)
Palicourea guianensis Aubl. 9282(5);
9193(2); 8664(2); 9018(7)
Palicourea klugii Standl. 8703(7)
Palicourea macrobotrys (RuRz & Pav\n)
Roem. & Schult. 9176(6)
Palicourea subspicata Huber 8752(3)
Pentagonia spaticalyx K. Schum. 8662(8);
9207(4)
Pentagonia subauriculata Standl. 8642(8)
Posoqueria latifolia (Rudge) Roem. &
Schult. 8900 (11)
Psychotria 8603 (?)
Psychotria 8618(1)
Psychotria 8789(4)
Psychotria 8791(3)
Psychotria 8891(1)
Psychotria 9019(1)
Psychotria borucana (A.Medina) C.M.
Taylor & Burger 9274(2)
Psychotria brachybotria Muell. Arg.
8963(6)
Psychotria caerulea RuRz & Pav\n
8971(5); 9061(6)
Psychotria carthaginensis Jacq. 8693(8);
8661(2)
Psychotria ferreyrae C. M. Taylor 8767(7);
8873(6)
Psychotria flaviflora (Krause) C.M. Taylor
8637(5)
Psychotria hoffmannseggiana (Roem. &
Schult.)Muell. Arg. 8700(4)
Psychotria laxa RuRz & Pav\n 8824(7)
Psychotria macrophylla RuRz & Pav\n
9003(6)
Psychotria marginata Sw. 8649(9)
Psychotria nudiceps Standl. 9058(2)
Psychotria pilosa RuRz & Pav\n 9059(9)
Psychotria poeppigiana Muell. Arg. 8741(3)

Psychotria racemosa (Aubl.) Raeuschel
8776(3)
Psychotria rhodothamne Standl. 8922(3)
Psychotria tenuicaulis Krause 8762(3);
9180(3)
Psychotria tessmannii Standl. 9184(2)
Psychotria trivialis Rusby 8919(8)
Psychotria viridis RuRz & Pav\n
8936(10)
Rudgea 8629(2)
Rudgea 8881(4)
Rudgea retifolia Standl. ? 8827(4)
Sabicea villosa Willd. ex Roem & Schult.
8966(10)
RUTACEAE
9237(10)
SANTALACEAE
Acanthosyris asipape 8950(12)
SAPINDACEAE
(by P. Acevedo)
Allophylus leucophloeus Radlk. 8866(10)
Allophylus pilosus (Macbr.) Gentry
8863(8)
Paullinia 8805(6)
Paullinia alata (RuRz & Pav\n) Don
8770(4); 9220(3)
Paullinia baileyi Standl. 9093(3)
Paullinia bilobata Radlk. 9262(2)
Paullinia bracteosa Radlk. 8774(3);
9097(2); 9127(3);
9263(4)
Paullinia cf. elongata Radlk. 9231(7)
Paullinia dasystachya Radlk. 8686(9);
8828(6); 8860(6)
Paullinia imberbis Radlk. 9222(7)
Paullinia killipii Macbr. 9115(2)
Paullinia neglecta Radlk. 9221(8)
Paullinia obovata RuRz & Pav\n
9260(2)
Paullinia pachycarpa Benth. 8912(3)
Paullinia selenoptera Radlk. 8702(6)
Paullinia serjaniaefolia Triana & Planchon
8722(5)
Paullinia tenera Poepp. & Endlch.
8831(3)
55

Pseudima frutescens (Aubl.) Radlk.
8829(7); 8865(3); 9090(3)
Serjania deltoidea Radlk. 9200(3)
Serjania pyramidata Radlk. 9112(3)
Serjania rubicaulis Radlk. 9079(1);
9173(2)
Talisia cerasina (Benth.) Radlk. 8908(2)
Talisia peruviana Standl. 8844(4)
Thinouia obliqua RuRz & Pav\n 8788(3)
Solanum oppositifolium RuRz & Pav\n

9020(3); 9284(6); 8750(2);
8845(8);9152(4)
Solanum pensile Sendtn. 9255(5)
Solanum robustifrons Bitter 9042(3);
9066(2)
Solanum sessile Dunal 9067(2)
Solanum supranitidum Bitter 9164(7)
Solanum thelopodium Sendtn 8960(7)
SAPOTACEAE
Ecclinusa ramiflora Mart. 9205(5)
Micropholis egensis (A.DC.) Pierre
8976(5)
Pouteria 8592(6);
Pouteria 9276(6)
STERCULIACEAE
(by L. Dorr)
Byttneria catalpaefolia Jacq. 8968(15)
Byttneria pescapraefolia Britt. 8739(9)
Herrania 8842(3)
Sterculia 8952(9)
Theobroma cacao L. 8718(2)
SOLANACEAE
(by M. Nee in part) 9091(1)
Brunfelsia mire Monachino 8588(4)
Cestrum silvaticum Francey 9278(5);
9120(11); 8970(15)
Cuatresia forsteriana A.T. Hunz.8723(11)
Cyphomandra pendula 8761(6)
Cyphomandra hartwegii (Miers.) Walp.
9245(5)
Cyphomandra tenuisetosa Bitter 9224(8)
Hawkesiophyton ulei (Dummer) A.T.
Hunz. 8937(4)
Juanulloa parasitica RuRz & Pav\n
9048(3)
Lycianthes inaequilatera (Rusby) Bitter
8725(6)
Physalis angulata L. 9225(5)
Solanum 8720(2)
Solanum 8747(1)
Solanum 8760(1)
Solanum 8803(7)
Solanum 8809(1)
Solanum 8810(2)
Solanum barbeyanum Huber 8757(6)
Solanum compressibaccatum Bitter 8857(4)
Solanum grandiflorum RuRz & Pav\n
9238(5)
Solanum lepidotum Dunal 9129(4)
Solanum mite RuRz & Pav\n 8945(9);
8745(2)
Solanum nigrum Dunal 8820(1)
56
THEOPHRASTACEAE
Clavija reflexiflora Killip 9271(2);
9122(2)
Clavija tarapotana Mez. 9022(9)
THYMELIACEAE
Daphnopsis peruviensis (Domke) Macbr.
8867(6)
Schoenobiblus peruvianus Standl.
8628(7)
TILIACEAE
(by L. Dorr)
Apeiba tibourbou Aubl. 9153(6)
ULMACEAE
Trema micranthum (L.) Blume 9246(5)
URTICACEAE
Pilea bassleriana Killip 8985(6)
Urera baccifera (L.) Gaud. 9004(2);
9119(3)
Urera caracasana (Jacq.) Gaud. ex Griseb.
9166(2)
VERBENACEAE
Aegiphila haughtii Mold. 8838(7);
8611(8)
Aegiphila peruviana Turcz. 8864(2);
9111(6)
Aegiphila sordida Mold. 8593(3)

Citharexylum poeppigii var. calvescens Walp.
8764(10)
Lantana 9161(1)
VITACEAE
Cissus lehmannii Bur. ex Lombardi
9089(8)
VIOLACEAE
Gloeospermum sphaerocarpum Triana &
Planch. 8674(8)
Leonia crassa L.B. Smith & Fdez.
8607(2); 8859(8)
Leonia glycycarpa RuRz & Pav\n 9236(4)
Rinorea pubiflora (Benth.) Sprague &
Sandw. var. grandifolia (Eichl.)
Hekking 9206(2)
Rinorea viridifolia Rusby 8639(10);
8826(10)
ZINGIBERACEAE
Costus sp. 1
Costus sp. 2
Costus sp. 3
Costus lima Schumann
Hedichyum coronarium Koen.
Renealmia sp. 1
Renealmia sp. 2
57
58
Flora II:
Biodiversity
Assessment
in the Lower
Urubamba
Region
Percy Nez
Facultad de Ciencias Biologicas,
Abad del Cusco
Severo Balden and
Hamilton Beltrn
Mayor de San Marcos
Introduction
The flora of southeastern Per

comprises a large number of species
(Wilson and Sandoval 1996, Nez
and Chong 1996). The topographic
irregularities of the eastern side of
the Andes creates different microhabitats where local plants and
animals survive. A high plant
diversity is usually correlated with a
high diversity of the organisms that
depend on them. Most animal
species feed on the leaves, flowers,
and fruits that the plants produce.
The greater the variety of plants in
an area, the greater the number of
animals they are likely to support.
To date, most studies on the

floristics of the Urubamba region
have been done in the upper portion. As part of the biodiversity
region, we sampled the vegetation
near the well sites San Martin-3 and
Cashiriari-2. Our principal objective
was to document most of the plant
species found in the area in a
relative short period of time. We
took several samples of each
species as voucher specimens.
These have been deposited at the
herbaria of the Museo de Historia
Mayor de San Marcos, the
Cusco in Per, and the Smithsonian
Institution. The work will serve to
complement studies of the Upper
Urubamba region.
Methods
As noted elsewhere by Acevedo

et al. (this volume) vegetation at
San Martin-3 is characterized by an
upland tropical forest mixed with
bamboo (Guadua sarcocarpa subsp.
sarcocarpa). The vegetation surrounding the drilling site at
59
The
bamboo is
a fast growing grass
that uses
other
plants to
climb. As
part of the
forest dynamics, the
bamboo
stems
frequently
break and
thus affect
the trees
near them.
60
Cashiriari-2 is characterized as a
pristine primary rainforest. The
bamboo is a fast-growing grass that
uses other plants to climb. Its stems
are about 7 cm in diameter, and
they can reach up to 25 m in height.
As part of the forest dynamics, the
bamboo stems frequently break and
thus affect the trees near them. We
observed that more than 10% of
individuals in biodiversity plot #1
at San Martin-3 had their tops
broken (Alonso et al. this volume).
Succession and forest dynamics may
be suppressed or enhanced by the
presence of bamboos, which may
affect tree regeneration for many
years until they flower and die.
Bamboos only flower once. The
effect on the forest is evident by the
many tree species that are present
only as juveniles and saplings
(Londoo and Peterson 1991, Judd
1992).
Field work enabled us to become familiar with the tree flora in
the biodiversity plots, supplemented
with general sampling in the vicinity
of the well sites. We conducted
meter-to-meter ground-truthing
surveys as far as one days walk
from the well sites, taking advantage of the rivulets as ways to
access the forest because the bamboo makes it very difficult to enter
in some areas. For example, it took
us an entire afternoon to obtain one
undescribed species of Pleurothyrium, a plant species of the
avocado family (Lauraceae).
Plant identification requires
experience. We used binoculars (10
x 40) to obtain visual access to the
canopies and telescoping poles to
cut small, representatives branches
in attempting to obtain fertile
material (i.e., flowers and fruits).
We frequently climbed trees using
climbing irons and a security belt.
We were able to get excellent plant

material by finding the plant specimen of interest in the forest and
carefully removing a few samples
without damaging the tree. Samples
were taken to the field laboratory
and stored in individual sheets of
newspaper to dry the material and
arrange it properly for museum use.
Once we accumulated several
samples, we packaged them, placed
them in plastic bags, added 70%
ethanol as a preservative, and sent
the bags to the National Herbarium
of the Museo de Historia Natural,
Marcos. The specimens were then
dried in an oven for 16 hours at
65o C. Given our experience, we
were able to identify most of the
plants using plant characteristics of
the live specimens. Organoleptic
characters of the tree trunks were
of crucial importance in our task.
Since many trees had no leaves, we
used the exudates of the trunk, the
color of the bark, and the shape of
the buttresses to determine the
family, the genus, and often the
species. We will use the plant
material sampled to complete the
identifications (Rohwer 1993, Tryon
and Stolze 1994, Terborgh et al.
1996).
Results and
Discussion
Of the samples taken in the

area of study, we found several
genera with three or more species
(i.e., sympatric species). We also
found several subspecies (i.e.,
geographical and morphological
differences within same species).
We observed the following examples in the study area: Perebea
guianensis, with the two subspecies
P. guianensis subsp. 1 and P. guianenis
subsp. guianensis (Moraceae); Guarea

macrophylla, with the three subspecies G. macrophylla subsp. pachycarpa,
G. macrophylla subsp. pendulispica,
and G. macrophylla subsp. tuberculata
(Meliaceae); and Pouteria torta, with
the three subspecies P. torta subsp.
glabra, P. torta subsp. pubescens, and
P. torta subsp. torta (Sapotaceae).
The group of Pouteria is differentiated in the field by presenting
glabrous leaves, pubescences, and
the shape of the bole. These are
relatively well-known in the tropics.
Guarea kunthiana has three distinct
formsG. kunthiana subsp. 1 (small
leaves), G. kunthiana subsp. 2
(gigantic leaves), and G. kunthiana
subsp. 3 (type 3, Meliaceae).
The flora of the Lower
Urubamba region is composed of a
mixture of species that are commonly found in other areas. For
example, the area of this study has
species typical of dry forests found
at altitudes of 700 m in Kiteni to
1100 m in Potrero, close to
Quillabamba, Department of Cusco,
Per. Botanical examples include
Astronium sp. 1 (undescribed,
Anacardiacaea), Bredemeyerea lucida
(Polygalaceae), Cybistax antisiphilitica
(Bignonaceae), Tabebuia ochracea (1
individual per ha), T. roseo-alba, and
T. serratifolia (Bignocaceae). Species
typical of montane cloud forest
include Bryophytes mosses
(Octoblepharum sp. 1), Begonia
parvifolia (Begoniaceae), Cinchona
sp. 1, (30-m trees, with one to two
individuals per ha, Rubiaceae),
Cyathea spp., Trichipteris sp. 1 (three
species of tree ferns, 25 individuals
per ha), Elleanthus sp. 1, and
Sobralia sp. 1 (Orchidaceae). Species associated with streams and
river banks include Chimarrhis sp. 1
(Rubiaceae), Cordia lomatoloba
(Boraginaceae), Cyathea spp., and
Trichipteris sp. 1 (tree ferns),

Guatteria acutissima (Annonaceae),
Lonchocarpus spiciflorus (Fabaceae),
Marila laxiflora (Clusiaceae),
Pentagonia parvifolia (Rubiaceae),
Piper reticulatum (Piperaceae),
Siparuna sp. 1 (Monimiaceae), and
Talauma amazonica (Magnoliaceae).
Less than 10 % of the tree
species found in the area are deciduous (i.e., trees that drop their
leaves seasonally). The deciduous
species include Ceiba pentandra
(Bombacaceae), Chorisia insignis
(Bombacaceae), Erythrina
poeppigiana, E. ulei (Fabaceae),
Schizolobium parahybum (Fabaceae),
Tabebuia ochracea, and T. serratifolia
(Bignoniaceae).
Tree-fall gaps and forest clearings have an important role in
maintaining species diversity.
Species commonly found in open
areas include Cecropia sciadophylla
(Moraceae), Citharexylum poeppigii
var. calvescens (Verbenaceae), Cordia
alliodora (Boraginaceae), Cyathea
boliviana (tree fern), Hyeronima
laxiflora, H. oblonga (Euphorbiaceae),
Inga edulis, I. punctata (Fabaceae),
Jacaratia digitata (Caricaceae),
Myriocarpa stipitata (Urticaceae),
Pourouma cecropiaefolia (Moraceae),
Urera baccifera, U. caracasana, and U.
laciniata (Urticaceae).
Several of the species sampled
were only classified as
morphospecies (i.e., individuals that
share the same morphological
characters are grouped in the same
species, for example Guarea spp. and
Pouteria spp.). Several individuals of
the genus Inga spp. (Fabaceae) were
very difficult to classify. However,
with the existing comprehensive
database, literature, and collection
of photographs, the most common
species will be determined. Several
of the species of this genus have
The flora
of the
Lower
Urubamba
region is
composed
of a mixture of
species
that are
commonly
found in
other
areas.
61
not been described; several of these

are known only by sterile individuals. It has been estimated that there
are more than 150 species of Inga
spp. in Per.
There are several species of
hemi-hepiphytes (i.e., species that
germinate and develop in the trees
and reach the ground by sending
roots). They are mostly represented
by herbaceous aroids, ferns,
gesneriads, and marcgravias. The
woody flora is represented by Ficus
maxima (Moraceae) and Schefflera
morototoni (Araliaceae). Other
examples include well-known
strangler trees such as Coussapoa
ovalifolia, C. villosa (Cecropiaceae),
Ficus juruensis, F. paraensis, F.
sphenophylla, and F. trigona
(Moraceae).
Palm trees dominate several
landscapes in southern Per. The
genus Iriartea spp. is a dominant tree
at the area of this study and in the
Rio Manu area. Palm species include Aiphanes aculeata, Iriartea
deltoidea (one of the most dominant)
and Socratea exhorriza (one individual in 1 ha). Other genera are
absent or represented by a single
individual, including the climbing
genera Desmoncus polyacanthos var.
polyacanthos (climbing palm).
Nez, P., and G. Chong. 1996.

Flora of the Camisea, Rio
Urubamba. In Proceedings from
the Workshop on Biological and
Cultural Diversity of the Lower
Urubamba, Per. Biodiversity
Programs, Smithsonian Institution,
Washington, DC.
Pitman, N., Terborgh, J., and P.
Nez V. In press. Tree rarity in
the upper Amazon.
Rohwer, J. 1993. A revision of
Nectandra (Lauraceae). Flora
Neotropica Monograph.
Terborgh, J., R. B. Foster, and P.
Nez V. 1996. Tropical tree
communities: A test of the
nonequilibrium hypothesis. Ecology
77: 561-567.
Tryon , R., and R. Stolze. 19891994. The ferns of Per. Fieldiana
Botany New series (Chicago).
Wilson, D., and A. Sandoval. 1996.
Manu: The Biodiversity of Southeastern Per. Editorial Horizonte/
Smithsonian Institution Press,
Lima.
References
Judd, S. 1992. Systematics of the

Guadua angustifolia complex
(Poaceae:Bambusoideae). Annals
of the Missouri Botanical Garden 79:
737-769.
Londoo, X., and P. Peterson. 1991.
Guadua sarcocarpa (Poaceae:
Bambuseae) a new species of
Amazonian bamboo with fleshy
fruits. Systematic Botany 16:
630-638.
62
Appendix 1. Flora of the upper and Lower Urubamba region of southeastern Per. The area covered is approximately 200 km 2.
DIVISION BRYOPHYTA
Marchantia chenopoda (18999)
DIVISION PTERYDOPHYTA
Adiantum sp. 1
Adiantum sp. 2
Adiantum sp. 3
Adiantum sp. 4
Adiantum latifolium aff. Lam. (10175)
Adiantum petiolatum Desv. (10253)
Adiantum poeppigianum Presl=
Phylitidis J.Sm.
Adiantum raddianum C.Presl. (10154)
Antrophyum lineatum ( Sw.) Kaulf.
(18198)
Asplenium sp. 1
Asplenium sp. 2
Asplenium sp. 3
Asplenium cristatum Brack (10134)
Asplenium dimidiatum Sw,
Asplenium kunzeanum
Asplenium quitense Hooker CB (1752)
Asplenium serra Lansd. & Fisch.
Asplenium ruizianum klotzsch? (10195)
Blechnum divergens (Kunze) Mett.
Bolbitis semipinnatifida (Fee) Alston
(10271)
Bolbitis serrata (Kuhn) Ching (12685)
Bolbitis serratifolia (Mert ex Kaulf.)
Schott (10224)
Campyloneurum angustipaleatum
(Alston) M.Mey ex Lellinger
(10199)
Campyloneurum nitidissimum (Mett.)
Ching var. abruptum (10243,
12750)
Ctenitis nigrovenia ( H. Christ) Copel
(12760)
Cyathea microdontha (Desv.) Domin
(12776)
Cyathea (small, tiny scales )
Cyathea (big scales 19200)
Cyclopeltis semicordta (Sw.) J. Sm.
Dryopteris patens
Dydimochlaena truncatula
Grammitis andicola Stolze
Grammitis blepharidea (Copel.) Stolze
Lomariopsis latipinna
Lomariopsis nicotianifolia
Lomariopsis nigropaleata
Microgramma reptans
Lycopodium (18284) sp. 1
Pecluma plumula
Polybotrya fractiserialis
Polypodium camptophyllarium var.
abreviatum
Polypodium furfuraceum
Pteris denticulata
Pteris petiolulata
Pityrogramma austroamericana
Pityrogramma calomelanos
Selaginella sp. 1
Selaginella sp. 2
Selaginella flavellata
Solanopteris bifrons (Hook.) Copel.
Tectaria draconoptera
Tectaria incisa
Thelypteris opulenta
Thelypteris sp. 1
Trachypteris pinnata
DIVISION GYMNOSPERMAE
CYCADACEAE
Zamia sp. 1 (12692, 19600)
GNETACEAE
Gnetum sp. 1
DIVISION ANGYOSPERMAE
ACANTHACEAE
Aphelandra (12813) sp. 1
Aphelandra sp. 2
Aphelandra pulcherrima (Jacq.)Kunth
(10120)
Dicliptera sp. 1
Encephalosphaera sp.
Fittonia alvivennis
Hansteinia sp. 1
Hygrophila sp. 1
Juruasia appendiculata
Justicia sp. 1
Justicia sp. 2
Justicia peruviana Lam. (12912)
Justicia rusbyana Lindau. V. A. W.
Wassell
Kalbreyeriella
Pachystachys (12947) sp. 1
Pachystachys coccinea
63
Appendix 1. Flora of the upper and Lower Urubamba region of southeastern Per. The area covered is approximately 200 km 2 (Cont.).
Pachystachys ossolae
Pseuderanthemum sp. 1 (12943)
Pseuderanthemum sp. 2
Mendoncia sp. 1
Mendoncia sp. 2
Mendoncia sp. 3
Mendoncia sp. 4
Mendoncia lindavii Rusby (10248)
Pulchranthus sp. 1
Ruellia sp. 1
Ruellia sp. 2
Ruellia sp. 3
Ruellia sp. 4
Ruellia sp. 5
Ruellia brevifolia ( Pohl) C.Ezcurra
(10101)
Ruellia geminiflora
Ruellia tarapotana
Stenostephanus thyrsoides
Sanchezia oblonga R. & P.
Sanchezia ovata Ruiz & Pav. (12923)
Sanchezia sp. 1
Suessenguthia vargasii Wasshausen
Thunbergia alata Bojer ex Sims
(10238, 12939)
ALISMATACEAE
Echinodorus sp. 1
AMARANTHACEAE
Alternanthera sp. 1
Amaranthus caudatus var. sanguinensis
Amaranthus spinosus L.
Chamissoa altissima HBK.
Cyathula achyranthoides (HBK) Moq.
Iresine sp. 1
Iresine diffusa H. B. ex Willd.
Pfaffia sp. 1
AMARYLLIDACEAE
Bomarea sp. 1
Bomarea amoena M.Roem. (10095)
Brunsvigia sp. 1
Crinum sp. 1
Eucharis sp. 1
Eucharis cyanosperma A.W. Meerow
Furcraea andina Trel.
Hippeastrum sp. 1
64
ANACARDIACEAE
Anacardium occidentale L.
Astronium sp. 1
Astronium graveolans
Mangifera indica L.
Mauria aff. heterophylla Kunth (10150)
Rhus juglandifolia Wall. ex D.Don
Spondias lutea Royen
Spondias mombin Jacq.
Tapirira guianensis Aubl.
Tapirira peckoltiana
ANNONACEAE
Anaxagorea pachypetala (Liels) R.E.
Fries
Anaxagorea sp. 1
Annona montana
Annona mucosa Jacq.
Crematosperma (10146) sp. 1
Crematosperma sp. 2 (manu)
Crematosperma sp. 3
Crematosperma sp. 4 (19374)
Cymbopetalum sp. 1
Cymbopetalum sp. 2
Duguetia (12880) sp. 1
Duguetia sp. 2
Duguetia sp. 3
Ephedranthus guianensis
Fusaea sp. 1
Guatteria sp. 1
Guatteria sp. 2
Guatteria sp. 3
Guatteria sp. 4
Guatteria sp. 5 (19313)
Guatteria acutissima
Guatteria dielsiana
Guatteria guentheri Diels (18185)
Guatteria microcarpa
Guatteria tomentosa Rusby (12848)
Malmea sp. 1
Malmea cauliflora
Malmea diclina
Oxandra sp. 1
Oxandra sp. 2
Porcelia nitidifolia R. & P.
Rollinia sp. 1
Rollinia sp. 2
Rollinia sp. 3
Rollinia cuspidata C. Martius (10123)
Appendix 1. Flora of the upper and Lower Urubamba region of southeastern Per. The area covered is approximately 200 km (Cont.).
Rollinia edulis
Rollinia mucosa (Jacq.)Baill (12855)
Rollinia pittieri
Rollinia ulei Diels
Ruizodendron sp.
Tryginaea sp.
Unonopsis sp. 1
Unonopsis sp. 2
Unonopsis floribunda Diels
Unonopsis mathewsii
Xylopia sp. 1
Xylopia sp. 2
Xylopia cuspidata
APIACEAE
Hydrocotile ranunculoides
APOCYNACEAE
Aspidosperma sp. 1 (10269)
Aspidosperma sp. 2 (19369)
Aspidosperma sp. 3 (19399, many
secondaries)
Aspidosperma megalocarpon Muell.
Arg.
Aspidosperma vargasii A.D.C.
Condylocarpon sp. 1
Forsteronia sp. 1
Forsteronia sp. 2
Himatanthus sucuuba (Spruce ex
Muell. Arg)
R.E.Woods.
Mandevilla callista Woodson (10108)
Mandevilla scabra (Hoffmannsegg ex
Roemer & Schultes)Schu (10194)
Mesechites trifida Muell.Arg.
Pacouria sp. 1
Prestonia sp. 1
Rauvolfia praecox
Secondatia sp. 1
Tabernaemontana sp. 1
Tabernaemontana sananho R. & P.
Thevetia peruviana Merrill
ARACEAE
Anthurium sp. 1
Anthurium sp. 2
Anthurium sp. 3
Anthurium sp. 4
Anthurium sp. 5
Anthurium sp. 6
Anthurium sp. 7
Anthurium clavigerum Poepp. (10168,
12733)
Anthurium croatii Madison
Anthurium gracile (Rudge) Schott
(10153, 12811)
Anthurium kunthii Poeppig (12752)
Anthurium pentaphyllum (Aubl.) G.
Don (10172)
Caladium bicolor (Ait.) Vent (10124)
Dieffenbachia sp. 1
Dieffenbachia sp. 2
Dieffenbachia sp. 3
Dieffenbachia sp. 4
Dracontium sp. 1
Dracontium sprucei
Heteropsis sp. 1
Heteropsis sp. 2
Homalomena sp. 1
Monstera sp. 1
Monstera sp. 2
Monstera sp. 3
Monstera sp. 4
Monstera lechleriana Schott (10171)
Monstera obliqua Mig.
Monstera pinnatifrons
Philodendron chanchamayense
Philodendron ernestii Engl.
Philodendron fibrillosum Poepp. &
Endl. (10185)
Philodendron latifolia (19116)
Philodendron undulatum Engl. (18125)
Pistia stratiotes L.
Rhodospatha sp. 1
Rhodospatha sp. 2
Spathiphyllum sp. 1
Stenospermation sp. 1
Syngonium sp. 1
Syngonium podophyllum Schott
(10169)
Syngonium yurimaguense Engl.
Xanthosoma sp. 1
Xanthosoma sp. 2
Xanthosoma pubescens Poepp. & Engl.
(18225)
65
ARALIACEAE
Dendropanax sp. 1
Schefflera morototoni (Aubl.) B.
Maguire
Gonolobus sp. 1
Matelea sp. 1
Sarcostemma sp. 1
Stenomeria sp. 1
ARECACEAE
Aiphanes aculeata (19349)
Aiphanes ernestii (Burret) Burret
Astrocaryum sp. 1
Astrocaryum chonta Mart.
Astrocaryum gratum
Bactris sp. 1
Bactris sp. 2
Bactris sp. 3
Bactris gasipaes HBK.
Chamaedorea angustisecta Burret
(10147)
Chamaedorea pinnatifrons Oerst.
Cocos nucifera L.
Desmoncus polycanthos (12938)
Euterpe precatoria Mart.
Geonoma sp. 1
Geonoma sp. 2
Geonoma jussieuana Mart.
Geonoma deversa Kunth
Hyospathe elegans Mart. (12697)
Iriartea deltoidea R. & P. Jessenia
bataua Barrret
Mauritia flexuosa L.f.
Oenocarpus mapora Karst.
Phytelephas macrocarpa R. & P.
Scheelea cephalotes Karst.
Socratea exorrhiza H. Wendl.
Socratea salazari H.E. Moore Syagrus
sancona
Wettinia augusta
ASTERACEAE
Acmella sp. 1
Adenostemma sp. 1
Adenostemma sp. 2
Ageratina sp 1.
Ageratum conyzoides Sieber ex Steud.
(12886)
Aspilia sp. 1
Baccharis salicifolia Nutt.
Baccharis trinervis Pers.
Bidens pilosa L.
Bidens squarrosa Less.
Clibadium vargasianum H. Robinson
Chromolaena sp. 1
Conyza sp. 1
Dasyphyllum sp. 1
Eclipta sp. 1
Eirmocephala cainarachiensis (Hieron)
H. Robinson (12915, 12918)
Elephantopus mollis HBK. (12885)
Erechtites hieracifolia Raf. ex DC.
Erechtites valerianaefolia DC.
Eupatorium crenulatum Gardn.
Eupatotium laevigatum Lam.
Eupatorium leptocephalum DC.
Fleischmannia microstemon (Cass.)R. M.
King & H. Rob. (12843)
Garcilassa sp. 1
Hebeclinium sp. 1
Ichthyothere sp. 1
Liabum acuminatum Rusby
Mikania sp. 1
Mikania sp. 2
Mikania cordifolia Willd.
Mikania guaco Humb. & Bonpl.
Mikania psilostachya DC. (10179,
12881)
Fulcaldea laurifolia Poir.
Gochnatia rusbyana Cabrera
Onoseris sp. 1
Oyedaea sp. 1
Piptocarpha poeppigiana Baker (18322)
Porophyllum ruderale Cass.
ARISTOLOCHIACEAE
Aristolochia (18293) sp. 1
Aristolochia sp. 2
Aristolochia (18338) sp. 3
Aristolochia sp. 4
Aristolochia iquitensis O.C.Schmidt
ASCLEPIADACEAE
Asclepias curassavica L.
Cynanchum sp. 1
Fischeria stellata (Vell. ) E. Fourn.
(10184)
66
Pseudelephantopus spicatus Rohr
Sonchus oleraceus L.
Struchium sp. 1
Tessaria integrifolia R.& P.
Trixix divaricata Spreng.
Vernonia cainarachensis Hieron.
Vernonia gracilis Poir.
Vernonia megaphylla Hieron.
Vernonia salzmannii DC.
Vernonia scorpioides Pers.
Viguiera sp. 1
Wulffia baccata (L.f.) Kuntze
BALANOPHORACEAE
Helosis cayennensis (Sw.)Spreng.
Ombrophytum sp. 1
Ombrophytum sp. 2
Ombrophytum peruvianum Poepp. &
Endl.
Ombrophytum violaceum B. Hansen
BASELLACEAE
Basellaceae sp. 1 (10210)
BEGONIACEAE
Begonia sp. 1
Begonia sp. 2
Begonia juninensis
Begonia parvifolia
Begonia patula Haworth
BIGNONIACEAE
Adenocalymma sp. 1
Adenocalymma sp. 2
Adenocalymma sp. 3
Adenocalymma sp. 4
Arrabidaea sp. 1
Arrabidaea sp. 2
Arrabidaea sp. 3
Arrabidaea corallina (Jacq.) Sandwith
Arrabidaea pearcei (Rusby) K. Schum.
ex Urb.(12840)
Arrabidaea poeppigii (A.DC.)Sandwith
Arrabidaea selloi (Spr. Eng.) Sandw.
(12717)
Callichlamys sp. 1
Ceratophyton sp. 1
Clytostoma sp. 1
Crescentia cujete Vell.
Cuspidaria emonsii A. Gentry (1284)

Cybistax antisyphilitica Mart.
Cydista aequinoctialis (L.) Miers (10167)
Cydista lilacina A. H. Gentry (12796,
12805)
Haplolophium nunezii A. Gentry
(12740)
Jacaranda copaia subsp. spectabilis
(Aubl.) D. Don
Jacaranda glabra
Lundia sp. 1
Macfadyena unguis-cati (L.) A. H. Gentry
(12803)
Mansoa sp. 1
Mansoa sp. 2
Martinella sp. 1
Musathia hyacinthina
Paragonia pyramidata (Rich.) Bureau
(12732)
Pithecoctenium crucigerum (L.) A. Gentry
Pleonotoma melioides(S. Moore) A. Gentry
(12823)
Spathicalyx xanthophylla (DC.) A. Gentry
(12799)
Sparattosperma sp. 1
Tabebuia capitata( Bur. & K. Schum)
Sandw.
Tabebuia ochracea Benth. ex Muell. Arg.
Tabebuia roseo-alba (Ridley) Sandw.
Tabebuia serratifolia (Vahl) G. Nicholson
(12910)
Tanaecium nocturnum Bur. & K. Schum
Tynanthus polyanthus (Bureau) Sandwith
(12758 , 12837, 12845)
Tynanthus schumannianus (Kuntze) A.H.
Gentry (12795)
Xylophragma pratense (Bur. & K. Schum.)
Spr.
BIXACEAE
Bixa orellana L.
Bixa platycarpa R. & P.
BOMBACACEAE
Cavanillesia umbellata R. & P.
Ceiba pentandra Gaertn.
Ceiba samauma (Mart.) K. Schumann
(10158)
Chorisia integrifolia
67
Chorisia insignis HBK.
Eriotheca globosa (Aubl.) A. Robyns
Matisia cordata Hum. & Bonpl.
Ochroma pyramidale
Quararibea wittii Schum & Ulbr. (10174)
BORAGINACEAE
Cordia alliodora Cham.
Cordia lomatoloba
Cordia nodosa Lam.
Cordia ucayalensis
Heliotropium sp. 1
Heliotropium indicum L.
Tournefortia sp. 1
Tournefortia sp. 2
Tournefortia sp. 3
BRAASSICACEAE
Rorippa clandestina (Spreng.) Macbride
BROMELIACEAE
Aechmea sp.1
Aechmea mertensii Schult. f.
Billbergia sp. 1
Billbergia cf. stenopetala Harms (10122)
Bromelia sp. 1
Bromelia sp. 2
Guzmania sp. 1
Neoregelia eleutheropetala
Pepinia tatzyanae H.Luther
Pitcairnia elongata L.B. Smith
Streptocalyx sp. 1
Tillandsia sp. 1
Tillandsia sp. 2
Tillandsia sp. 3
BURSERACEAE
Crepidospermum sp. 1
Protium puncticulatum
Protium tenuifolium Engl.
Trattinickia aspera
Tetragastris sp. 1
CACTACEAE
Disocactus sp. 1
Epiphyllum phyllanthus Haw.
Rhipsalis sp. 1
Selenicereus sp. 1
68
CAMPANULACEAE
Centropogon roseus Rusby (12913)
Centropogon sp. 1
Centropogon urubambae (19289, 19599)
CANNACEAE
Canna iridiflora R. & P.
CAPPARIDACEAE
Capparis flexuosa Vell. (12669)
Capparis macrophylla
Cleome aculeata L.
Cleome spinosa Sw.
Cleome viridifloraSchreb. (10109)
Podandrogyne sp. 1
CARICACEAE
Carica papaya L.
Carica microcarpa
Jacaratia digitata Solms. ( 10132)
CARYOCARACEAE
Anthodiscus klugii Standl. ex Prance
(18095)
Caryocar amygdaliforme
Caryocar pallidum
CARYOPHYLLACEAE
Drymaria
CELASTRACEAE
Maytenus magnoliifolia
COMBRETACEAE
Combretum
Combretum
Combretum fruticosum
Combretum laxum
Terminalia catappa
Terminalia oblonga
Thiloa glaucocarpa (10257)
COMMELINACEAE
Commelina longicaulis
Tradescantia zanonia
COCHLOSPERMACEAE
Cochlospermum orinocense
Appendix 1. Flora of the upper and Lower Urubamba region of southeastern Per. The area covered is approximately 200 km (Cont.).
CONNARACEAE
Connarus
Rourea sp. 1
CONVOLVULACEAE
Dicranostylis
Ipomoea aegyptia
Ipomoea coccinea
Ipomoea nil
Ipomoea phyllomega
Merremia macrocalyx
CUCURBITACEAE
Anguria sp. 1
Citrullussp. 1
Fevillea sp. 1
Gurania spinulosa (10246)
Melothria (12706) sp. 1
Psiguria bignoniacea R.P. Wunderlin
CRYSOBALANACEAE
Couepia latifolia
Hirtella racemosa
Hirtella triandra Swartz (18272)
(18105 ) sp.
Licania elata
Parinari parilis Macbride
CYCLANTHACEAE
Asplundia sp. 1
Asplundia peruviana Harling
Carludovica palmata Ruiz & Pav. (10126)
Cyclanthus bipartitus Poit.
Ludovia (18181) sp. 1
CYPERACEAE
Carex sp. 1
Cyperus ferax Benth.
Cyperus friburgensis Boeck. (10261)
Cyperus hermaphroditus Standley
Cyperus flavus Boeck.
Cyperus miliifolius Poepp. & Kunth ex
Kunt (10213)
Cyperus odoratus Osbeck
Cyperus regiomontanus Britton (12916)
Eleocharis sp.1
Fimbristylis dichotoma Vahl.
Rhynchospora umbraticola
Scleria (18187) sp. 1
DICHAPETALACEAE
Dichapetalum sp.
Tapura sp. 1
Tapura peruviana var. petioliflora
DILLENIACEAE
Curatella americana L.
Doliocarpus sp. 1
Doliocarpus dentatus
Tetracera parvifolia
DIOSCOREACEAE
Dioscorea sp. 1
Dioscorea sp. 2
Dioscorea acanthogene Rusby (12700)
Dioscorea decorticans C. Presl ( 12945)
Dioscorea samydea C. Mart. ex Griseb
(10241, 12836)
Dioscorea trifida (18928)
EBENACEAE
Diospyros sp. 1 (pubescent)
Diospyros subrotata
ELAEOCARPACEAE
Sloanea sp. 1
Sloanea sp. 2 (pubescent: 19344, 19355,
19476)
Sloanea fragans Rusby
Sloanea picapica
Sloanea sinemariensis
Muntingia calabura L.
ERYTHROXYLACEAE
Erythroxylum sp. 1
Erythroxylum sp. 2
Erythroxylum coca var. coca Lam.
Erythroxylum macrophyllum
Erythroxylum ulei O. E. Schultz
EUPHORBIACEAE
Acalypha sp. 1
Acalypha sp. 2
Acalypha benensis Britton ( 10161)
Acalypha cuneata Poepp. (10160, 12860,
12921)
Acalypha diversifolia Jacq. (10250)
Acalypha mapirensis Pax (12814)
Acalypha stricta Poepp. (10230)
69
Acalypha stenoloba Muell. Arg. (12920)
Acalypha villosa Jacq.( 12818)
Alchornea castaneaefolia A. Juss.
Alchornea glandulosa
Alchornea grandiflora Muell. Arg. (12834)
Alchornea triplinervia Muell. Arg.
Boconia alterna
Caryodendron orinocense
Cnidoscolus sp. 1
Conceveiba guianensis (19229)
Croton sp. 1
Croton sp. 2
Croton gracilipes Baill. (10228)
Croton matourensis
Croton lechleri Muell. Arg.
Croton tessmanniiMansfeld (18998)
Euphorbia hirta L.
Euphorbia heterophylla Desf.
Glycidendron amazonicum
Hevea brasiliensis Muell. Arg.
Hieronima laxiflora (18936)
Hura crepitans L.
Manihot brachyloba Muell. Arg. (12844)
Margaritaria nobilis L.f. (12856)
Mabea sp. 1
Mabea nitida
Pachystroma longifolium I. M. Johnst.
(10135, 12715)
Phyllanthus carolinianusBlanco
Phyllanthus lathyroides HBK.
Phyllanthus orbiculatus Rich.
Phyllanthus stipulatus (Raf.) G. Webster
(12873)
Ricinus communis L.
Sapium sp. 1
Sapium eglandulosum Ule
Sapium glandulosum Morong
Sapium ixiamasense Jabl.
Sapium laurifoium Griseb.
Sapium marmieri Huber
Senefeldera inclinata Muell. Arg.
FABACEAE
Acacia sp.1 (12673)
Acacia sp. 2
Acacia sp. 3
Amburana cearensis (Fr. Allem.) A.C.
Smith
Andira inermis HBK.
70
Andira surinamensis
Bauhinia guianensis Aubl.
Bauhinia tarapotensis
Bauhinia ungulata L.
Calliandra sp. 1
Calliandra glyphoxylon Spruce ex Benth.
Cassia sylvestris
Cedrelinga catenaeformis Ducke
Clitoria sp. 1
Clitoria sp. 2
Clitoria (18128) sp. 3
Copaifera reticulata
Coumarouma sp. 1
Cracca toxicaria Kuntze
Cratylia (Dioclea) argentea Kuntze
Crotalaria anagyroides HBK.
Crotalaria incana L.
Crotalaria nitens HBK.
Dalbergia sp. 1
Desmodium sp. 1 (18110)
Desmodium affine Schlecht.
Desmodium axillare DC.
Desmodium cajanifolium DC.
Desmodium tortuosum DC.
Dioclea sp.1 (19429)
Dioclea arborea
Dipteryx micrantha Harms (= D. odorata)
Erythrina
sp. 1
Erythrina poeppigiana Skeels
Erythrina ulei O.E. Schulz
Hymenaea sp. 1
Hymenaea sp. 2
Indigofera subfruticosa
Inga sp. 1 (19228)
Inga sp. 2 (19294, 19411)
Inga sp. 3 (19383)
Inga sp. 4 (19397)
Inga sp. 5 (little wings, 19198, 19339)
Inga sp. 6 (4-7 leaflts, pair)
Inga sp. 7 (6 leaflts, terete, 19438)
Inga sp. 8 (pubescent 7-8 leaflts, 19197)
Inga acrocephala
Inga adenophylla Pittier
Inga affinis Steud (10267)
Inga aliena Macbride
Inga chartacea (19296)
Inga cinnamomea
Inga edulis Mart. (18115)
Inga feullei DC. (10117)
Inga gracilies
Inga hirsuta G. Don
Inga macrobotrys Penn. (19258)
Inga macrophylla (19426)
Inga mathewsii Benth.
Inga pavoniana (19420)
Inga puctata (19219, 19358)
Inga quaternata
Inga ruiziana G. Don
Inga semialata Mart (18117)
Inga sertulifera (19359)
Inga setosa G. Don
Inga stipulacea
Inga stipularis (19352)
Inga thibaudiana (= I. peltadenia)
Inga umbellifera
Leucaena trichodes Benth.
Lonchocarpus sp. 1
Lonchocarpus sp. 2
Lonchocarpus spiciflorus Mart. ex Benth.
Machaerium sp. 1
Machaerium (10268) sp. 2
Myroxylum balsamum Harms (12770)
Mucuna sp. 1
Mucuna sp. 2
Mucuna rostrata Benth.
Ormosia sp. 1 (hairy, 19483)
Ormosia bopiensis
Ormosia coccinea Jacks.
Pachyrrhizus ajipa
Parkia nitida Miq.
Phaseolus adenanthus G.F.W. Mey
Piptadenia sp. 1
Piptadenia adiantoides var. peruviana
(18336)
Piptadenia colubrina Benth.
Piptadenia grata Macbride
Pithecellobium sp. 1
Pithecellobium longifolium (HBK.) HBK.
Standl.
Platypodium affinis cf. (12726)
Platymiscium sp. 1
Pueraria sp. 1
Phyllocarpus riedeliiTul.
Prosopis chilensis Stuntz
Pterocarpussp. 1
Pterocarpus rohrii
Rhynchosia sp. 1
Samanea saman (Jacq.)Merr. (12826)
Schizolobium parahybum Blake

Sclerolobium bracteosum
Senna sp. 1
Senna hirsuta var. hirta H. S.Irwin & R.
Barneby (12917)
Senna reticulata (Willd.) H.S. Irwin &
R.C. Barneby
Senna ruiziana (G.Don) H.S. Irwin &
R.C. Barneby
Senna spectabilis var. spectabillis (D.C.)
(12794)
Senna sylvestris (Vellozo) H.S. Irwin &
R.C. Barneby
Stryphnodendron sp. 1
Swartzia arborescens (18967, 19300)
Vatairea peruviana (19455)
Vigna repens Baker
Zygia latifolia
Zygia macrophylla (19398)
FLACOURTIACEAE
Banara guianensis Aubl.
Banara nitida
Carpotroche longifolia Benth.
Casearia decandra 19407
Casearia gossipiosperma Briquet (12702)
Casearia megacarpa Cuatr.
Casearia ulmifolia Cambess.
Hasseltia floribundaH.B.K.
Laetia procera Eichl. 18320
Lunania parviflora Spruce ex Benth
Mayna sp. 1
Mayna sp. 2
Mayna odorata Aubl.
Tetrathylacium macrophyllum Poepp. &
Endl.
Xylosma intermedium Griseb.
GENTIANACEAE
Escobedia scabrifolia R. & P.
Lisianthus alatus Aublet
Voyria sp. 1
GESNERIACEAE
Besleria sp. 1
Besleria sp. 2 (10237)
Besleria sp. 3
Besleria sp. 4
Codonanthe uleana Fritsch
71
Codonanthopsis sp.
Columnea sp. 1 (18090)
Columnea sp. 2
Columnea sp. 3
Drymonia penduliflora/pendula 18920
Drymonia semicordata (Poepp.) Wiehler
Episcia fimbriata
Gloxinia perennisDruce
Koellikeria erinoides (DC.) Mansf.
Phinaea sp. 1
GUTTIFERAE
Calophyllum brasiliense Camb.
Chrysoclamys ulei Engl.
Clusia sp. 1
Garcinia macrophylla
Decaphalangium peruvianum Melch.
Marila laxiflora Rusby 18196
Rheedia sp. 1
Rheedia sp. 2
Rheedia floribunda Planch. & Tr.
Symphonia globulifera L.f.
Vismia sp. 1
Vismia sp. 2
Vismia sp. 3
Vismia gracilis Hieron.
HAEMODORACEAE
Xiphidium coeruleum Aubl. (19705)
HERNANDIACEAE
Sparatanthelium tarapotanum
HIPPOCRATEACEAE
Anthodonsp. 1
Cheiloclinium cognatum
Hippocratea volubilis Sw.
Salacia gigantea
HUMIRIACEAE
Humiriastrum excelsa (Ducke) Cuatrec.
Vantanea sp. 1
72
Casimirella sp. 1 (19362)

Citronella incarum (19364)
Leretia sp. 1
IRIDACEAE
Eleutherine bulbosa Urb.
LABIATAE
Hyptis sp. 1
Hyptis spicigera Lam.
Hyptis mutabilis Brig.
Hyptis pectinata Poit.
Ocimum micranthum Willd.
Salvia occidentalis Sw.
LAURACEAE
Aniba sp. 1
Aniba guianensis (silvery below)
Aniba taubertiana Mez (12714)
Endlicheria sp. 1 (18102)
Endlicheria dysodantha (19323)
Endlicheria formosa (19435)
Mezilaurus sp. 1
Nectandra furcata Nees
Nectandra longifolia (19213)
Nectandra, terceary reticulate coriaceus
(19447)
Ocotea sp. 1
Ocotea comphomoroea (19322)
Persea americana Mill.
Persea gratissima Gaertn. f.
Pleurothyrium poeppigii (19400)
LECYTHIDACEAE
Cariniana sp. 1
Couroupitasp. 1
Eschweilera sp. 1
Eschweilera (camisea) sp. 2
Eschweilera (cumpirusiato) sp. 3
Grias peruviana
JUGLANDACEAE
Juglans boliviana
Juglans neotropica Diels
LILIACEAE
Smilax sp. 1
Smilax sp. 2
Smilax sp. 3
Veratrum candidum
ICACINACEAE
Calatola venezuelana Pittier (18254)
LOGANIACEAE
Potalia amara Aubl.
Strychnos sp. 1
Strychnos sp. 2
Strychnos tarapotensis Sprague & Sanwith
(10139, 12743)
LORANTHACEAE
Gaiadendron sp. 1
Phthirusa pyrifolia (Kunth) Eichler
(10116, 12810)
Psittacanthus sp. 1
Strutanthus sp. 1
LOASACEAE
Mentzelia aspera Cav.
LYTHRACEAE
Adenaria floribunda HBK.
Cuphea cordata R. & P.
Lafoensia punicifolia
MALVACEAE
Anoda cristata Schlecht.
Gossypium peruvianum Cav.
Hibiscus rosa- sinensis L.
Pavonia spinifex Cav.
Pavonia paniculata Cav.
Sida cordifolia Forsk.
Sida spinosa L.
Sida rhombifolia L.
MALPIGHIACEAE
Banisteria metallicolor A. Juss.
Banisteriopsis muricata (Cav.) Cuatr.
(10240)
Bunchosia glandulifera ( Jacq.) Kunth
(10155)
Byrsonima arthropoda A. Juss. (10206)
Heteropteris sp. 1
Hiraea sp. 1
Stigmaphyllon sp. 1
MAGNOLIACEAE
Talauma amazonica (18977, 19382)
MARANTHACEAE
Calathea sp. 1
Calathea comosa Lindl.
Calathea crotalifera Watson
Ctenanthe sp. 1
MARCGRAVIACEAE
Marcgravia sp. 1
Marcgravia crenata Poepp. ex Wittm.
MELASTOMATACEAE
Bellucia grossulariodes Trisna
Blakea mexiae Gleason
Graffenrieda (18250) sp. 1
Miconia sp. 1
Miconia sp. 2
Miconia sp. 3
Miconia sp. 4
Miconia sp. 5
Miconia sp. 6
Miconia sp. 7
Miconia sp. 8
Miconia bubalina -19477
Miconia calvescens DC
Miconia cf. cretacea Gleason (10197)
Miconia paleacea Cogn. (18127)
Miconia peruviana Naudini (12901)
Miconia persicariaefolia Cogn. ex Britton
Miconia procumbens (19181)
Miconia triplinervia
Tibouchina longifolia Baill.(18120)
Triolena amazonica
MELIACEAE
Cabralea canjerana (19230)
Cabralea canjerana subsp. canjerana
(19430)
Cedrela sp. 1
Cedrela fissilis Vell (10156, 12819)
Cedrela odorata R. & P.
Guarea sp. 1
Guarea sp. nov. (19341)
Guarea glabra Vahl (10205, 12709)
Guarea gomma Pulle (10159)
Guarea guidonia (L.) Sleum.
Guarea kunthiana A. Juss.
Guarea kunthiana (giant leaves, type 2)
Guarea kunthiana (type 3)
Guarea macrophylla (18992)
Guarea pterorachis Harms (18914)
Trichilia elegans subsp. Elegans
(10143, 12864)
Trichilia quadrijuga
Trichilia pallida
73
Trichilia pleeana (A. Juss.) C. DC.
(12680)
Trichilia poeppigiana
Trichilia septentrionalis
MENISPERMACEAE
Abuta sp. 1
Abuta grandifolia (Mart.) Sandwith
Anomospermum grandifolium Eichl.
(18173)
Borismene sp. 1
Chondodendron tomentosum
Curarea sp. 1 (12765)
Curarea sp. 2
Disciphania heterophylla R.C. Barneby
MONIMIACEAE
Mollinedia sp.
Siparuna sp. 1
Siparuna sp. 2
Siparuna sp. 3
Siparuna sp. 4
Siparuna sp. 5 (19199)
Siparuna crassifolia Perkins
Siparuna cuspidata
MORACEAE
Artocarpus incisa L.f.
Batocarpus amazonicus (Ducke) Fosberg
(18148)
Brosimum acutifolium
Brosimum alicastrum Sw. (10267)
Brosimum parinaroides Ducke (18212)
Brosimum utile (19440)
Castilloa ulei Warb.
Cecropia sp. 1
Cecropia sp. 2
Cecropia engleriana
Cecropia flagillifera Trec.
Cecropia membranacea Trec.
Cecropia polystachya Trecul (10121)
Cecropia sciadophylla Mart.
Cecropia tessmannii Mildbr.
Coussapoa sp. 1
Coussapoa villosa Poepp. & Endl.
Clarisia biflora R. & P.
Clarisia racemosa Ruiz & Pavon (10193,
12721)
Ficus sp. 1
74
Ficus sp. 2
Ficus sp. 3
Ficus sp. 4
Ficus insipida Willd.
Ficus gomelleira Hort. Monac. ex Kunth
& Bouche
Ficus macbridei
Ficus maxima
Ficus paraensis Miq.
Ficus sphenophylla
Helicostylis tomentosa
Maclura tinctorea D. Don (12725)
Maquira calophylla
Maquira coriacea (19301)
Maquira guianensis
Naucleopsis sp. 1
Naucleopsis glabra
Naucleopsis krukovii (Standl.) C.C. Berg.
Naucleopsis pseudonaga
Naucleopsis ternstroemifolia
(Mildbr.) C.C. Berg.
Olmedia aspera Poepp. & Endl.
(=Trophis caucana)
Perebea angustifolia
Perebea guianensis Aubl.
Perebea humilis
Perebea tesmannii
Pourouma cecropiaefolia Mart.
Pourouma cucura (golden hairy)
Pourouma guianensis
Pourouma mollis Trec.
Pourouma palmata Poepp. & Endl.
Poulsenia armata
Pseudolmedia laevigata
Pseudolmedia laevis (R. & P.) Macbride
Pseudolmedia macrophylla
Sorocea sp. 1
Sorocea guilleminiana Gaudich. (12677)
Sorocea pileata Burger
Sorocea steinbackii
MUSACEAE
Heliconia sp. 1
Heliconia sp. 2
Heliconia sp. 3
Heliconia sp. 4
Heliconia apparicioi H. de Souza
Barreiros
Heliconia episcopalis Vell.
Heliconia hirsuta L.f.
MYRISTICACEAE
Compsoneura capitellata Warb.
Iryanthera juruensis (19360)
Iryanthera laevis
Otoba glycicarpa
Otoba parvifolia
Virola sp. 1
Virola sp. 2
Virola calophylla Warb.
Virola duckei A.C. Smith
Virola flexuosa
Virola sebifera Aublet (10211)
MYRSINACEAE
Cybianthus (19469)
Myrsine sp. 1
Myrsine sp. 2
Stylogyne ambigua
Rapanea oligophylla Mez.
MYRTACEAE
Acca sp. 1
Calyptranthes sp. 1 (19390)
Calyptranthes longifolia
Eugenia sp. 1 (19220)
Eugenia sp. 2 (tiny-wavy leaves, 19433,
19443)
Eugenia L. (12722)
Myrcia sp. 1 (18177)
Myrcia sp. 2 (19335)
Myrcia sp. 3 (white below, 19439,
19446)
Psidium acutangulum (19353)
Psidium guajaba L.
Syzgium jambos (L.) Alston
NYGTAGINACEAE
Guapira sp. 1 (elliptic, 19342)
Neea sp. 1 (hairy, long ovobate, 19472)
Neea sp. 2 (19425)
Neea sp. 3 (19223, 19357)
Neea chlorantha
Neea hirsuta (19331)
Reichenbachia hirsuta Sprengel
(undescribed, related species?)
(10166)
NYMPHACEAE
Nymphaea sp. 1
OCHNACEAE
Godoya obovata R. & P.
Ouratea
Sauvagesia sp. 1
OLACACEAE
Heisteria sp. 1
Heisteria acuminata
Minquartia guianensis Aubl.
ONAGRACEAE
Ludwigia sp. 1
Ludwigia sp. 2
Ludwigia sp. 3
Ludwigia octovalvis (Jacq.) Raven
OPILIACEAE
Agonandra (12669)
Agonandra (19361)
ORCHIDACEAE
Bletia catenulata R. & P.
Cochleanthes amazonica (Reichb. f. &
Warsc.) R.E. (18954)
Epidendrum sp. 1
Epidendrum sp. 2
Epidendrum coronatum R. &P.
Epidendrum rigidum Schlechter
Gongora sp. 1
Huntleya sp. 1 (18949)
Macroclinium sp. 1
Maxillaria sp. 1
Ornithocephalus sp. 1 (12723)
Ponthieva sp. 1 (18263)
Polystachya sp. 1 (12877)
Polystachya sp. 2
Rodriguezia lanceolata
Spiranthinae sp. 1
Stanhopea hassleriana
Vanilla sp. 1
OXALIDACEAE
Biophytum sp. 1
Biophytum soukupii A. Lourteig
Oxalis spruceana Progel (10100)
75
PASSIFLORACEAE
Dilkea retusa (19183)
Passiflora sp. 1
Passiflora sp. 2
Passiflora auriculata kunth (12838)
Passiflora coccinea Blanco (18328)
Passiflora coriaceaJuss. (10110)
Passiflora edulis fo. Flavicarpa
Degener (10119)
Passiflora ligularis A. Juss.
Passiflora riparia Mart. ex Mast (10218)
PHYTOLACCACEAE
Gallesia integrifolia (Spreng.) Harms
(12768)
Hilleria latifolia H. Walter
Phytolacca rivinoides Kunth & Bouche
(12905)
PIPERACEAE
Peperomia sp. 1
Peperomia sp. 2
Peperomia macrostachya A. Dietr.
Peperomia quesita Trel.
Peperomia serpens C. DC.
Piper sp. 1
Piper sp. 2
Piper augustatum Rudge (10216)
Piper laevigatum HBK.
Piper reticulatum L.
POACEAE
Andropogon bicornis L.
Andropogon condensatus Kunth
Andropogon leucostachyus Kunth
Aristida torta (Nees) Kunth
Axonopus chrysoblepharis (Lag.) Chase
Axonopus scoparius (Flff) Kuhlm
Arundinella berteroniana (Schult.)
Hitchc. & Chase
Cynodon dactylon (L.) Pers.
Cenchrus echinatus L.
Cenchrus hillebrandianus Hitchc.
Chloris distichophylla Lag.
Chloris polydactyla (L.) Sw.
Chloris radiata ( L.) Sw.
Coix lacryma-jobi L.
Cryptanthus sp. 1
Digitaria sanguinalis (L.) Scop.
76
Digitaria violascens Link

Eragrostis ciliata
Eriochloa dystachya Kunth
Eriochloa punctata (L.) Desv.
Eleusine indica (L.) Gaertn.
Gouinia latifolia (Griseb.) Vasey
Guadua sp. 1
Guadua sp. 2
Guadua sarcocarpa subsp. sarcocarpa
Guadua weberbaueri Pilg.
Gynerium sagittatum var. sagittatum(Aubl.)
P. Beauv.
Gynerium sagittatum var. pubescens
Hackelochloa granularis (L.) Kuntze
Heteropogon contortus (L.) P. Beauv. ex
Ruem.
Hyparrhenia bracteata (Humb. & Bonpl.
ex Willd.) Stapf
Ichnanthus minarum (Nees) Dll
Ichnanthus panicoides P. Beauv.
Ichnanthus leiocarpus(Spreng) Kunth
(10214)
Imperata tenuis Hack.
Leptochloa dominguensis (Jacq.) Trin.
Leptochloa virgata (L.) P. beauv. (10128)
Lasiacis divaricata (Steud ex Lechler)
Hitch. (12909)
Lithachne pauciflora (Sw.) P. Beauv.
Melinis minutiflora P. Beauv.
Olyra sp. 1
Olyra fasciculata Trin.
Olyra latifolia L.
Oplismenus hirtellus (L.) P. Beauv.
Panicum capillare L.
Panicum maximum Jacq.
Panicum stoloniferum Poir (10254)
Panicum trichanthum Nees (12809)
Pariana (10270) sp. 1
Pariana zingeberina Rich. ex Doell
Paspalum cerecioides
Paspalum conjugatum Bergius
Pennisetum sp. 1
Phragmites australis (Cav.) Trin. ex Steud.
Pharus latifolius L. (10186)
Rhipidocladum sp. 1 (10208)
Setaria geniculata P. Beauv.
Setaria poiretiana( Schult.) Kunth
(12816)
Setaria sulcata Raddi
Setaria vulpiseta (Lam.) Roem & Schult.
Trachypogon montufari (Kunth) Nees
Trachypogon plumosus (Humb. & Bonpl.
ex Willd. Nees)
Trichachne insularis (L.) Nees
Tripsacum piosen
Urochloa brizantha (Hchstetter ex A.
Richard) R. Webster (10236)
POLYGALACEAE
Bredemeyera lucida
Moutabea sp.
Polygala sp. 1
Polygala sp. 2
Polygala sp. 3
POLYGONACEAE
Coccoloba sp. 1
Coccoloba (18165) sp. 2
Coccoloba (18218) sp. 3
Coccoloba mollis (19405)
Triplaris americana Rob. Schomb.
Triplaris pavonii Meissn.
Triplaris peruviana Fisch. & Mey ex C.A.
Mey
Triplaris poeppigiana Wedd.
Triplaris setosa Rusby
Triplaris vestita Rusby
PORTULACACEAE
Talinum paniculatum Jacq. (12940)
PROTEACEAE
Oreocallissp. 1
Roupala montana
QUIINACEAE
Lacunaria sp. 1
Quiina macrophylla (19129, 19409)
Quiina peruviana
RANUNCULACEAE
Clematissp. 1
RHAMNACEAE
Gouaniasp. 1
Gouania lupuloides Urb.
Rhamnidium elaeocarpum Reiss. (12730)
Ziziphus cinnamomum
ROSACEAE
Prunus sp. 1
RUBIACEAE
Rubiaceae (19337)
Bathysa sp. 1
Bathysa peruviana Krause
Borreria ocimoides DC
Calycophyllum acreanum
Calycophyllum spruceanum (Benth.) K.
Schum.
Capirona decorticans Spruce
Cephaelis tomentosa Vahl.
Chimarhis sp. 1
Cinchona sp. 1
Condaminea corymbosa DC
Condaminea microcarpa DC
Duroia sp. 1
Hamelia axillaris Sw.
Isertia laevis (Triana) B.M. Boom
Genipa americana
L.
Geophila repens (L.) I.M. Johnston
Macrocnemum roseum Wedd.
Manettia cordifolia Mart.
Mitracarpum hirtum DC
Loretoa peruviana Standl.
Palicourea conferta (Benth.) Sandwith
Pentagonia parvifolia Steyerm.
Pogonopus speciosus K. Schum.
Psychotria sp. 1
Psychotria sp. 2
Psychotria sp. 3 (19462, 19318, 19327)
Psychotria deflexaDC.
Psychotria officinalis Raeusch.
Psychotria platypoda DC.
Psychotria remota Benth.
Randia armata DC.
Richardia scabra L.
Relbunium hypocarpium Hemsl.
Rudgea sp. 1
Rudgea cornifolia (Humb. & Bonpl. ex
Roem. & Schult. ) Standley
Sickingia tinctoreaK. Schum.
Uncaria guianensis J.F. Gmel.
Uncaria tomentosa DC. (18291)
Warszewiczia coccinea Kl.
Warszewiczia cordata Spruce ex k.
Schum.
77
RUTACEAE
Amyris macrocarpa
Angostura sp. 1
Dictyoloma peruviana Planch.
Erythrochiton brasiliensis Nees & Mart.
Raputia sp. 1
Zanthoxylum sp. 1
Zanthoxylum sp. 2 (12751)
Zanthoxylum culantrilo HBK.
Zanthoxylum ekmanii kellermanii (Urb.)
Alain
Zanthoxylum huberi
Zanthoxylum rhoifolium (19297, 19386)
Zanthoxylum sprucei
SALICACEAE
Salix humboldtiana Willd.
SAPINDACEAE
Allophylus sp. 1
Allophylus pilosus (Macbride) A.H. Gentry
Allophylus scrobiculatus
Cupania cinerea
Dilodendron bipinnatum Radlk.
Dodonaea viscosa Jacq.
Paullinia bracteosa Radlk.
Paullinia aff. carpopoda Cambess.
Paullinia sp. 1
Paullinia sp. 2
Paullinia frutescens
Paullinia obovata Pers.
Paullinia setosa Radlk.
Sapindus saponaria L.
Serjania sp. 1
Talisia sp. 1 (19324)
Talisia cerasina
Toulicia reticulata Radlk. (19389)
SAPOTACEAE
Chrysophyllum venezuelanense (19407)
Ecclinusa guyanensis
Manilkara bidentata (A.D.C) A. Cheval.
Michopholis guyanensis Pierre (18099)
Pouteria bilocularis
Pouteria caimito Radlk.
Pouteria ephedrantha
Pouteria procera
Pouteria reticulata
78
Pouteria torta Radlk.

Pouteria trilocularis
Sarcaulus (19366)
SCROPHULARIACEAE
Lindernia thouarsi (Cham &
Schlechtend. ) Edwin
Scoparia dulcis L.
SIMAROUBACEAE
Picramnia sp. 1
Simaba sp. 1
Simaruba amara Aubl.
SOLANACEAE
Cestrum sp. 1
Cestrum arboreum
Cestrum megalophyllum Dun.
Cuatresia fosteriana Hunziker
Cyphomandra sp. 1 (19200)
Cyphomandra sp. 2 (19292)
Cyphomandra splendens Dun.
Juanulloa parasitica(18990)
Larnax subtriflora Miers
Lycianthes sp. 1
Lycopersicon cf. peruvianum Mill.
Markea ulei (Dammer) Cuatrec.
Nicotiana tabacum L.
Solanum sp. 1
Solanum riparium Pers.
Solanum nigrum Acerb. ex Dun.
Solanum sessile R. & P.
Solanum topiro Humb. & Bonpl. ex
Dun.
Witheringia solanacea Miers ex Benth. &
Hook. f.
STAPHYLEACEAE
Huertea glandulosa
STERCULIACEAE
Byttneria sp. 1
Byttneria catalpaifeolia Jacq.
Byttneria pescapraeifolia
Guazuma crinita Mart.
Guazuma ulmifolia Lam.
Herrania sp. 1
Herrania sp. 2
Melochia mollis Tr. & Planch.
Melochia villosa Fawc. & Rend
Pterygota amazonica L.O. Wms. (12695)
Sterculia sp. 1
Sterculia apetala Karst.
Theobroma sp. 1
Theobroma cacao L.
STYRACACEAE
Styrax tessmannii Perkins
TEOPHRASTACEAE
Clavija sp. 1 (19189)
Clavija tarapotana Mez (10196)
TILIACEAE
Apeiba sp. 1
Apeiba membranacea Spruce ex Benth
Apeiba tibourbou Aubl.
Heliocarpus americanus L.
Luehea divaricataMart.
Luehea paniculata Mart.
Luehea speciosa Willd.
Lueheopsis sp. 1
Tilia sp. 1
Triumfetta semitriloba Jacq.
TRIGONIACEAE
Trigonia sp. 1 (12879)
TROPAFOLACEAE
Tropaeolum sp. 1 (18981)
TURNERACEAE
Turnera weddelliana Urb. & Rolfe
Urera laciniata Wedd.

VERBENACEAE
Aegiphila sp. 1
Citharexylum poeppigii
Petrea maynensis Huber
Stachytarpheta cayennensis Schau.
Vitex cymosa Bert. ex Spreng.
Vitex pseudolea Rusby
VIOLACEAE
Leonia glycycarpa R. & P. cauliflorous
Leonia glycicarpa R. & P. ramiflorous =
L. racemosa
Rinorea sp. 1 (12729)
Rinorea viridifolia (18922)
Rinerocarpus ulei (8924)
VITACEAE
Cissus sp. 1
VOCHYSIACEAE
Qualea sp. 1
Vochysia sp. 1
Vochysia sp. 2
ZINGIBERACEAE
Costus sp. 1
Costus sp. 2
Costus sp. 3
Costus lima Schumann
Hedichyum coronarium Koen.
Renealmia sp. 1
Renealmia sp. 2
ULMACEAE
Ampelocera sp.
Ampelocera eduntula
Celtis iguanea
Trema micrantha Blume
URTICACEAE
Boehmeria sp. 1 (12914)
Myriocarpa stipitata Benth.
Urera sp. 1
Urera sp. 2
Urera baccifera Gaudich.
Urera caracasana Griseb.
79
80
Floristic
Composition,
Structure,
and Diversity
Assessment
in the Lower
Urubamba
Region
Alfonso Alonso, Francisco
Dallmeier, Shahroukh Mistry,
Christopher Ros, and James
Comiskey
Program, (SI/MAB)
Percy Nez
Abad del Cusco
Jos Santisteban
Departamento de Entomologa,
Universidad de Trujillo
Gorky Valencia
Abad del Cusco
Severo Balden and
Hamilton Beltrn
Mayor de San Marcos
Introduction
Although the biological diversity in tropical forests has been

documented at many sites, the
need for long-term monitoring of
biodiversity in tropical forests over
time is essential to understanding
the structure and dynamics of such
complex ecosystems (Dallmeier and
Devlin 1992). A fundamental role
of such monitoring programs is to
measure and assess the status of
both plant and animal species at the
study site. Much of the initial work
in these studies is devoted to the
examination of vegetation and the
structure of the forest. This information is then used to study patterns of plant demography, species
turnover, succession, and other
processes key to understanding how
forest dynamics work and how they
respond to disturbances. The
biodiversity plots described here are
part of a global network of longterm biodiversity monitoring plots.
Because the system is standardized,
results from plots within the network can be compared (Dallmeier
et al. 1996).
81
Figure 1. Diagram showing the location of the biodiversity monitoring test plots, well sites, and local
settlements in the Lower Urubamba region.
One of the largest areas of
species richness in the tropics is the
Amazon region of South America.
Studies at Manu National Park in
Per have shown this area to be one
of the worlds highest in biological
diversity (Wilson and Sandoval
1996). More than 1147 species of
vascular plants have been recorded
from this region so far. The diversity of the vegetation provides a
rich environment for high faunal
diversity as well. This paper describes the results of a floristic
assessment at gas well sites and the
immediate vicinity in the Lower
Urubamba region, Cusco, Per. This
area exhibits a mixture of both
82
lowland tropical and Andean vegetation types. The objectives of the

study were to examine the area for
plant species diversity and forest
structure and compare the results
with other sites such as the plots in
Manu.
Methods
The area of study is located in

the northern part of the Cusco
Department. This part of Per has a
diverse floristic composition composed of a mixture of several types
of vegetation. Representatives of
species of tropical dry forest and
montane forest are found (Nez et
al. this volume) as well as a very
strong influence of the flora of the

Amazon Basin.
The two well sites that were
the focus of our work are San
Martin-3 and Cashiriari-2. The
vegetation at San Martin-3 is characterized by an upland tropical
forests mixed with bamboo (Guadua
sarcocarpa). The vegetation at
Cashiriari-2 is characterized as a
primary rainforest. We established
two biodiversity plots at San Martin-3 and one at Cashiriari-2 (see
Fig.1). To select the locations for
the plots, we set a multi-scale plot
to determine areas of high diversity.
Plot Selection
and Establishment
The forest plots were established according to Dallmeier
(1992), wherein botanists select
sites in the area of study using
cartographic information, remote
sensing, aerial photographs, and
field verification. At the sites, field
teams delineate a 1-ha plot (10,000
m2) and divide it into 25 quadrats,
each 400 m2 in size. Close supervision is needed to ensure the least
amount of disturbance to the
vegetation when the plots are set.
The survey team also takes level
measurements at each of the quadrat corners, producing a detailed
topographic map of the plot. Exact
coordinates are determined with the
aid of a geopositioner. The boundaries of the quadrats are demarcated using string, to be removed
later, making orientation easier
within the plot.
Field Measurements
Tree tagging and identification
begins after the corner stakes of the
quadrats are set and the strings tied.
The process includes locating,
measuring, marking, and mapping
all trees with a diameter at breast

height (dbh) >4 cm. Diametric tape
is used to measure dbh, avoiding
any protrusions on the trunk.
Smaller trees are measured with
calipers. Where multiple stems
occur, individual diameters are
measured. Trees are tagged with an
aluminum label that should face
toward the base line of the plot and
set with a nail above the point of
measurement. The nails serve as a
general guide for future measurements (Dallmeier et al. 1993 a,b).
Trees are tagged with a different number consisting of a sequence of 3 double digits. Using
(01-24-09) as an example, the first
2 numbers (01) designate the zone
where the plot is located. The
second set (24) identifies the quadrat. The last 2 numbers (09) represent an individual tree within the
quadrat. No other tree receives this
number. In each quadrat, the tree
numbers start at one and continue
until the last tree is labeled.
We mapped the position of the
trees with a quadrat mapping team
of three people. Two of them stood
at the corners of the quadrat, while
the third moved between the trees
to be measured. Electronic range
finders measured the distance from
each tree to two adjacent corners.
The A and B values recorded are
later used, along with the diameter,
to calculate the exact position of
the tree. The field mapping teams
used different preprinted data
sheets for each quadrat to record
each trees position, dbh, and height
as well as pertinent notes about any
tree or quadrat. After the mapping
teams completed data gathering,
botanists and voucher samplers
recorded the identities of the trees
on the sheets, normally using six-
The
vegetation
at San
Martin-3 is
characterized
by an
upland
tropical
forests
mixed with
bamboo
(Guadua
sarcocarpa).
83
We have
found that
efficient
data management is
key to a
successful,
long-term
forest
biodiversity
monitoring
program.
84
letter codes for the different species

mapped in the plot. Information
from the finished sheet was then
entered into BioMon (see data
management section below) by data
entry teams. For sub-quadrat mapping, the sheets were placed on a
small digitizing tablet that allows
the calculation of the X and Y
coordinates to be transferred directly to the database. This procedure further reduces data entry
errors and increases the speed and
efficiency of data management.
BioMon generates a map of each
quadrat, which field personnel use
for on-site verification of the
correct location of the measured
trees.
Voucher Specimen Sampling
Voucher specimens of the
measured and mapped trees are the
most valuable information for
further study of forest biodiversity.
A minimum of three herbarium
specimens are always sampled; most
are sterile. The botanists sample
and identify the specimens, assisted
by two or three experienced tree
climbers who go after the more
inaccessible samples. Field specimens are held together with flagging
tape labeled with the four doubledigit tree number. The samples are
sorted at the base camp, trimmed,
and placed between absorbent
paper. The tree number, as well as
the botanists collection number,
identify the samples, which are
stacked and bundled, placed in
plastic bags, and preserved with a
solution of 50% alcohol so that
they will not decompose during the
trip to the herbarium.
On arrival at the herbarium, the
samples are placed in driers separated by corrugated plates. They
should be fully dry within two days,
at which point they are sent to the

respective experts to confirm field
identifications. The final stage of
this process is to mount and make
photocopies of the specimens.
Data Management
Data gathered from the
biodiversity plots are entered in the
Biological Monitoring Database
(BioMon). This program arose from
a need for a flexible database to
manage the information coming
from the international network of
biodiversity research sites. The
priority was to provide a speedy and
efficient turn-around time between
the gathering of data in the field
and publication of the information.
We have found that efficient data
management is key to a successful,
long-term forest biodiversity monitoring program (Dallmeier et al.
1993 a,b).
Guiding principles in designing
BioMon were modularity, ease of
use, and compatibilityall aimed at
facilitating quick and accurate
cross-site comparisons. Off-theshelf software was incorporated,
creating portable links that, once
fully documented, are made available to other researchers. The core
database operates in Paradox;
statistical analysis is augmented in
Systat. Graphics for the publication
are produced using Corel Draw, and
report formatting is carried out with
Ventura Desktop Publisher.
User friendly menus mean
inexperienced users need only
minimal training to operate the
system. BioMon was developed
with the field work in mind, where
researchers and students may
manipulate more than 20,000
independent forest observations per
ha over three to five days. Field and
office modules are the two main
components of the system. Their

structures are similar, but not their
intents and capabilities. The field
module manages information for a
specific site; the office-based
module manages and analyzes data
for all the sites, allowing the production of diverse publications. A
series of linked tables provides the
framework for the database. Information from the tables is made
available through forms that can be
called up on the screen.
BioMons database application
language made it possible to create
data validity checks that come into
play during data entry and editing,
thus enhancing the integrity of the
data. Once the data has been
entered, it can easily be viewed and
edited; more information can be
added or corrections made. Sections
of the data may also be printed,
providing a hard copy of the database content and aiding in data
verification. We recommend that
back-up copies be made, updated,
and stored on both the computer
and diskettes.
Once in the office, data management proceeds in the office
module. Information from each
forest site is stored within its own
directory in a series of tables with
identical structures so that crosssite comparisons can be made. Data
entered on portable computers in
the field can be downloaded via a
network connection or from the
backup diskettes. The office module
is an extension of the field module,
accommodating database management as well as statistical manipulation, map creation, and publication
development. We are automating
the movement of data between the
modules as much as possible to
improve efficiency and consistency
in the production of user and field
guides and other scientific publications. BioMon allows the production of complete field reports at
each 1-ha plot as soon as data entry
and verification is completed. The
reports provide feedback to make
on-the-spot decisions as to whether
additional information from the
field is needed. They also provide
immediate information on tree
species composition, structure, and
diversity.
Results and
Discussion
Because the biodiversity plots

at San Martin-3 and Cashiriari-2
have different topographic features,
one would expect different vegetation types. Plots #1 and #2, at San
Martin, are located in rugged terrain
with many hills and valleys. On the
ridges of these hills, thickets of the
bamboo Guadua sarcocarpa are often
encountered. In fact, this species of
bamboo accounts for up to 59% of
all individuals in the plots. Its role
in determining the structure of the
forest is likely to be quite strong.
During field work, it became apparent that heavy rains and wind
knocked bamboo down and that
this in turn had an effect on neighboring tree species. We frequently
found trees without a crown, and
the immediate effect on such trees
is a reduction in photosynthetic
products, thus reducing tree growth.
We also observed that several
individuals damaged by bamboo
died from the damage. The third
biodiversity plot (# 3) at
Cashiriari-2 is quite different from
the San Martin-3 plots. Cashiriari is
typical lowland forest with few
ridges or hills. The vegetation lacks
bamboo; species such as Grias
85
250
3000
200
150
100
50
40
Total Basal Area
3500
# of Individuals
# of Species
300
2500
2000
1500
1000
500
0
0
P lot 1
(S M -3)
P lot 2
(S M -3)
S ite
P lot 3
(CA-2)
35
30
25
20
15
10
5
0
Plo t 1
(SM -3)
Plo t 2
(SM -3)
S ite
Plo t 3
(C A-2 )
Plo t 1
(SM -3)
Plo t 2
(SM -3)
Plo t 3
(C A-2 )
S ite
Figure 2 (left). Number of woody plants at each of the three plots (SM-3 = San Martin-3 and
CA-2 = Cashiriari-2). Figure 3 (center). Number of individual woody plants at the three sites.
The bars for Plots 1 and 2 show the number of individuals with and without the bamboo species
included. Plot 3 had no bamboo. Figure 4 (right). The total basal area of the three plots.
peruviana and Calatola venezuelana
are quite common.
The total number of species at
the three plots was surprisingly
different. Plots #1 and #3 have
similar number of species (258 and
271, respectively) while plot #2 had
a much lower count224 species
(Fig. 2). The total number of individuals at these sites was also
markedly different. If the number
of Guadua are included, then plots
#1 and #2 had 2975 and 2585
plants, respectively; plot #3 had
1522. When Guadua is not included
in the analysis, plot #1 has 1650
individuals, and plot #2 has 1260
(Fig. 3). In other words, Guadua
accounted for 45% of the individuals in plot #1 and 58% in plot #2.
This dominance by Guadua, especially in plot #2, may account for
the low number of other species
and their low densities at this site.
The influence of Guadua is also
seen in the total basal area measurements for the three plots. Plots #1
and #2, with high numbers of
Guadua stems, have low basal areas
whereas plot #3 has a much higher
value (Fig. 4). The bamboo apparently affects not only the number of
86
other species in a plot, but their size

as well.
A total of 16 species occurred
at densities of more than 1% (of all
non-bamboo individuals) in the
plots. Other than Guadua, the most
common species in plot #1 were
Iriartea deltoidea, Matisia cordata,
Rinorea viridifolia, and Nectandra
longifolia (Fig. 5). In plot #2, only 14
species occurred at more than 1%
with Iriartea deltoidea and Matisia
cordata the most common plants
(Fig. 6). Plot #3 had a total of 21
species that occurred at densities of
more than 1% (Fig. 7). Again,
Iriartea deltoidea was common along
with Grias peruviana and Calatola
venuzuelana. The plot at Cashiriari-2
displayed much more equity in
densities among species than did
the plots at San Martin-3, although
at San Martin, there were only a few
common species, with the rest
occurring at low densities.
A similar pattern is observed
when examining species with the
largest dbh in the three plots. In
plot #1, with the exception of Ceiba
pentandra and Tabebuia ochracea (large
trees that only occurred once), all
species had an average dbh of
Inga sp1
17
Neea sp1
17
Sorocea piliata
17
Plot 1 (SM-3)
Cyathea sp1
18
Protium puncticulum
18
24
Pseudomelia laevis
26
Guarea kunthiana2
30
Lunania parvifolia
Guapira sp1
34
35
Sorocea steinbackii
36
Pentagonia parvifolia
39
Guarea kunthiana
Nectandra longifolia
47
73
Rinorea viridifolia
92
Matisia cordata
96
Iriartea deltoidea
0
10
20
30
40
50
60
70
80
90
100
# of Individuals
Figure 5. Species in Plot 1 comprising more than 1% of the total number of plants in the plot.
Note that bamboo individuals (1325 of 2975) have been excluded.
Cyathea sp1
Heisteria acuminata
Guapira sp1
Lonchocarpus sp1
Lonchocarpus spiciflorus
Perebea guianensis
Neea sp1
Otoba parviflolia
Sorocea steinbackii
Lunania parvifolia
Guarea kunthiana
Iriartea deltoidea
Matisia cordata
13
P lo t 2 (S M -3 )
13
14
15
16
16
21
22
31
31
36
69
87
0
10
20
30
40
50
60
70
80
90
100
# of Individuals
Note that bamboo individuals (1512 of 2585) have been excluded.
87
15
15
16
16
18
18
19
21
22
Inga sp1
Quararibea witii
Bauhinia tarapotensis
Sapium marmieri
Hasseltia floribunda
Otoba parvifoli
Cyathea sp2
Eugenia sp1
Lunania parvifolia
Guarea macrophylla
Guarea kunthia na
Lonchocarpus spiciflorus
Matisia cordata
Rionorea viridifolia
Chrysochlamys membranacea
Pentagonia parvifolia
Rinorea guianensis
Trichillia poeppigii
Iriartea deltoidea
Calatola venezuelana
Grias peruviana
Plot 3 (CA-2)
28
30
31
38
39
46
48
48
50
65
66
77
0
10
20
30
40
50
60
70
80
90
100
# of Individuals
0.2 m to 0.5 m. Nearly all of the
large species in plot #2 were in the
same range, but in plot #3, most of
the large species ranged from 0.4 m
to 1.0 m in dbh while Ficus mathewsii
reached 1.25 m dbh.
It is apparent that many of the
differences recorded at the three
plots reflect the presence of
Guadua. The plot with the most
Guadua, plot #2, also contained the
least number of tree species, the
least number of individuals, the
smallest total basal area, and the
smallest average dbh. The plot with
no Guadua (plot #3) had the highest
number of species, the most basal
area, and many tree species with
large dbh. The presence of Guadua,
either physically or via its competitive edge during the dry season, is
certainly felt in the area of study. It
is particularly well adapted to steep
slopes and ridges and ephemeral
water supply. The lack of such
harsh topography at Cashiriari-2 is
88
likely the reason that bamboo is

absent there (Figs. 8 and 9).
The presence of Guadua by
itself does not explain the differences between plots #1 and #2,
which are located within 300 m of
each other. Plot #2 does have
considerably more bamboo, and the
effects of this on the other vegetation are apparent from the above
descriptions. The preference by
Guadua for that plot may be caused
by subtle differences in the topography of the two sites or the availability of moisture. Plot #1 has less
steep slopes, which may explain its
reduced Guadua density. As more
sites are analyzed, the relationships
among Guadua and the other vegetation should be clarified.
Certain similarities between the
forest plots in Urubamba and the
other SI/MAB plots in Manu are
present. The number of species and
their densities are quite comparable.
The major difference is that in
Manu, many species have a dbh of
Figure 8. Horizontal and vertical vegetation profiles at San Martin-3 well

site, Lower Urubamba, Cusco, Per.
Figure 9 Schematic representation of the topography of biodiversity Plots

#1 and #2.
0.8 or higher whereas at these sites
only the rare species such as Ficus
mathewsii have large stems. This
difference may have to do with the
correlation between dbh and bamboo. The presence of Guadua
drastically reduced the girth of
coexisting tree species. In plots
where bamboo is absent, such as
the Manu plots and the one at

Cashiriari-2, the girth of trees is
considerably larger. The effect of
bamboo on the distribution and
growth these forest plants will be
clarified with future research.
89
References
Dallmeier, F. 1992. Long-term

monitoring of biological diversity
in tropical forest areas: methods
for establishment and inventory of
permanent plots. MAB Digest 11.
UNESCO, Paris.
Dallmeier, F., and F. Devlin. 1992.
Forest diversity in Latin America:
Reversing the losses? Journal of
Forest Science 5: 232-270.
Dallmeier F., R. Foster, and J.
Comiskey. 1993a. Users guide to
the Manu Biosphere Reserve
Biodiversity Plots, Per. Vols. I
and II. Smithsonian Institution,
Washington, DC.
Dallmeier F., R. Foster, and J.
Comiskey. 1993b. Field guide to
the Manu Biosphere Reserve
Biodiversity Plots, Per. Vols. I, II,
III, IV. Smithsonian Institution,
Washington, DC.
90
Dallmeier, F., M. Kabel, and

R. Foster. 1996. Floristic composition, diversity, mortality, and
recruitment on different substrates: lowland tropical forest,
Pakitza, Rio Manu, Per. Pages
61-77 in Manu: The Biodiversity
of Southeastern Per. (D. E.
Wilson and A. Sandoval, eds.) Ed.
Horizonte, Lima.
Wilson, D. E., and A. Sandoval.
1996. Manu: The Biodiversity of
Southeastern Per. Ed. Horizonte,
Lima.
.
Long-term
Vegetation
Monitoring
Plan
Francisco Dallmeier,
Shahroukh Mistry, and
James Comiskey
Program (SI/MAB)
Introduction
Biological systems, such as

tropical forests, are extremely
complex networks of biotic and
abiotic interactions. While all
components of a complex system
are essential, plants comprise
perhaps the most important structural and function component of a
forest ecosystem. Their roles are
many. The fact that plants absorb
large amounts of carbon dioxide
and produce oxygen to support
animal life is well known. So is the
fact that all the energy available for
animals comes either directly or
indirectly from plants; their role as
producers makes all animals dependent on them for survival. But
plants also provide lesser known
benefits. They are extremely good at
cycling nutrients in the soil, and in
some cases such as leguminous
plants, they increase the nitrogen
content of soil and make it more

fertile. Plants can also control the
climate at a local and global scale,
and they prevent erosion, maintain
temperature and humidity levels,
and protect the forest ecosystem
from disturbances caused by floods
or drought (Primack 1995).
Forest plants are crucial to
multi-taxa monitoring as well. Some
plant species such as fig trees are
considered keystone forest species because of their role in supporting numerous species of birds
and bats during seasons when fruits
are not readily available (Terborgh
1986). Without keystone species,
many bird and bat population levels
would decline, leading to decreases
in populations of plants that depend on these animals for seed
dispersal or pollination (Cox et al.
1991).
The botanical implications of
forest biodiversity and the importance of forest plants to humans
cannot be understated. There are
tremendous economic, medical, and
agricultural benefits to studying
forest plants. In addition to their
value in finding cures for diseases,
91
Long-term
forest inventories can
provide
managers
with important information
needed to
establish
conservation
priorities
and serve as
the base of
future work.
92
tropical forest plants provide a

natural genetic warehouse that
helps maintain many domesticated
agricultural species. For example,
many of the central American
banana plantations were recently
contaminated by a disease that
could only be stopped by creating
new hybrids with wild varieties of
bananas from tropical forests.
Without such wild plants, the future
of many agricultural crops could be
in jeopardy (Hladisk et al. 1993).
To conserve tropical forest
diversity and promote sustainable
use of forest resources, it is necessary to understand how forests
change. The permanent plot inventory system proposed here provides
a baseline distribution of tree
species and describes habitats
within a particular site. Monitoring
change in these plots over time will
enable an understanding of the
impacts of natural and humancaused disturbances on species and
community composition. Data
produced from monitoring can also
be used to predict future changes,
based on current understanding of
the ecosystem processes.
Long-term forest inventories
can provide managers with important information needed to establish
conservation priorities and serve as
the base of future work. Without
adequate inventories, managers
cannot measure change over time,
identify gaps in existing knowledge,
or set priorities to fill these gaps.
Good inventory data are key to
successful research projects, conservation, and management. Permanent inventory plots can provide
data that are otherwise unobtainable and that, when maintained and
measured repeatedly, become more
valuable over time.
Long-term inventories can also

be used to help integrate management of protected areas with
compatible development on surrounding lands. As an example, data
gathered from permanent inventory
plots in temperate forests have long
been used to project future growth
and yields on managed timberlands,
information that is critical in planning for sustainable development.
Forest plants form the base of
all food webs and support the
diversity of animal life found in
tropical forests. The need to identify and monitor the status of plants
in a forest is essential in any
biodiversity monitoring program.
Many of the patterns seen in the
abundance and distribution of
animals can be correlated to the
structure and health of the forest
plants. To truly understand forest
diversity and dynamics, a detailed
vegetation monitoring program has
to be established. An outline of
how such a program would be
implemented in the area of gas well
sites in the Lower Urubamba region
of Per is presented below.
Monitoring Program
Objectives
Continue obtaining vegetation baseline information at
the well sites, the proposed
pipeline route, and the general Block #75.
This process will continue the
assessment and surveys already
initiated. It allows for quick identification of different forest sites,
species diversity estimations, and
characteristics of the vegetation.
The amount of data will increase
through the incorporation of more
sites and the sampling of data on
species dynamics.
Gain a better understanding

of habitat types, vegetation
types, and spatial distributions of species in the area of
influence of the well sites and
pipeline and the general area
of influence of the gas development project.
It is important that the implications of human activity on vegetation patterns and forest regeneration be thoroughly studied. Changes
in forest structure caused by human
activity at the well sites and along
the pipeline will induce regeneration of plants in the disturbed areas.
The species that dominate these
sites will be pioneering secondgrowth species that will form a
forest quite different from the
surrounding undisturbed forest. The
influence of this successional
sequence will have significant
impact on the animals currently
inhabiting the area. It is thus imperative that the vegetation in the
undisturbed forest as well as regenerating forest be studied and analyzed.
Obtain quantitative information about how representative
the forest monitoring plots
are compared to the multiscale vegetation plots and the
larger unsampled landscape.
Vegetation studies require data
from numerous sites in the forest to
get a representative sample. Since
this forest is very large and it is
possible to sample only a few
hectares, it is important that the
sites be representative of the larger
area. Vegetation sampling will thus
be performed at various scales,
from small 0.1-ha and 1-ha plots to
large 50-ha plots.
Details of the methodologies
used to gather the needed data and
analyze them are presented below.
Monitoring Program
Components
Training and education
Training provides assurance of
reliability of the information produced. During the first part of the
monitoring phase of the project,
staff will be trained and educated in
all aspects of vegetation monitoring: measuring vegetation, identification, processing and curation of
specimens, establishment of multiscale vegetation plots and larger SI/
MAB biodiversity monitoring plots,
training of para-taxonomists for
plant identification, data entry and
management through the BioMon
software program, and data analysis
and interpretation. Training will be
conducted in Per and at appropriate international locations such the
Smithsonian Institution in Washington DC and other counterpart
organizations. Education of local
staff will require travel by specialists to conduct in-country training.
Compilation of baseline vegetation information for well
sites, pipeline area, and general area of influence of the
project.
This component will be accomplished through the following steps:
field sampling program, specimen
processing, specimen identification,
data analysis and interpretation,
report preparation, and publication
of reports. The approaches are a
combination of tested methodologies that provide information on
vegetation community richness,
composition, structure and dynamics of species assemblages in representative habitats of the area. The
combined approaches will provide
quantitative (species richness) and
qualitative (basal area, frequency,
density, dominance) information
93
that allows comparison among sites

and within the general area.
The
methods
outlined
here are
part of
the methodology
developed
and used
by
SI/MAB
researchers at
biodiversity
plots
throughout the
world
Monitoring Program
for Well Sites,
Pipeline, and
Area of Influence
In the same manner as the
assessment, the monitoring activities also require a sampling program, specimen processing, specimen identification, data analysis
and interpretation, and the preparation of publication and reports.
Based on the information gathered
during the vegetation assessment
phase, the monitoring program will
be designed and established. Additional studies will also be conducted on seed dispersal and regeneration processes in areas affected
by the well sites and the pipeline.
Vegetation
Monitoring Strategy
The methods outlined here are
part of the methodology developed
and used by SI/MAB researchers at
biodiversity plots throughout the
world (Dallmeier et al. in prep.).
Abiotic Sampling:
Gathering Baseline Data
Initially, considerable energy
will be invested in obtaining
baseline information at the plots
and in measuring and monitoring a
few environmental parameters.
These data can help identify and
describe the site more thoroughly,
and they will provide greater understanding of the effects of abiotic
factors such as rainfall and soil
conditions on vegetation and other
taxa.
Climate data. It is necessary
that as much climate data as possible be gathered. The relationships
94
among precipitation, temperature,

and vegetation can be quite significant, yet they are often ignored in
many vegetation studies. The
inclusion of climate data is essential to an understanding of changes
in vegetation over time and in
comparing vegetation among sites.
A weather station will be placed at
the study site to gather data on
rainfall, temperature, and wind.
Additional probes may be placed in
different strata of the forest for
comparative purposes.
Soil composition. Analysis of
soil at various locations will provide
important information about the
forest and the vegetation patterns.
Random soil cores along transects
across the biodiversity plots will be
undertaken. A typical soil auger will
be used for such sampling and the
soil stored in zip-lock bags or whirlpacks for analysis. Such soil data
will be gathered at least once in
setting up a plot and, if possible,
whenever the plot is revisited.
Typical soil analysis include estimating sand/silt/clay composition
and organic material as well as
quantifying various inorganic
parameters such as nitrogen, phosphorus, potassium, etc.
Available light. When obtaining the vegetation data, the amount
of light reaching the forest floor
should also be measured. Light
penetration in forests is important
in determining regeneration patterns, especially in gaps. The species composition in gaps can often
be quite different from the neighboring forest. Dramatic differences
in species abundance may not be
easily understood without the
corresponding light information. A
hand-held light meter will provide
quick and accurate measurements
of light.
Canopy cover. The percent

cover, a measure of direct skylight
available at any point on the
ground, is also a measure frequently
used in forest research. This will be
measured using densiometers,
optical devices that permit viewing
of a portion of the canopy and
estimating the percent of the
canopy that is open.
Monitoring Plot Data
Plot locations will be precisely
measured with the most accurate
means available. Whenever possible
a hand-held GPS (global positioning
system) unit will be used to measure
coordinates. In addition, the exact
elevation of each permanent
marker will be established, using a
GPS or a pre-calibrated altimeter.
Alternatively, it is possible to
measure the angle and distance of a
plot marker from a site of known
elevation. Details will also be
provided on the exact location of
each permanent plot marker. Other
parameters that will be included are
slope and aspect (direction the
slope faces) of the plot.
Background Data
At the time of each plot establishment, details about the history
of the land, past use by indigenous
groups and/or industry, and any
obvious disturbance effects such as
past storms, grazing, or logging will
be documented. All available
information on the flora and fauna
of the region will be documented;
especially important are rare, endemic or threatened species in the
area.
Continue assessment of
species diversity at well sites and
proposed pipeline. The acquisition
of comprehensive and accurate
plant identification requires specific
methodologies for surveys. The

details of where, when, and how
plant specimens are sampled can
significantly influence the effectiveness of sampling efforts. For example, most plants are identified by
their reproductive parts (flowers
and fruits), and it is thus necessary
to ensure that sampling efforts
coincide with plant phenology. To
this end, we use a detailed methodology for locating, sampling, and
preserving plant specimens. The
methodologies presented below
have been adapted from the Missouri Botanical Gardens Field
Techniques.
Locating the areas in a forest to
sample plants can be the most
challenging part of sampling. Although the forest area of interest is
the primary focus for sampling,
other surrounding areas should also
be studied. Forest edges, tree-fall
gaps, and clearings are of particular
importance because they provide
access to branches of canopy tree
species that would be inaccessible
in the mature forest. The added
light in these areas also provides
suitable conditions for flowering
and fruit production. However,
many uncommon and rare species
are often found in older forests, not
in disturbed areas; thus, detailed
sampling should not be excluded.
The number of plant samples
depends on the future use of the
specimens. Duplicates will be
gathered to distribute to museums
of all institutions involved in the
study as well as for museum exchange programs and identification
by experts. It may not be possible to
sample a sufficient number of
fertile (with flower and fruit)
samples, but it is more important
that time and energy be spent on
locating at least one fertile speci-
95
Climbing
equipment
and other
tools will be
required to
access the
canopy
branches
of plants
and gather
leaf and
flower/fruit
samples.
96
men and then obtaining duplicates.

Representative materials of all
stages of the plant will be sampled
as leaf form and structure vary with
age.
Climbing equipment and other
tools will be required to access the
canopy branches of plants and
gather leaf and flower/fruit
samples. Plant material will be kept
in plastic bags during sampling and
then later placed in sheets in a
wood press. The samples will be
preserved with diluted alcohol or a
formaldehyde solution. Later they
will be dried over a heat source to
complete the preservation process.
Throughout, care will be taken
to record and document each
specimen. Information such as
location, elevation, date, identification, collector, preservation technique, etc. will all be recorded and
stored in a computer database,
allowing for future data management.
Assessment at multi-scale
vegetation plots to select longterm monitoring sites. To assess
and monitor the vegetation of
Lower Urubamba region, the forest
needs to be sampled in a precise
and scientific manner. Care will be
taken to assure that each habitat
type, as identified by satellite
imagery and ground observations, is
equally sampled (Stohlgren et al. In
press a). Within each habitat type,
the location of plots will be selected at random to avoid bias in
site selection caused by subjectivity
(e.g., preference for locating near a
trail).
The first stage in vegetation
assessment and monitoring is the
establishment of five to ten small
(20 m x 50 m) plots per habitat
type. Some of these have already
been set up at the two well sites
and more are being added. These

plots are multi-scale in that they
sample vegetation in sub-plots of
different sizes (Stohlgren et al.
1995). Within each 1000-m 2 plot,
ten 1 m 2 subplots, two 10-m 2 subplots and one 100-m2 sub-plot are
also sampled. This sampling design
allows for analysis of plant species
diversity, the calculation of speciesarea curves, and the extrapolation
of diversity information to larger
areas of forest (Stohlgren et al. In
press b). These assessment plots
also allow for the selection of sites
to establish future 1-ha permanent
monitoring plots.
One-ha biodiversity monitoring plots for long-term studies.
The fundamental unit for measuring
forest biodiversity is the 1-ha plot
that is used throughout the SI/MAB
global network of plots (Dallmeier
1992). This has become the standard size for many forest plots and
can thus be used for comparisons
among sites. At the Urubamba
study area, these plots will be
established in each of the vegetation types. The number and location
of the plots will depend on the
information provided by the analysis of the smaller multi-scale plots.
The 1-ha plots will provide
sites for the study of plant demography, dynamics, and diversity.
Typically these plots are 10,000 m 2,
but the actual configuration will
depend on the topography of the
area. Each of the plots is divided
into 25 smaller 20-m 2 quadrants.
Each quadrat is subsequently
divided into four 5-m 2 subquadrants. The entire plot and all the
quadrants will be surveyed by a
field team using a theodolite or
surveying compass. Permanent
markers will be placed at each
quadrat corner, and the entire plot
will be corrected for changes in

slope.
Within each plot, each tree
above 10 cm in diameter at breast
height is identified, measured, and
tagged, and its exact location is
mapped. All of this information is
then entered into a database program (BioMon) that analyzes the
structure, diversity, and dynamics
of the plot. The information provided by this program is then used
to produce the users guide as well
as the data for further statistical
analysis. These plots will be surveyed at regular intervals to measure changes in forest structure and
species dynamics.
The 1-ha plots will also be used
for intense sampling and monitoring
of multi-taxa information. These
plots provide an ideal site for the
study of other taxa such as insects,
birds, and mammals. Together with
the vegetation data, these datasets
can provide important information
on the correlation among different
groups of animals and plants and
act as predictive tools in studying
the densities of organisms.
Establishing 25- and 50hectare plots to study forest
structure and dynamics at a
large scale. The Center for Tropical Forest Science has established a
series of large plots around the
world (Condit In press). These plots
are typically about 50 ha in size and
require considerable effort to setup
and maintain. However, the yields
from such large plots can be significant. For example, the site in
Sarawak, Malaysia, has 1175 species and 350,000 individual stems.
Large plots allow detailed population monitoring of many species,
especially those that are rare in the
forest. Rare species, by definition,
are hard to find, and in a small plot
they may be very few or altogether

absent. Large plots also provide
demographic data at a scale that is
quite useful, and they can be used
to monitor vegetation responses to
climate change or human disturbance.
These large forest dynamics
plots (FDPs) serve two broad
functions. First, demographic tree
data provide a baseline to monitor
and better understand forest dynamics and regeneration, the maintenance of species diversity, extinction processes, and the impact of
climate and land use changes on
tropical ecosystems. Second, the
network of sites provides baseline
data for targeted research on sustainable forest management. FDPs
supply critical dataoften in
conjunction with other socioeconomic and silvicultural data
for determining the biologically
sustainable harvest rates for timber
and non-timber forest products;
selecting native tree species for the
reforestation of degraded areas; for
statistically evaluating sampling
protocols for monitoring
biodiversity; and designing reserves
and protected areas to conserve
biodiversity. Each of these research areas relies heavily on the
information produced by the FDPs
regarding abundance, size/distribution, productivity, regeneration, and
spatial distribution. Because forest
dynamics plots follow a standardized methodology, analytical and
management tools developed from
one site can be transferred to
another. The ultimate goal of these
targeted research programs is to
provide a scientifically rigorous and
economically sound basis for devising strategies and incentives to
conserve, monitor, and manage
tropical forest biodiversity.
These plots
provide an
ideal site
for the
study of
other taxa
such as
insects,
birds, and
mammals.
97
A similar FDP could be set up

in the Lower Urubamba study area.
The location will be determined by
the multi-scale plots described
above. The size of the plot will be
determined by the location, but is
likely to be between 25 and 50 ha.
This plot will complement the
smaller plot types; combined, they
will provide a unique opportunity to
study forest ecology at different
scales. Such a system will prove
useful in extrapolating vegetation
patterns from the plots to the entire
forest.
The methodology used to
establish the large plot is similar to
that described for the 1-ha plots. It
requires a survey team using theodolites and stakes to mark out the
entire area. The standard 50-ha plot
is 500 m x 100 m; delineating it can
take a crew of 5 persons 5 months
to complete (the 1-ha SI/MAB
plots normally require 1 to 4 days
to complete, depending on the
topography). Once the large plot is
demarcated into a 20 x 20 m grid,
the measurement of vegetation can
begin. In some 50-ha plots, all
stems greater than 1 cm have been
measured and tagged, promoting
detailed studies of seedling demography and recruitment. Data gathered from this plot will be maintained in a computer database for
analysis.
Studying forest regeneration
patterns in the well and pipeline
areas. One of the fundamental
questions facing ecologists is forest
regeneration. The ability of a forest
to respond to natural and anthropogenic disturbances, is a process that
requires considerably more research.
Of particular importance is the
ability to predict how a forest
ecosystem will respond to perturbations, how long the response might
98
take, what species are most important in the regeneration process,

and how this process might affect
the resident fauna (Mistry 1995).
The study of the successional
processes in a forest have been well
documented, yet few cases exist
that document the natural regeneration process after clearcutting
within a dense tropical forest.
The Urubamba project presents
a unique opportunity to study the
process by which tropical forests
regenerate over time. Since the role
of seed dispersal agents is integral
to this process, it also permits us to
study an important animal-plant
interaction to determine which
animal species are important seed
dispersers. We plan to set up additional (10 to 20) vegetation plots to
examine the regeneration process.
This involves locating disturbed
areas, either along the pipeline or
on the edge of the well sites. The
plots will be relatively small (0.1
ha) but will measure and map seeddeposition patterns and seedling
development. Within these plots,
seed trays will be placed to measure
seed rain from dispersers as well as
wind-dispersed seeds. The seeds
will be identified either by structural characteristics or by germinating them in a makeshift greenhouse.
Studies will also be conducted to
see which animal species disperse
the seeds into the openings. The
plots will be regularly monitored to
measure growth and mortality of
seedlings, providing a measure of
species turnover and successional
sequence. Comparisons will also be
made with areas where the regeneration process has been aided by
human intervention to examine
differences in the successional
sequence.
Development of users
guides for monitoring plots.
Users guides contain basic biological information about permanent
plots at long-term research sites.
The purpose is to assist in conducting more effective research and
training. Each users guide is accompanied by a complementary
field guide containing a condensed
version of information from the
users guide for use in the field.
Together, these documents make
available to researchers all the basic
information gathered in the plots.
They are designed for loose-leaf
binders to accommodate continuous
updating as more information
becomes available.
A consistent format is followed
in producing the users guides. The
first section provides a detailed
explanation of the site ecology and
gives a brief overview of the areas
climate, topography, soils, vegetation, fauna and history. The second
section details the methodology for
establishing a permanent plot,
presented in a readily understandable manner for researchers planning to use the same methodology
in other areas. Section three consists of the specific information
gathered at the plot, providing a
foundation for comparative analysis
of the plots vegetation. The users
guides include a complete set of
maps for each quadrat in the plot,
along with data sheets and a full
size map of the plot. Whenever
possible, the guides also include a
mini-herbarium of tree specimens, designed to help in
identification.
References
Condit, R. In press. Tropical Forest

Census Plots: Methods and results
from Barro Colorado Island and a
comparison with other plots.
Landes Publishers.
Cox, P.A., T. Enquist, E.D. Pierson,
and W.E. Rainey. 1991. Flying
foxes as strong interactors in South
Pacific island ecosystems: A
conservation hypothesis. Conservation Biology. 5: 448-454.
Dallmeier, F., J. Comiskey, and
S. Mistry. In preparation. Methodology for the creation and inventory of long-term plots.
Dallmeier, F., ed. 1992. Long-term
Monitoring of Biological Diversity
in Tropical Forest Areas: Methods
For Establishment and Inventory
of Permanent Plots. MAB Digest
11. United Nations Educational,
Hladick, C.A. et al., eds. 1993.
Tropical forests, people and food.
Parthenon Publishing and
UNESCO. Paris.
Mistry, S. 1995. Seed dispersal
patterns and their relationship to
disturbance levels and seasonality.
Ph.D. dissertation. University of
New Mexico, Albuquerque.
Primack, R. B. 1995. A Primer of
Conservation Biology. Sinauer
Associates, Sunderland,
Massachusetts.
Stohlgren, T. J., M. B. Falkner, and
L. D. Schell. 1995. A modifiedWhittaker nested vegetation
sampling method. Vegetatio 117:
113-121.
a. Rapid assessment of plant
diversity patterns: A methodology
for landscapes. Ecological Monitoring
and Assessment.
99

b. Multi-scale sampling of plant
diversity: Effects of the minimum
mapping unit. Ecological
Applications.
Terborgh, J. 1986. Keystone plant
resources in the tropical forest.
100
Pages 330-344 in Conservation

Biology: The Science of Scarcity
and Diversity. (M. E. Soule, ed.)
Sinauer Associates, Sunderland,
Massachusetts.
Arthropods:
Biodiversity
Assessment
in the Lower
Urubamba
Region
Jos Santisteban
Departamento de Entomologa,
Universidad Nacional de la
Libertad (Trujillo)
Gorky Valencia
Abad del Cusco
Alfonso Alonso
Program (SI/MAB)
Introduction
The South American neotropical rain forest is one of the most

complex and diverse ecosystems on
earth. However, knowledge and
understanding of this vast reservoir
of life is still very much incomplete
(Gentry 1990). Given the continuous, accelerated destruction of
tropical rain forests and the associated decline in earths biodiversity,
it is increasingly important to study
these areas to inform management
decisions about conservation and
rational use.
In discussing animal diversity,

most people usually think in terms
of vertebrate species such as large
mammals, birds, and reptiles. These
constitute what can be called the
macroscopic fauna of tropical
forests. However, they represent
only a small fraction of forest
biological richness. The microscopic
forest fauna attracts less attention
than the macroscopic fauna, with
the exception of a few cases of
medically or economically important species. The vast bulk of
microscopic fauna is made up of
myriad small to very small organisms with secretive life styles.
The principal element of the
micro-fauna is the group known as
arthropods, which represent the
largest component of biological
diversity on the planet. Approximately 75% of all described animal
species to date belong to this group,
a proportion that increases as more
studies in tropical regions are
undertaken. In recent years, research on tropical forest canopies
has produced a great number of
arthropod species new to science,
and the findings may result in a
101
Temporal
changes in
arthropod
activity
occur daily,
as different
species are
active during the day
and night.
102
drastic increase in current estimates

of biological diversity found in
these forests. Besides the fact that
their species richness far exceeds
other groups, arthropods also
account for more biomass than that
of all vertebrates in a given area
(Wilson 1987).
Arthropods are characterized
by bodies covered with hardened
walls and locomotory appendices
that are divided into distinct segments or parts. Arthropods include
well-known and common animals
such as butterflies, flies, mosquitoes, bees, wasps, dung beetles,
plant bugs, spiders, scorpions,
mites, crabs, crayfishes, and lobsters and a huge number of less
conspicuous organisms.
Their generally diminutive size
does not stop arthropods from
performing critical roles in forest
dynamics and function (see note at
end of paper). Arthropods play a
key role in the structure and dynamics of forest ecosystems. They
are important as herbivores, consuming the living tissue of higher
plants and therefore regulating plant
growth; as predators in a broad
sense, including species that are
parasites on others and thus regulating animal populations; as pollinators, assuring plant reproduction;
and as nutrient recyclers involved in
decomposition of dead organic
matter. Arthropods are also very
important as vectors of diseases
and pathogens to animals and
humans.
Most arthropod species are
sensitive to environmental changes,
both spatial and temporal. Environmental gradients such as changes in
water availability and small differences in temperature will limit
species distribution. Temporal
changes in arthropod activity occur
daily, as different species are active

during the day and night. The
composition and density of arthropod communities for a given place
also change throughout the year;
many species have higher densities
during the rainy season, while other
species are more abundant during
the dry season. Habitat heterogeneity increases the number of species
at a larger scale; structural patterns
of the forest have a profound
influence on arthropod composition
and abundance, as these animals are
found from deep in the soil up to
the top of tree canopies. Because of
their small size and varied ecological requirements, arthropods exploit
and dwell in every conceivable
resource in the forest, including
higher plant tissues, flowers and
fruits, soil, leaf litter, fallen logs,
dead and live animals, nests, moss
and fungi, temporary and permanent standing water.
The neotropical rain forests in
southeastern Per are some of the
richest in the world, and have been
the focus of considerable interest in
recent years. Nevertheless, relatively few studies have concentrated on the arthropod fauna.
Several studies on the entomofauna
of the Tambopata Reserved Zone
were published in the Revista
Peruana de Entomologa Vol. 27,
1984, and included papers on
Odonata (Paulson 1984),
Cicindelidae (Pearson 1984),
Asilidae (Fisher 1984), Tabanidae
(Wilkerson and Fairchild 1984),
Lepidoptera (Lamas 1984a, 1984b),
and Euglossine bees (Dressler
1984). A recent volume has been
published on the studies carried out
at the Pakitza station in the Manu
National Park (Wilson and
Sandoval 1996). Arthropod studies
in that volume are for butterflies
Table 1. Arthropods sampled using different protocols in the Lower Urubamba

region.(* = estimated number of specimens)
Sweeping
General Winklers Pitfalls /Beating Light Baits Malaise Total
Taxon
Collembola
19
25
15
59
Ephemeroptera
40
35
75
Odonata
1
10
11
Orthoptera
265 22
8
89
25
544
Acrididae
1
Mantodea
89
89
Gryllidae
2
12
89
Tetrigidae
4
6
1
Tettigonidae
1
19
Blattodea
30
7
13
2
1
53
Phasmatodea
15
16
Phasmidea
1
Isoptera
93
13
106
Heteroptera
152
4
2
11
8
1
130
318
Coreidae
3
1
Lygaeidae
1
Miridae
1
Ochteriidae
Pentatomidae
4
2
225
1
Homoptera
53
2
1250 1653
Cercopidae
3
Cicadelloidea
83
Membracidae
1
3
7
19
Dermaptera
4
1
10
44
8
1034 200
Coleoptera
144
54
649
*1400 2977
17
Carabidae
10
Chrysomelidae
85
1
Cicindelidae
Coccinellidae
2
1
Curculionidae
46
Driliidae
1
Elateridae
6
Elmidae
5
(Robbins et. al. 1996), spiders
(Silva and Coddington 1996),
Cicadoidea (Pogue 1996), Mutilidae
(Quintero and Cambra 1996),
Carabidae (Erwin 1996),
Trichoptera (Flint 1996), and
Odonata (Louton et al. 1996). The
Lower Urubamba region was recently been the subject of a workshop on biological and cultural
diversity (Udvardy and Sandoval
1996), but no assessment of the

arthropod fauna was included.
Difficulties with site-intensive
arthropod studies in a tropical area
are several and important, the main
one being the large amount of
material that any intensive sampling
effort will produce in a very short
period of time (Table 1). Coupled
with this specimen abundance is the
taxonomic status of the various
103
Table 1. Arthropods sampled using different protocols(Cont.).

Taxon
Endomychidae
Erotylidae
Lampyridae
Lycidae
Nitidulidae
Phengodidae
Scarabaeidae
Scolitidae
Staphylinidae
Tenebrionidae
Psocoptera
Neuroptera
Chrysopidae
Megaloptera
Trichoptera
Lepidoptera
Plecoptera
Hymenoptera
Formicidae
Diptera
Immatures
Sweeping
General Winklers Pitfalls /Beating Light Baits Malaise Total
56
2
151
10
36
121
46
2
72
Araneae
19
Scorpionida
1
Amblypigi
2
Opilionida
3
Acarina
3
Pseudoscorpionida
Chilopoda
Diplopoda
Crustacea
8
Mollusca
Total
2443
208
63
1
15
10
6
25
408
609
11
1569
18
277
9
397
80
10
113
201
176
8
33
42
27
31
17
58
2
16
1
945
1118
2541
groups found in the survey. Many

arthropod groups are still very
poorly known, or the expertise to
identify them properly to species
level is wanting. A good number of
species are probably new to science,
particularly among the smaller
forms. One way to cope with these
challenges is to set up an assessment program using a simple but
flexible storage and retrieval sys104
1
5
23
140
50
295
4
5
1
12
23
797
4
25 75
*2500 2790
650
*4000 957
295
147
1
2
4
36
1
18
77
10
26
1
1
1
1
197
2025
16
20
2175
11444
tem, which could allow the incorporation of new data as it becomes

available. A well-curated reference
collection in the host institution is a
necessity for this endeavor, as is a
good logistic set-up for getting the
selected taxa to workers around the
world. Given the paucity of taxonomic expertise in many arthropod
groups, a world-wide network of
specialists needs to be established
for the final study and analysis of

this material.
In this paper, the results of a
preliminary survey on the arthropod
fauna in the area of the exploratory
wells in the Lower Urubamba
region is presented. The bulk of the
material is still under study, and the
preliminary results presented are
based on the division of material
into higher groups. This initial
sorting should be followed with a
detailed study of selected taxa,
where expertise is available or
which may be selected as indicator
groups for particular questions
related to the environment.
Methods
Arthropod sampling and preparation were made following standard methods for this group
(Oldroyd 1970, Martin 1977). The
strata selected for sampling were
the forest floor and leaf-litter layer
immediately above it, the understory layer, and the first layer of
vegetation. Special habitats were
also surveyed, including fallen logs
and fungi. The sampling techniques
selected were primarily passive; i.e.,
traps or devices that capture
arthropods without intervention by
the operator except for setting up
and servicing the trap. This was
necessary to make the sampling
effort repeatable for comparisons
among sites. This standardized
methodology should be the basis for
an arthropod sampling protocol
under development. Standardization
here refers to a set of criteria for
placing the traps or choosing the
right substrate, as well as using
devices of the same dimensions or
characteristics. The objective is to
minimize the sampling bias associated with more active, albeit less
repeatable, sampling activities.
Every collector has inherent subjective traits that will influence the
outcome of sampling and will make
comparison among samples by a
second collector less likely. Techniques are selected so that overlap
among samples is minimized for
maximum efficiency (Table 1).
Arthropods on the Forest
Floor and in Leaf Litter
Separation of small arthropods
found on the forest floor and in leaf
litter from the substrate is very time
consuming and inefficient if done
by hand. For these faunal elements,
two techniques were used: pitfall
traps and Winkler separators. Pitfall
traps consists of small, open containers buried in the forest floor up
to their openings. The mouth of the
trap should be level with the forest
floor, and the container should have
a killing and preserving solution.
Traps should be protected from
rainfall. Chance encounters of
moving arthropods on the forest
floor with these devices will result
in some of them falling onto the
trap. The containers used were
simple plastic cups of approximately 10 fluid ounces in capacity;
mouth diameter was 8 cm; the
preserving solution used was 70%
ethanol. A total of 40 such traps
were used at each of the 2 well
sites (San Martin-3 and Cashiriari2). Distance between traps was
about 10 m. Traps were left for 48
hours before servicing.
Sifter and Winkler separators
help remove the specimens from the
forest litter. They rely on the creation of an artificial environmental
gradient, usually moisture, against
which the arthropods will move. An
area of 1 m 2 is considered standard
for studies of soil arthropod communities. All litter and material is
Separation
of small
arthropods
found on
the forest
floor and in
leaf litter
from the
substrate is
very time
consuming
and inefficient if
done by
hand.
105
Sweeping
net and
beating
sheet
techniques
were used
to sample
understory
arthropods.
For flying
insects,
malaise,
butterfly,
and pan
traps were
used.
removed from this area and placed

on the sifter, which consists of a
cloth funnel with two rings placed
on the upper part, the lower ring
provided with a 5-mm wire mesh.
The litter is placed on top of the
mesh and vigorously shaken with
the aid of two wooden handles. The
smaller particles and arthropods are
sifted into the funnel where they are
trapped. Each sample is individually
tagged, bagged, and taken to the
field laboratory where the material
is placed in the Winkler separator.
This device consists of a
funnel-shaped cloth enclosure with
a suspended mesh bag inside. The
sifted material is placed inside the
mesh bag. A container with 70%
ethanol (such as a simple plastic
cup) is placed at the bottom of the
funnel to sample the falling
arthropods. Samples were left in the
Winklers for 48 hours. This method
is best suited for relatively mobile
organisms (i.e. ants), but given
enough time, slow animals will also
move away from the gradually
drying substrate and can thus be
sampled (Edwards 1991).
Understory Arthropods
Sweeping net and beating sheet
techniques were used to sample
understory arthropods. This is a
qualitative technique, somewhat
subjective as operator bias (e.g.,
operator preferences for handling
the net or sheet) will influence the
outcome. The sweeping net method
employs an insect net with a bag
made of heavy-duty cloth and a
strongly built ring and handle. The
net is vigorously swept against the
vegetation, arthropods fall into the
bag from the force of the impact,
and the specimens at the bottom of
the bag are transported to the
laboratory, where they are placed in
106
a cyanide killing jar or directly into

alcohol. The beating sheet technique uses an 80 x 80 cm piece of
cloth stretched over a x-frame that
can be improvised from thin,
straight branches. The sheet is
placed under a suitable habitat (a
brush, small branch, or any agglomeration of plant material separated
from the ground), and the target is
hit with a heavy stick. The force of
the impact will cause arthropods to
fall onto the clear surface of the
sheet, where they can be easily
picked up with forceps or an aspirator and placed directly into alcohol.
Flying Arthropods
For flying insects, malaise,
butterfly, and pan traps were used.
Some of these traps create mechanical barriers to flight, while
others employ attractants. Malaise
traps were used inside the
biodiversity plots to sample flying
insects through the lower strata of
the forest. The trap is designed to
intercept the insects and direct
them upwards to a confined space
where a collecting container with
70% ethanol awaits. Most insects
meeting the trap will fly up, go into
the chamber, and fall into the
alcohol. The technique provides a
large amount of material with little
effort, but the success of the
method depends on how long the
trap is left to work (Townes 1972).
Three to five traps were used per
plot; the sampling period until
service was 48 hours; and four
replicates or samples per trap per
plot were obtained.
The butterfly traps used consisted of a mesh cylinder with an
opening at the base. The trap was
usually placed about 5 m from the
ground, and suitable bait (usually
fermenting fruit) was placed at the
bottom of the cylinder. Butterflies

and other flying insects are attracted by this substance and enter
the trap though the opening at the
base. As they depart after feeding,
they usually will fly upwards and
become trapped in the cylinder. At
servicing, the traps are carefully
lowered and the insects inside are
sampled by hand. Five traps per
plot were used, and each trap was
serviced after 48 hours; four repetitions per trap per plot were done.
Pan traps are large containers
(pans) about 25 cm in diameter with
the interior surface painted bright
yellow. The purpose is to attract
visually cued insects flying through
the forest to the coloration of the
pan.
Light traps were used to sample
elements of the that are difficult to
sample through other techniques.
The success of such traps depends
on understanding the response of
many night flying insects that are
attracted by lights. Different light
wave lengths will produce different
results. White fluorescent light is
usually effective, but a better
option may be the so-called
black-light. We used both. We
simply placed a fluorescent light or
black-light tube in the middle of
the biodiversity plots with a portable power source (battery) and
positioned a white blanket or sheet
behind it to enhance light reflection
and provide a surface for insect
landings. The insects attracted to
the light were picked by hand and
placed directly into a killing jar
(Lepidoptera, Orthoptera) or into
70% ethanol. Samples were separated every 30 minutes over twohour periods, usually from 7:00 pm
to 9:00 pm. A total of three
samples every 48 hours were obtained using this method.
Arthropods Attracted to
Specific Substances: Baits
Many forest arthropods are
specialized to particular resources
and can thus be ecologically characterized. Among these are groups
that feed on dead animal bodies
(carrion feeders), mammal waste
materials (dung specialists), and
decaying or fermenting fruit (fermenting fruit specialists). Such
species can be sampled by setting
traps with the appropriate attractants. In this report, authors present
more detailed descriptions of these
traps and the results they produced.
Arthropods in
Specialized Habitats
Arthropods may be found in
specialized habitats such as fallen
logs in various stages of decomposition on the forest floor, fungi,
bromeliads, etc. At San Martin-3,
the bamboo internodes are hollow
and filled with water. When an
opening is made on the internode
wall by the activity of monkeys or
birds, the internode filled with
water provides suitable habitat for a
number of aquatic arthropods or
arthropods whose immature stages
require contained water for development. In most cases of specialized habitats, standardization is less
obvious; therefore it is very important to record particularities of the
sampling unit and the materials
used to extract the arthropods to
make comparisons possible. In case
of fallen logs, the type of wood and
its size and stage of succession as
well as factors such as humidity
and shade will influence the fauna
to be found.
General Sampling
Here, we refer to sampling that
is not considered part of the stan-
107
108
dardized methodology, but is

instead qualitative and not repeatable. It may be as basic as going
into the forest and seeking
arthropods in different microhabitats, walks though the forest
with a head lamp at night, etc.
Depending on the expertise, this
method can produce reliable data.
The amount of materials sampled is
usually less than with other
methods and the effort higher. In
this study, we considered sampling
of arthropods that were attracted to
camp lights as part of general
collection techniques.
of lot and sublot numbering was

used for specimen data recording
and retrieval, thus incorporating
identification data as it becomes
available and allowing for tagging
and further division of samples
down to individual specimens if
necessary. Data elements in the
system are designed to be compatible with international standards
and recommendations (Hawksworth
and Mound 1991). A more detailed
presentation of the data base
system and associated specimen
handling protocol is presented in
Appendix 1.
Sample Processing and

Data Base Management
Materials sampled and processed in the field were stored in
two ways: dry and in fluids. Storage
systems depend on the type of
organism, determination requirements, and collecting technique
employed. Materials best kept dry
are the Lepidoptera (including
butterflies and moths), the Odonata
(dragonflies and damselflies), and
some groups of Orthoptera (green
katydids, praying mantises). Alcohol
will fade the insects colors, so we
preserved samples in 70% ethanol
and stored them in whirl-packs.
This method is best suited for
handling specimens in the field, and
it offers several advantages over
more traditional glass or plastic
vials; e.g., the packs weigh less and
take up less space. Problems with
dehydration were minimal because
this was temporary storage. We
sorted and tagged all samples for
location, date of collection, and
sampling technique. This combination provides a unique code for
each sample and gives ready information on sampling technique,
habitat, and seasonality. A system
Preliminary Identification
and Deposit of Voucher
Material
Preliminary sorting and determination of sampled specimens was
attempted in the field. Sorting of
sampled material is a very timeconsuming and labor-intensive
activity when dealing with forest
arthropod samples. Completed in
the setting of a fully equipped
laboratory, this initial sorting to
ordinal level will make it possible to
target selected taxa for further
processing and study, while at the
same time yielding a gross estimate
of the arthropod biodiversity in the
study area. Voucher material will be
deposited in the collection at the
Marcos, Lima, Per and at the
Washington DC, USA.
Assessment of Arthropods:
Major Groups
The preliminary results of the
different sampling techniques are
presented in Table 1. The table
presents cumulative totals for each
of the samples. Numbers refer to

the actual number of specimens in
each sample for each taxonomic
group. Some of the major taxonomic groups are further divided
into the most distinctive taxonomic
units. This first sorting level gives a
rough estimate of the arthropod
community present in the study area
during the sampling period.
Note: For example, there are
only a few exceptions to the small
or very small size of land
arthropods. Included are the giant
tropical walkingstick, which can
reach 30 cm in length; aquatic bugs
of the genus Lethocerus, which can
attain a body size of 12 cm and
prey on small fishes and frogs;
species of beetles such as Dinastes
rhinoceros, which can have a body
length of 16 cm including the horn;
species of the African beetle
Megasoma, which are the heaviest of
all insects (up to 100 g in weight);
some South American nocturnal
moths of the genus Tasmania, with
wingspans up to 30 cm;
megalomorph spiders, whose legs
can span 30 cm and who prey on
small vertebrates; and tropical
centipedes that may have bodies 25
cm in length.
References
Dressler, R. L. 1984. Euglossine

bees (Hymenoptera: Apidae) of
the Tambopata Reserved Zone,
Madre de Dios, Per. Revista
Peruana de Entomologa 27: 75-79.
Edwards, C.A. 1991. The assessment of populations of
soil-inhabiting invertebrates.
Agricultural Ecosytems and Environment 34: 145-176.
Erwin, T.L. 1996. Natural history
of the carabid beetles at the
BIOLAT biological station, Rio
Manu, Pakitza, Per. Supplement

I. Additional records. Pages
of Southeastern Per. (D. Wilson
Horizonte/Smithsonian Institution
Press, Lima.
Fisher, E. M. 1984. A preliminary
list of the robber flies (Diptera:
Asilidae) of the Tambopata Reserved Zone, Madre de Dios, Per.
Revista Peruana de Entomologa 27:
25-36.
Flint, O.S., Jr. 1996. The
Trichoptera collected on the expeditions to Parque Manu, Madre de
Dios, Per. Pages 369-430 in
Manu: The Biodiversity of Southeastern Per. (D. Wilson and A.
Sandoval, eds.) Editorial
Press, Lima.
Gentry, A. H., ed. 1990. Four Neotropical Rainforests. Yale University Press, New Haven & London.
Hawksworth, D. L., and L. A.
Mound. 1991. Biodiversity databases: the crucial significance of
collections. Pages 17-29 in The
Biodiversity of Microorganisms
and Invertebrates: Its Role in
Sustainable Agriculture.(D.L.
Hawksworth, ed.) C. A. B. International, Wallingford, UK.
Lamas, G. 1984a. The Castniidae
and Sphingidae (Lepidoptera) of
Madre de Dios, Per: A preliminary list. Revista Peruana de
Entomologa 27: 55-58.
Lamas, G. 1984b. Los Papilionoidea
(Lepidoptera) de la Zona
Reservada de Tambopata, Madre
de Dios, Per. I: Papilionidae,
Pieridae y Nymphalidae (en parte).
59-74.
Louton, J. A., R. W. Garrison, and
O. S. Flint. 1996. The Odonata of
109
Parque Nacional Manu, Madre de

Dios, Per; natural history, species
richness and comparisons with
other Peruvian sites. Pages
of Southeastern Per. (D. Wilson,
Press, Lima.
Martin, J. E. H. 1977. Collecting,
preparing and preserving insects,
mites and spiders. The insects and
arachnids of Canada, Part 1.
Research Branch Canada Department of Agriculture Publication
1643.
Oldroyd, H. 1970. Collecting,
Preserving and Studying Insects.
Hutchinson, London.
Paulson, D. R. 1984. Odonata of
the Tambopata reserved Zone,
Madre de Dios, Per. Revista
Pearson, D.L. 1984. The tiger
beetles (Coleoptera, Cicindelidae)
of the Tambopata Reserved Zone,
Madre, Per. Revista Peruana de
Pogue, M.G. 1996. Biodiversity of
Cicadoidea (Homoptera) of
Pakitza, Manu Reserved Zone and
Tambopata Reserved Zone, Per:
a faunal comparison. Pages
Press, Lima.
Quintero, D., and T.R.A. Cambra.
1996. Contribucion a la
sistematica de las mutilidas
(Hymenoptera) del Per, en especial las de la estacion biologica
BIOLAT, Pakitza, Rio Manu.
Pages 327-358 in Manu: The
Biodiversity of Southeastern Per.
(D. Wilson and A. Sandoval, eds.)
Editorial Horizonte/Smithsonian
Institution Press, Lima.
110
Robbins, R.K., G. Lamas, O.H.H.

Mielke, D.J. Harvey, and M.M.
Casagrande. 1996. Taxonomic
composition and ecological structure of the species-rich butterfly
community at Pakitza, Parque
Nacional del Manu, Per. Pages
Press, Lima.
Silva, D., and J. Doddington. 1996.
Spiders of Pakitza (Madre de Dios,
Per): species richness and notes
on community structure. Pages
Press, Lima.
Townes, H. 1972. A light-weight
Malaise trap. Entomological News 83:
239-247.
Wilkerson, R. C., and G. B.
Fairchild. 1984. A checklist and
generic key to the Tabanidae
(Diptera) of Per with special
reference to the Tambopata Reserved Zone, Madre de Dios.
37-54.
Wilson, E.O. 1987. The little things
that run the world (the importance
and conservation of invertebrates). Conservation Biology 1:
344-346.
Manu: The Biodiversity of Southeastern Per. Editorial Horizonte/
Lima.
of the Lower Urubamba, Per.
Smithsonian Institution, Washington, DC.
Appendix 1. Database system developed for data recording and information retrieval for the arthropod specimens sampled in the Lower Urubamba
region.
Introduction
Site-intensive arthropod sampling at a tropical forest location
will produce large amounts of
material and associated data. Any
sampling program of arthropods
will entail a number of steps as
follows:
1. sample collection and specimen sorting,
2. preliminary determination
and further specimen sorting (species level) and preparation,
3. establishment of reference
collection for study site arthropods,
and
4. shipment of selected representative taxa for specialist identification and incorporation into
system.
Data management for each one
of these steps is based on a simple
yet powerful system. System design
should be continuous for all or most
of the steps in the process, transference of data should be very simple,
and repetition should be avoided.
This can be readily achieved with
present day technology and an
adequate design.
The following system has been
used and is still under development
for sampling arthropods (inventory
level) at the well sites in the Lower
Urubamba region. This brief presentation refers only to the first two
steps in the process. Please note
that the system involves entering
data into computers in the field.
Given the nature of magnetic media
and the weather hazards of various
study locations, extreme caution
should be taken to adequately
safeguard original data and back-up
copies. This system is intended to
replace most routine data recording

usually done by hand in field notebooks.
Sample Tagging and
Specimen Sorting
Specimens sampled are tagged
according to three criteria: locality
of collection, date of collection,
and sampling technique employed.
The principle of lots and sub-lots
was used: each sample from a
particular method constitutes a lot;
e.g., each of the 40 pitfall samples
corresponds to a particular lot for
that method. Different samples
(lots) obtained at the same locality
the same day and using the same
method are distinguished with
arabic numerals. This combination
provides a unique code that can be
associated with the specimens and
entered into the computer. At the
same time, it is expected the code
will give ready information on these
same criteria (site, date, sampling
method) without even having to
access the system.
Initial codes for each sample
consist of 1 letter, 3 digits, and 1
letter. An example is S331M, where
the first letter designates the site or
locality, the numbers indicate
month and day of sampling, and
the last letter indicates the sampling
method. Codes for site and sampling technique are: S= San Martin3; A= Cashiriari-2; G= general or
occasional sampling; L= light traps;
P= pan traps; M= malaise traps; F=
pitfall traps; W= Winkler samples;
S= sweeping and beating; C= baits;
B= berlese extractors (not used in
this phase); R= creek or stream;
111
Appendix 1. Database system developed for data recording (Cont.).

N=sampling at night on trails using
head lamps; X= special habitats.
Combination two of these
acronyms may be possible. However, restricting the codes for
sampling method to one letter is
still desirable. Therefore, the previous code reads as sample collected
in San Martin-3, May 31st, obtained
with a malaise trap. In cases of
multiple samples, a two-digit number is added at the end: S331M01,
where 01 corresponds to the
numeral given to each of the different samples obtained that day using
the malaise trap. After initial tagging and data entry, samples are
sorted to higher taxonomic groups,
each one representing a sublot,
which are distinguished from each
other by a consecutive numbering
system. Three digits are assigned for
sublot numbering (from 001 to
999), and a hyphen separates a lot
from a sublot; e.g., S331M01-003.
This constitutes the final code for
each of the sorted sublots. The
system allows for tagging of less
inclusive sublots down to species
level (each particular species from a
lot will become a sublot). Further
sorting of samples can be done at
any time, but the system should be
consulted before separating sublots
into species, etc. Each unique sublot code should provide easy access
to data associated with the material
and minimize the amount of work
for data entry. It should also provide the means to produce specimen labels with very little effort.
Database System Design
and Data Categories
The database system designed
for arthropod sampling consists of
three databases. The model is
112
relational, using a combination of

specimen-based data for samples
and taxon-based data for inventory
level. The particulars of system
design are related to the program
selected. Any commercially available relational database could be
used, depending on hardware
availability and resources. The
system used in this study was
Borlands Paradox, later transferred
to Claris FileMaker Pro.
A. Lots Database
This database consists of the
following fields:
Field Name, Field Type, Description, Lot/Sublot, Alphanumeric, Lot and sublot number,
Number, Number, Number of
individuals in sublot, Site, Text,
Locality name, Date, Sampling date,
Sampler, Text, Name(s) of
Sampler(s), Technique, Text, Sampling technique used, Habitat, Text,
Specific habitat or micro-habitat,
Plot No., Number, Plot number,
Quad. No., Number, Quadrat
Number, Prelim. ID, Text, Preliminary ID (should be linked to species
data base), ID. by, Text, Name of
investigator, Notes, Text/Memo.
Data common to each particular site or location may be kept in a
separate database, such as country,
province, specific locality, latitude
and longitude, elevation, etc. Key
fields should be lot/sublot (unique),
and several of the fields should be
set up for automatic entry when a
new record is added (site, date).
Use of pull-down menus or choice
lists is highly recommended (sampler, sampling method, person
determining sample). For the separation of processed sublots into less
inclusive groups (or individual
Appendix 1. Database system developed for data recording (Cont.).

species), the original record can be
duplicated and only relevant fields
changed.
B. Species Database
This database will keep the list
of individual species as they are
recognized and entered into the
system. The initial sorting level may
be to morphospecies. Codes for
species may be done by a four-letter
code for taxonomic group (class,
order, family) followed by species
number (example: Mant sp01). The
data base consists of the following
fields:
Field Name, Field Type, Description, Prelim. ID, Text, Preliminary ID, link to lots data base,
Genus, Text, Genus name, Species,
Text, Species name, Spauthor, Text,
Species author name, Class, Text,
Class name, Order, Text, Order
name, Family, Text, Family name,
Authority, Text, Authority for
taxonomic group, Notes, Text/
Memo, Miscellaneous notes.
The key field is Prelim. ID. This
allows linkage to the lots database,
so that queries based on recognized
species may be done and data
retrieval accomplished for identified
specimens.
C. Taxa Acronyms Database

This is an auxiliary database
consisting of a comprehensive list
of higher taxonomic groups with
four-letter made-up acronyms for
use in preliminary identifications.
Using species numerals and higher
taxonomic category acronyms
(usually families) provides a unique
system for species designation until
a more definitive identification is
available. The fields for this data
base are:
Field Name, Field Type, Description, Taxa, Text, Taxon acronym, Taxa name, Text, Full taxon
name, Class, Text, Class name,
Order, Text, Order name, Family,
Text, Family name, Notes, Text/
Memo, Miscellaneous notes, Empty,
Text.
113
114
Diurnal
Butterflies
(Lepidoptera:
Rhopalocera):
Biodiversity
Assessment
in the Lower
Urubamba
Region
Alfonso Alonso
Program (SI/MAB)
Gorky Valencia
Facultad de Ciencias Biolgicas,
Abad del Cusco
Introduction
Diurnal butterflies constitute

one of the best studied groups of
animals among the invertebrate
fauna. Adult butterflies are popular
with people because of their
brightly colored wings and their
observable daily activities that
include feeding on nectar as they
visit gardens. Indeed, butterfly
farming is becoming a source of
alternative income in many parts of
the world (Pyle 1981), including

southeastern Per (Farfan and
Valencia 1994). The number of
butterflies in a given area is
usually high; thus, people from
many cultures have carried out
studies in butterfly biology.
As with other insects, diurnal butterflies have a life cycle
that can be completed in a few
months. A butterflys life cycle
has four distinct stages, beginning with deposition of an egg
on a host plant by a female
butterfly. After a few days of
embryonic development, a
caterpillar (i.e., larva) hatches to
feed on the plant. The study of
caterpillars is fascinating since
they present different morphologies to reduce the potential of an
attack by predators. These range
from being cryptic (i.e., protective resemblance or camouflage)
to highly aposematic (i.e., colorful attraction combined with
defensive chemical compounds)
and from having spines and hairs
to being totally devoid of these
aspects. By feeding on plants,
caterpillars have to deal with the
115
After the
larval
feeding
and
growing
stage, the
butterfly
begins
metamorphosis to
produce
an adult
winged
butterfly.
116
chemical composition of the secondary compounds that plants produce to reduce herbivore attack.
Thus, caterpillar studies have
enriched our understanding of
animal/plant interactions, symbiotic
association, and the ecology and
behavior of parasitoids and predators (DeVries 1987, 1997).
After the larval feeding and
growing stage, the butterfly begins
metamorphosis to produce an adult
winged butterfly. When the caterpillar completes its feeding, it usually
leaves the host plant and wanders
to find a spot to form a pupa or
chrysalis. The pupa is immobile to
facilitate the intense physiological
change that the butterfly is experiencing. Upon emergence from the
pupa, the sexually mature stage of
the butterfly starts, in which the
insect is capable of flight, mating,
and reproduction, while it can not
develop further growth. As with all
insects, the body of an adult butterfly is composed of a head, thorax,
and abdomen. The wings are attached to the thorax; the antennae
and feeding parts are attached to
the head. Butterflies use their wings
to fly, but also to gain energy
(Opler and Malikul 1992).
Butterflies are highly dependent
on the microclimatic conditions of
the habitat where they live. As with
most insects, butterflies are coldblooded; they do not have an
internal mechanism to produce
energy to maintain their body
temperature at a constant level. To
fly and perform other daily activities such as nectar-feeding, egglaying, and mating, butterflies must
warm up their bodies to a certain
level. This can be easily accomplished by basking in the sun.
Thus, on sunny days many more
species of butterflies are active and
can be observed. Butterflies do not

fly on rainy days; they hide under
leaves or roost on tree trunks.
Diurnal butterflies play important roles in ecosystems because
they serve as plant pollinators and
plant feeders (i.e., herbivores) as
well as prey for hundreds of parasitoid species of wasps and flies that
feed on them. A parasitoid is an
organism that feeds only on one
host and kills it. In comparison,
parasites feed on portions of the
host but do not directly cause the
hosts death. Butterflies are also
preyed upon by other animals such
as birds and dragonflies (Brower
1984, Alonso and Marquez 1994).
Adult butterflies have different
ways to reduce predation. Chemical
compounds stored in their body
parts make them distasteful to
vertebrate predators. They may also
resemble other organisms that may
be toxic, and they have the capability to resemble structures and thus
blend with the environment (e.g.,
mimicking a dead leaf). The community of butterflies is highly
dependent on the healthy status of
an ecosystem. Given their strong
reliance on food plant availability
and microclimate conditions,
diurnal butterflies can serve as early
detectors of changes in ecosystem
functioning (Lamas 1986).
Within the scope of the general
biodiversity assessment in the
Lower Urubamba region, we studied
the community of butterflies at San
Martin-3 and Cashiriari-2. Our
principal objective was to document
the species present at these sites
and provide basic information
about the natural history of some
of the species. Our study is of
particular interest since the highest
diversity of butterflies in the world
has been documented in the upper
part of the Amazon River in Colombia, Ecuador, Per, and Brasil

(Emmel and Austin 1990, Robbins
et al. 1996). As other sites in
tropical areas are studied, it is likely
that new species will be found.
thorax. We then stored each specimen in a glassine envelope. These

specimens will be mounted and
deposited in the Natural History
Museum, Universidad Nacional
Mayor de San Marcos.
Methods
Baited Traps
These traps consist of a cylindrical net (0.90 m height, 0.20 m
diameter) placed above a platform
containing a bait. Most aerial
arthropods have the tendency to fly
upward after feeding or when they
are startled, except in a few instances when the insects drop to the
ground. Thus, traps are designed to
allow the butterfly to access the
bait, but then capture it as it flies
up after feeding. Baits usually
attract particular groups of butterflies, mostly within the
Nymphalinae and Satyrinae subfamilies. In the Lower Urubamba,
we set baits consisting of rotting
fruit (fermented papaya, banana,
pineapple) and rotten fish. We
placed six traps in the forest and
along trails. The traps were most
effective during the first two days.
After that, their effectiveness
decreased. The chemical composition of several plants can also serve
to attract butterflies. Most butterflies require specific chemical
compounds during their life cycle,
especially for reproductive activities
such as gonad and spermatophore
development. Pyrrolizidine alkaloid
(PA) producing plants such as
Heliotropium indicum (Boraginaceae)
have been used to attract
Ithomiinae butterflies (Medina et al.
1996, Robbins et al. 1996). PAs are
used for sexual courtship and
chemical defense.
The most effective ways to

study butterflies are to observe
them in their natural habitats,
sample them, take photographs of
them, and visit them in museum
collections (gathered from sampling). Sampling butterflies requires
a great deal of interest, dedication,
and funding, as well as knowing
how to capture, kill, and preserve
the insects.
Butterfly biologists have
followed the protocols described
below for many years. The protocols
have been improved over time, and
they are useful in sampling entire
communities of butterflies in an
ecosystem.
Hand Sampling with Nets
At Urubamba, we netted
butterflied with a circular green silk
bag, 60 centimeters (cm) in diameter and 90 cm deep, set on a pole 2
m in length. Butterflies are captured
in flight or while resting, feeding, or
drinking water. Once a butterfly
entered the net, we flipped the net
bag over to reduce the possibility of
escape. For butterflies perched on
the ground, we dropped the net
over them and held the tip of the
net in the air in such a way that the
insect flew or crawled up into the
bag. There are different ways of
killing the specimens. Most entomologists prefer to use killing jars
with ethyl acetate or to freeze the
butterflies if possible. We immobilized the thoracic muscles of the
butterflies by gently squeezing their
The most
effective
ways to
study
butterflies
are to
observe
them in
their
natural
habitats,
sample
them, take
photographs of
them, and
visit them
in
museum
collections.
117
Papilronidae
Pieridae 3%
Lycaenidae 5%
1%
Hesperidae
6%
Heliconiidae
28%
Riodanindae
9%
Ithominae
12%
Panainae
1%
Satyrinae Brassolinae
Charaxinae
Morphinae
10%
3%
3%
2% Apaturinae
1%
Nymphalinae
16%
Figure 1. Proportion of the genera found per family and subfamily of

diurnal butterflies in the Lower Urubamba region.
Finding and
Rearing Caterpillars
This method is useful but very
time consuming. Most butterflies
have a limited number of food
plants that they can feed on as
larvae. Once a caterpillar is found
feeding on the plant, it is taken to a
laboratory where it is reared. It is
best to sample caterpillars that are
about to pupate. It is also important
to take a viable sample of the plant
on which the caterpillar is found to
provide food for it. This protocol
aids in understanding the full life
cycle of butterflies. Butterflies are
usually known only from the adult
stage, but this protocol makes it
possible to record developmental
time and caterpillar behaviors.
Finding Dead Butterflies
Most predators of adult butterflies feed only on the soft tissues of
the organism. Thus, it is common
118
and relatively easy to find the

brightly colored wings of dead
butterflies on the ground. One
study (Alonso and Marquez 1994)
used this protocol to identify more
than 40 species that had been prey
for dragonflies in a tropical dry
forest in Costa Rica.
Taking Photographs
Taking pictures is also a good
way to document species and their
behaviors (some species, however,
can only be identified by dissecting
them). Butterfly photography is
growing in popularity and is an
effective method to amass a collection.
We conducted our sampling
from February-June, 1997, during
the wet/dry season transition. No
specimens were taken during rainy
days, when butterfly activity is
reduced. Our sampling with insect
nets was done near the system of
trails that were developed for
research at both sites, as well as

inside the biodiversity plots established for multi-taxa monitoring. We
also sampled near the opened area
of the platform operation center at
the well sites and along several
streams near our base camp.
We classified specimens into
morphospecies, a system that
separates different species based on
their morphology. In most groups of
invertebrates, specific names can
only be given by specialists. Our
specimens were also compared to
the plates published in DeVries
(1987, 1997) and Tyller et al.
(1994). Some specimens where also
compared with the butterfly collection of the Museo de Historia
Mayor de San Marcos.
Results and
Discussion

morphospecies of diurnal butterflies
Papilronidae
2% Hesperidae
Pieridae
5%
Lycaenidae 3%
2%
Riodanindae
13%
(Appendix 1). We found 134

Nymphalidae species, 22
Riodinidae, three Lycaenidae,
sixPieridae, three Papilionidae, and
eight Hesperiidae. The most diverse
subfamilies within the Nymphalidae
were the Nymphalinae with 25
species, Satyrinae with 49 and
Ithomiinae with 31 (Figs. 1, and 2).
Many species within the lycenid,
riodinid, and hesperiid butterflies
are mostly active during the transition period between the dry and the
wet seasons; hence, we did not
sample them. Most pierids appear to
be species of open habitats such as
tree-fall gaps and river edges. In our
study area, the drilling platform was
the only open area where this
species could have been trapped;
thus, its densities were very low. We
captured mostly satyrids in our
baited traps as well as several
species of nocturnal butterflies (i.e.,
moths; see Grados this volume).
The most common species captured
Heliconiidae
5%
We
sampled
a total
of 176
morphospecies of
diurnal
butterflies.
Nymphalinae
14%
Charaxinae
5%
Apaturinae
1%
Morphinae
2%
Brassolinae
3%
Ithominae
17%
Panainae
1%
Satyrinae
27%
Figure 2. Proportion of the species found per family and subfamily of

diurnal butterflies in the Lower Urubamba region.
119
We
sampled
more than
15% of
the
butterfly
species
expected
to be
found in
the area.
120
in the traps were Cissia cer. libye,

Chloreuptychia spp., and Taygetis spp.
One of the most intriguing
aspects of butterfly biology in
tropical areas is the similarity of
morphology and color among
unrelated groups of species. Some
butterfly species store toxic compounds that make them toxic to
potential predators. Other species
of butterflies that are palatable to
predators may come to resemble
toxic butterflies through natural
selection over time (e.g., the celestial blue of the genus Chloreuptychia
spp within the Satyridae, as well as
the darker patterns of Cissia and
Taygetis and the translucent and
color patterns of Oleria, Melinaea,
and Mesosemia).
We noticed a very different
community of butterflies when the
rainy season was over. Many species
of butterflies are closely linked to
environmental conditions in relation
to their activity patterns. In April
and May when the intensity of rain
decreased, we observed an increase
in the number of active butterfly
species. This may be the result of
adult butterflies moving into the
area as others disperse in search of
new areas to colonize. Or environmental changes (e.g., increased
sunlight reaching the ground) could
have encouraged the pupae to end
their dormant stage and emerge as
adults. Further studies are needed
to test these hypotheses.
We sampled more than 15% of
the butterfly species expected to be
found in the area. Sampling will
have to take place at the sites
during several seasons and over
several years to construct a picture
of total butterfly diversity in the
area. Robbins et al. (1996) found
the highest diversity1300 identified speciesof diurnal butterflies
registered in the world during five

trips in four years to Pakitza in
Manu National Park.
References
Alonso-Mejia, A., and M. Marquez.

1994. Dragonfly predation on
butterflies in a tropical dry forest.
Biotropica 26: 341-344.
Brower, L. P. 1984. Chemical
defenses in butterflies. Symposium
of the Royal Society of London 11:
110-134.
DeVries, P. 1987. The butterflies of
Costa Rica and their natural history. Volume I: Papilionidae,
Pieridae, and Nymphalidae.
Princeton University Press,
Princeton.
DeVries, P. 1997. The butterflies of
Costa Rica and their natural history. Volume II: Riodinidae.
Princeton University Press,
Princeton.
Emmel, T. C., and G. T. Austin.
1990. The tropical rain forest
butterfly fauna of Rondonia,
Brazil: species diversity and conservation. Tropical Lepidoptera 1:
1-12.
Farfan, M., and G. Valencia. 1994.
Caracteristicas generales de la
comercializacion de lepidopteros
en la zona de uso multiple
amazonico de la Reserva de la
Biosphera de Manu. Resumenes
del VII Congreso Iberoamericano
de Biodiversidad y Zoologia de
Vertebrados, Piura, Per.
Lamas, G. 1986. Mariposas, Ciencia
y Desarrollo. Medio Ambiente 13:
17-18.
Medina, M. C., R. K. Robbins, and
G. Lamas. 1996. Vertical stratification of flight by ithomiine butterflies (Lepidoptera: Nymphalidae)
at Pakitza, Manu National Park,
Per. Pages 211-215 in Manu: The

(E. Wilson and A. Sandoval, eds.)
Ed. Horizonte, Lima.
Opler, P. A., and V. Malikul. 1992.
Peterson Field Guide to the Eastern Butterflies. Ed. Houghton
Mifflin Co., New York.
Pyle, R. 1981. Butterflies: now you
see them. International Wildlife 11:
4-11.
Robbins, R K., G. Lamas, O. H
Mielke, and D. Harvey, and M. M.
Casagrande. 1996. Taxonomic
composition and ecological structure of the species-rich butterfly
community at Pakitza, Parque

Nacional del Manu, Per. Pages
of Southeastern Per. (E. Wilson
and A. Sandoval, eds.) Ed.
Horizonte, Lima.
Tyller, H., K. Brown, and K. Wilson. 1994. Swallowtail Butterflies
of the Americas: A Study in Biological Dynamics, Ecology, Diversity, Biosystematics, and Conservation. Scientific Publishers, Inc.,
Gainesville, Florida.
121
Appendix 1. Diurnal butterflies of the Lower Urubamba region

(morphospecies of the diurnal butterfliesOrder Lepidoptera: Division
Rhopalocerasampled and alphabetized within families and subfamilies;
butterflies were sampled in San Martin-3 and Cashiriari-2; the month column refers to the time when butterflies were sampled).
San Martin - 3
I) Nymphalidae
A) Heliconiinae
Dione af. juno
Dryas sp. 1
Heliconius sp. 1
Heliconius sp. 2
Heliconius sp. 3
Heliconius sp. 4
Heliconius sp. 5
Heliconius pos. demeter
Heliconius pos. sara
B) Nymphalinae
Nymphalinae sp. 1
Adelpha sp. 1
Adelpha sp. 2
Adelpha sp. 3
Batesia sp. 1
Callicore sp. 1
Callicore sp. 2
Callicore sp. 3
Colobura sp. 1
Diaethria sp. 1
Epiphile sp. 1
Eresia afin mechanitis
Eunica pos. excelsa
Hamadryas sp. 1
Historis dious
Marpesia sp. 1
Panacea prola
Pyrrhogyra sp. 1
Pyrrhogyra sp. 2
Pyrrhogyra sp. 3
Siproeta epaphus
Siproeta stelenes
Temenis sp. 1
Temenis af. laothoe
Tigridia sp. 1
C) Limenitidinae
D) Charaxinae
Hypna sp. 1
Hypna clytemnestra spp.
Memphis sp. 1
Memphis sp. 2
122
3
1
1
4
2
2
1
1
1
2
1
2
1
2
1
2
1
1
2
1
1
8
1
3
1
3
2
1
1
Cashiriari - 2
1
1
1
3
1
1
1
1
1
1
Month
April-June
May
February
April-June
April-May
April
May
Feb-March
March
April
April
April
June
May-June
May
April
June
June
March-April
February
April-May
May
June
March
May
May
February
April-May
June
March
Feb-April
March
May
April-May
March-May
May
February
April-June
Appendix 1. Diurnal butterflies of the Lower Urubamba region (Cont.).

San Martin - 3
Memphis sp. 3
Memphis sp. 4
Memphis sp. 5
Memphis sp. 6
Siderone sp. 1
E) Apaturinae
Doxocopa sp. 1
F) Morphinae
Antirrhea sp. 1
Morpho sp. 1
Morpho sp. 2
Morpho zephyritis
G) Brassolinae
Caligo sp. 1
Caligo illioneus
Caligo pos. memnon
Eryphanis sp. 1
Opsiphanes sp. 1
H) Satyrinae
Satyrinae sp. 1
Satyrinae sp. 2
Amphidecta sp. 1
Amphidecta cer. pignerator
Cissia sp. 1
Cissia sp. 2
Cissia sp. 3
Cissia sp. 4
Cissia sp. 5
Cissia sp. 6
Cissia sp. 7
Cissia sp. 8
Cissia sp. 9
Cissia sp. 10 A
Cissia sp. 11 B
Cissia sp. 12 C
Cissia sp. 13 D
Cissia sp. 14 E
Cissia sp. 15 F
Cissia sp. 16
Cissia sp. 17
Cissia sp. 18
Cissia cer. libye
Cissia cer. tiessa
Cithaerias aurorina
Chloreuptychia sp. 1
2
1
2
2
5
2
1
Cashiriari - 2
1
1
Month
May
May
June
June
April-May
June
April
April-June
April
May-June
1
1
1
1
6
February
April
April
April
March-May
2
1
1
April
April
April
April
Feb-March
Feb-March
March
April-May
April
April-May
April
April-May
April-May
April-May
May
April-May
April-May
April-June
April-May
June
June
June
April-May
April-May
April
Feb-May
April-May
April
2
2
1
2
6
5
2
3
3
3
1
2
4
9
4
1
25
3
3
16
4
2
3
1
2
123

San Martin - 3
Cithaerias aurorina
Cyllopsis sp. 1
Cyllopsis sp. 2
Euptychia sp. 1
Euptychia sp. 2
Haetera sp. 1
Pierella sp. 1
Pierella sp. 2
Pierella sp. 3
Pierella sp. 4
Taygetis sp. 1
Taygetis sp. 2
Taygetis sp. 3
Taygetis sp. 4
Taygetis sp. 5
Taygetis sp. 6
Taygetis sp. 7
Taygetis sp. 8
Taygetis sp. 9
Taygetis cer. penelea
I) Danainae
Eutresis sp. 1
J) Ithomiinae
Ithomiinae sp. 1
Ithominae sp. 2
Ithominae sp. 3
Ithominae sp. 4
Episcada sp. 1
Episcada sp. 2
Episcada sp. 3
Greta sp. 1
Heterosais sp. 1
Hypoleria sp. 1
Hypoleria virginia
Hypoleria vitiosa
Hypothyris sp. 1
Ithomia sp. 1
Melinaea sp. 1
124
3
16
4
2
1
4
2
1
3
1
1
1
12
9
5
1
1
1
2
8
9
1
1
1
Cashiriari - 2
Month
April
Feb-May
April-May
April
April
April-May
April-May
April
April-May
April
April-May
March
February
April-May
April-May
June
Feb-March
March
March
March
April-May
April-May
April-May
May
April
March
1
1
April-May
February
May
June
June
April
April-May
April
June
June
April
February
April
June
April
1
3
1
1
3
1
1
1
1
1
1
1

San Martin - 3
Melinaea sp. 2
Melinaea sp. 3
Melinaea sp. 4
Mechanitis sp. 1
Mechanitis sp. 2
Oleria sp. 1
Oleria sp. 2
Oleria sp. 3
Oleria sp. 4
Oleria cer.denuta
Oleria denuta ssp.
Pteronymia sp. 1
Pteronymia sp. 2
Pteronymia sp. 3
Rhodussa sp. 1
Scada sp. 1
2) Riodinidae
Riodinidae sp. 1
Riodinidae sp. 2
Riodinidae sp. 3
Emesis mandana ssp.
Eurybia elvina
Eurybia pos. unxia
Isapis sp. 1
Mesosemia sp. 1
Mesosemia sp. 2
Mesosemia sp. 3
Mesosemia sp. 4
Mesosemia sp. 5
Mesosemia sp. 6
Mesosemia sp. 7
Napaea sp. 1
Napaea sp. 2
Nymphidium sp. 1
Nymphidium sp. 2
Parcela sp. 1
Synargis sp. 1
Synargis sp. 2
Theope sp. 1
3) Lycaenidae
Lycaenidae sp. 1
Thecla sp. 1
Thecla sp. 2
Cashiriari - 2
1
5
1
1
1
21
1
2
1
3
1
1
1
1
4
4
1
1
1
1
1
1
1
2
1
21
2
2
1
1
1
1
3
1
1
2
1
2
1
1
Month
February
April
April-May
April
May
April-June
April-May
April
April
Feb-March
June
February
April-June
April
February
April
February
February
February
April
May
April
May
April-May
April-May
April-May
April
April
May
May
April-May
April
May
June
May
April
April
June
May
April
June
125

San Martin - 3
4) Pieridae
Aphrissa sp. 1
Appias sp. 1
Eurema sp. 1
Perrhybris sp. 1
Perrhybris af. pyrrha
Phoebis argante argante
5) Papilionidae
Battus polydamas sim athyamus
Eutresis sp. 1
Heraclides thoas cyniras
6) Hesperidae
Hesperidae sp. 1
Acolastus sp. 1
Acolastus sp. 2
Cobalus sp. 1
Heteropia sp. 1
Proteides sp. 1
Thymele sp. 1
Timochares sp. 1
126
1
2
1
Cashiriari - 2
1
1
1
1
1
May
June
June
April
February
May
April
April
1
1
2
1
1
1
1
1
Month
March
February
June
May
April
June
June
May
Nocturnal
Butterflies
(Lepidoptera:
Heterocera:
Ctenuchinae):
Biodiversity
Assessment
in the Lower
Urubamba
Region
Juan Grados
Mayor de San Marcos
Introduction
Systematic biological assessments in southeastern Per have

found a great invertebrate diversity
that has not been reported in any
other neotropical site or the world
(Wilkerson and Fairchild 1985,
Fisher 1985). Of the invertebrate
groups, the butterflies are the best
known group in Per even though
they have not been studied in more
than half of that country (Lamas
1989a). In Tambopata and Manu
National Park, two of the most-
studied tropical areas in Per, a

great diversity of diurnal butterflies
has been found1249 species have
been reported at Tambopata (Lamas
personal communication), while
1300 species have been reported at
Manu (Robbins et al. 1996). Many
of those species were new records
to science.
Nocturnal butterflies (i.e.,
moths) have been studied less, but
they are also very diverse. Among
these, the Arctiidae group is primarily distributed in the neotropics.
The Ctenuchinae, a subfamily of
the Arctiidae, has many members
with diurnal activity and morphological characteristics that make
them resemble wasps (Scoble
1992). Only a few studies have
focused on nocturnal butterflies in
Per. Lamas (1985, 1989b) studied
the Castniidae, Saturniidae, and
Sphingidae families in southeastern
Per; Yabar (1989) sampled nocturnal lepidoptera in Sahuayaco (La
Convencion, Cusco) when studying
the Plusiinae (Noctuidae). Still,
there are no significant samples of
nocturnal lepidoptera for the Lower
Urubamba region. This paper
127
discusses the assessment of nocturnal butterflies in that area.
Methods
It is
estimated
that 150
species of
Ctenuchinae
can be
found in
this area,
especially
if sampling
is done
year round
in both wet
and dry
seasons.
The study was conducted at the

San Martin-3 well site, dominated
by bamboo (Guadua sarcocarpa).
Sampling was done over 18 days
(March 18-April 8, 1997) at the end
of the wet season. Since many
Ctenuchinae are attracted to light, I
used light traps with a sheet for
sampling (mercury lights for 6 days
and ultraviolet lights for 12 days).
Traps were set at different sites to
optimize opportunities to sample a
higher diversity of nocturnal butterflies; included were the tops of hills
near the camp and primary forest.
Specimens attracted to the lights
were captured and placed in a lethal
chamber. Some specimens were
sampled during the day.
Primary Forest
At the beginning of the sampling period, I sampled the eastern
part of the biodiversity plot #1
near a 4-m wide stream. Only a few
species were captured because the
ambient temperature was relatively
low and the night skies were clear.
Primary Forest
Associated with Bamboo
Light traps were placed in the
high spots at the limits of the
cleared area near the camp, where
G. sarcocarpa dominates. The higher
location of these traps allowed the
light to travel further than the traps
in primary forest, and many more
individuals were attracted. Most of
the specimens were sampled at
these locations.
128
Results and
Discussion
During the 18 days of field

work, I sampled more than 600
specimens; I have identified 87
species to date (Appendix 1). Only
four species were captured during
the dayHistiaea proserpina
(Hbner 1827), Bodosa tina (Walker
1854), Agyrtidia uranophila (Walker
1866), and Androcharta meones
(Cramer 1780).
The fact that I was able to
sample 87 species in 18 days
coupled with the fact that only 103
species of Ctenuchinae are known
to date for nearby Tambopata
(Grados unpublished data) indicates
that the Lower Urubamba region
could be one of the most speciose
parts of Per. It is estimated that
150 species of Ctenuchinae can be
found in this area, especially if
sampling is done year round in both
wet and dry seasons. To optimize
the capture of these butterflies, it is
necessary to use natural and artificial baits within the primary forest,
where many species can be found
that are not attracted to lights. It is
also important to sample adjacent
areas near the camp.
References
Fisher, E. 1985. A preliminary list

of the robber flies (Diptera:
Asilidae) of the Tambopata Reserved Zone, Madre de Dios, Per.
25-36.
Lamas, G. 1985. The Castniidae
and Sphingidae (Lepidoptera) of
Madre de Dios, Per: A Preliminary List. Revista Peruana de
Lamas, G. 1989a. Un estimado del

grado de cobertura geogrfica de la
colecta de mariposas (Lepidoptera)
en el Per. Revista Peruana de
Lamas, G. 1989b. Lista preliminar
de los Saturniidae, Oxytenidae,
Uraniidae y Sematuridae (Lepidoptera) de la Zona Reservada de
Tambopata, Madre de Dios, Per.
57-60.
Robbins, R., G. Lamas, O. Mielke,
D. Harvey, and M. Casagrande.
1996. Taxonomic composicion and
ecological structure of the species
of the rich butterfly community at
Pakitza, Parque Nacional del
Manu, Per. In Manu: The
(D.E. Wilson and A. Sandoval,
eds.) Editorial Horizonte, Lima.
Scoble, M. J. 1992. The Lepidoptera: Form, Function and

Diversity. Oxford University Press,
London.
Yabar, E. 1982. Contribucin al
Conocimiento de los Plusiinae
(Lepidoptera: Noctuidae) del
Cusco (Per). Revista Peruana de
Wilkerson, R., and G. Fairchild.
1985. A checklist and generic key
to the Tabanidae (Diptera) of Per
with special reference to the
Tambopata Reserved Zone, Madre
de Dios. Revista Peruana de
129
Appendix 1. The Ctenuchinae in the Lower Urubamba region (specimens

from the San Martin-3 well site, sampled March 18-April 8, 1997).
Pseudosphex sp.
Sarosa acutior (Felder, 1874)
Isanthrene sp.
Autochloris cf. cuma (Druce, 1897)
Phoenicoprocta analis (Schrottky, 1909)
Phoenicoprocta cf. sanguinea (Walker,
1854)
Loxophlebia pyrgion (Druce, 1884)
Loxophlebia triangulifera (R. Felder,
1869)
Cosmosoma flavothorax (Rothschild,
1910)
Cosmosoma remotum (Walker, 1854)
Cosmosoma stibosticta (Butler, 1876)
Cosmosoma telephus (Walker, 1854)
Cosmosoma oroyanum (Rothschild, 1911)
Cosmosoma cf. phoenicophora (Dognin,
1909)
Cosmosoma sp. 1
Cosmosoma sp. 2
Chrostosoma decisa (Walker, 1864)
Chrostosoma cardinalis (Schaus, 1898)
Lepidoneiva erubescens (Butler, 1877)
Nyridela chalciope (Hubner, 1827)
Leucotmemis varipes (Walker, 1854)
Mystrocneme varipes (Walker, 1854)
Dixophlebia quadristrigata (Walker, 1864)
Psoloptera thoracica (Walker, 1854)
Histiaea proserpina (Hubner, 1827)
Bodosa tina (Walker, 1854)
Agyrtidia uranophila (Walker, 1866)
Macrocneme sp. 1
Calonotus sp.
Hypocladia militaris (Butler, 1877)
Ecdemus hypoleucus (Herrich-Schaffer,
1854)
Ecdemus pereirai (L. Travassos, 1940)
Teucer carmania (Druce, 1883)
Teucer glaucopis (R. Felder, 1869)
Episcepsis capyscoides (Dognin, 1911)
Episcepsis demonis (Druce, 1896)
Episcepsis cf. lenaeus (Cramer, 1780)
Episcepsis scintillans (Rothschild, 1911)
Episcepsis sp. 1
Episcepsis sp. 2
Androcharta diversipennis (Walker, 1854)
Androcharta meones (Cramer, 1780)
Pseudaclytia opponens (Walker, 18649)
130
Athyphopsis modesta (Butler, 1878)

Napata albimaculata (Hampson, 1901)
Napata walkeri (Druce, 1889)
Schasiura mimica (Butler, 1877)
Osmocneme bracata (Draudt, 1915)
Patreliura capys (Cramer, 1775)
Loxozona nitens (Rothschild, 1912)
Sciopsyche tropica (Walker, 1854)
Tipuloides rubriceps (Dognin, 1912)
Aclytia heber (Cramer, 1780)
Aclytia reducta (Rothschild, 1912)
Aclytia cf. petra (Schaus, 1911)
Epidesma hoffmannsi (Rothschild, 1912)
Epidesma klagesi (Rothschild, 1912)
Epidesma ursula (Stoll, 1781)
Neacerea lemoulti (Draudt, 1915)
Neacerea macella (Dognin, 1911)
Eucereum facundum (Draudt, 1915)
Eucereum leria (Druce, 1884)
Eucereum maricum (Cramer, 1775)
Eucereum maja (Druce, 1884)
Eucereum marmoratum (Butler, 1877)
Eucereum metoidesis (Hampson, 1905)
Eucereum obscurum (Moschler, 1872)
Eucereum pagina (Hampson, 1898)
Eucereum cf. pseudoarchias (Hampson,
1898)
Eucereum punctatum (Guerin, 1844)
Eucereum pyrozonum (Hampson, 1911)
Eucereum quadricolor (Walker, 1855)
Eucereum cf. scyton (Cramer, 1777)
Eucereum setosum (Sepp., 1830)
Eucereum varium (Walker, 1854)
Eucereum sp. 1
Eucereum sp. 2
Eucereum sp. 3
Eucereum sp. 4
Correbia lycoides (Walker 1854)
Correbia cf. minima (Druce 1903)
Correbia sp. 1
Correbidia calopteridia (Butler 1878)
Correbidia cf. terminalis (Walker 1856)
Hyaleucerea erythrotelus (Walker 1854)
Hyaleucerea gigantea (Druce 1884)
Hyaleucerea vulnerata (Butler 1875)
Ants
(Hymenoptera:
Formicidae):
Biodiversity
Assessment
in the Lower
Urubamba
Region
Leeanne Alonso
Conservation International,
Washington, DC
Alfonso Alonso
Program (SI/MAB)
Introduction
Traditionally, biological assessments and monitoring programs

have focused on large organisms
such as plants, birds, and mammals
and have devoted little attention to
the small creatures (Wilson 1988).
However, invertebrates are by far
more speciose and numerically
dominant and contribute more
biomass to ecosystems than the
larger taxa (Wilson 1987, Gaston
1991). Conservation planning can
benefit greatly by including the
diverse invertebrate group, which
occupies a great number of functional niches and microhabitats

throughout the world. There is great
potential for the use of arthropods
in conservation planning (Kremen
et al. 1993). Among other reasons,
invertebrates are ideal because they
are readily observed, easily
sampled, and often can be monitored as sensitive indicators of
environmental change.
The family Formicidae, the
ants, is particularly well suited for
assessment and monitoring programs. Ants are ideal bioindicators
because they are diverse (approximately 9500 described species in 13
subfamilies) and found in abundance in almost every habitat in the
world (Holldobler and Wilson
1990). Ants are numerically dominant in many tropical forest canopy
studies, comprising 19% to 50% of
the individual arthropods (Basset et
al. 1992). A study in the Brazilian
Amazon found ants contributed
80% of the insect biomass and
more than 30% of all animal biomass (Fittkau and Klinge 1973). Ants
are important ecologically because
they function at many levels in an
131
Ants are
important
ecologically
because
they
function at
many levels
in an
ecosystem
as predators,
prey, and
detritivores.
Ants also
have diverse
associations
with plants;
they are seed
dispersers,
protectors,
and
herbivores.
132
ecosystemas predators, prey, and

detritivores. Ants also have diverse
associations with plants; they are
seed dispersers, protectors, and
herbivores.
Since ants are common and
apparent in almost every habitat,
they are readily sampled. Several
sampling techniques can be used
depending on the purpose of the
project, the habitat, and available
resources. Ants are not as hyperdiverse as some other insect groups
(i.e., Coleoptera), so they can be
sorted easily to morphospecies by
parataxonomists and then identified
with available keys and the assistance of systematists. Many ant
species are highly sensitive to
microclimate and habitat structure
and thus respond rapidly to environmental changes (Majer 1983,
Andersen 1990, Bestelmyer and
Wiens 1996).
Since they are social insects,
ants live in colonies that range from
a few to several million sterile
worker ants serving one to several
queens. The social structure of ant
colonies allows sampling of numerous worker ants without significantly affecting the reproductive
unit of the colony. Ant species vary
greatly in their nesting and feeding
requirements. They can be found
living in all strata of the environmentarboreally in the forest
canopy and lower vegetation,
terrestrially in the leaf litter and
rotting wood on the ground, and
subterraneanly in the soil. If the
goal is to inventory all ant species
in an area, different methods must
be used to sample each of these
strata. Ants feed on a diverse array
of foods; some are primarily
nectivorous, feeding at floral and
extrafloral nectaries of plants or at
homoptera exudates, while others
are strictly predatory. Some specialize on termites or other ants. Many

ant species are omnivorous, feeding
opportunistically on any live or
dead insect they encounter and
occasionally on plant material
(nectar, seeds, or tissue). The
feeding habits of ants can be exploited to sample them at baits,
especially because most ant species
recruit nest mates to food sources
so that numerous ants can be
sampled or followed to locate the
nest.
Two standardized, quantitative
ant-sampling methods (described in
more detail under Methods below)
were used at San Martin-3 to
sample the leaf-litter and grounddwelling ant fauna. The first
method consisted of intensive
searching in 1-m2 plots of leaf
litter to sample ant nests located in
the plots. Sampling nests allows for
analysis of the abundance of ants at
the site (see Data Analysis below).
The second sampling method was
recently developed by a group of
biologists, specializing in the study
of ant taxonomy and ecology, as a
standardized protocol for sampling
soil and leaf- litter ants (Agosti et
al. In preparation). The protocol is
designed to allow inclusion of ants
and other ground-dwelling
arthropods in biodiversity inventory
and monitoring programs. It incorporates two standardized sampling
methodsleaf-litter extraction and
pitfall trapswhich are easy to
implement.
Both of the sampling methods
used in this study are quantitative
and can be repeated over time,
which makes them appropriate for
monitoring. Future sampling of ants
in the same location using these
methods can help detect and moni-
tor any change in the ant community at the biodiversity plot.
Methods
The richness and abundance of

ground-dwelling ants was investigated in the forest near the San
Martin-3 camp from March 8-18,
1997. Ants were sampled in
biodiversity plot #1(Alonso et al.
this volume), located approximately
500 m downhill from the camp.
Intensive Sampling of Ant
Nests in Leaf Litter and Soils
Ant nests were sampled using
an intensive method that samples
both ant nests and stray, foraging
ants. Two 150-m transects were
established across the biodiversity
plot. Sampling points were located
every 10 m along the transects, for
a total of 30 points. A 1-m2 quadrat
was marked out at each point and
all leaf litter, twigs, and rotting logs
in the plot were examined for ants.
Cookies were then scattered over
the plot to attract ants so that they
could be followed back to their
nests. All ant colonies and stray
ants found between leaves in the
litter, in twigs, or in the soil were
sampled with forceps and preserved
in vials of 70% ethanol.
Standardized Protocol for
Ground-dwelling Ants
Ants were sampled using the
standardized protocol at 10-m
intervals along four 100-m transects
(10 points per transect). Two
transects were located inside the
biodiversity plot, and two were
located outside of the plotone
aligned parallel to the northern side
of the plot and the other aligned
parallel to the western side. At each
sampling point, two sampling
techniques were employed. First
was leaf-litter sifting (with later

extraction using Winkler sacks). A
1-m 2 area of leaf litter (including
twigs and rotting branches) was
sampled at each point and sifted to
remove all large leaves and pieces
of wood. The sifter consisted of a
1-cm 2 wire mesh screen enclosed in
a cotton sack with two handles for
shaking. The sack was tied off at
the bottom to retain the sifted litter
and then opened to move the litter
to a bag for transport to the laboratory, where the contents were
carefully transferred to a cloth mesh
bag over a white plastic sheet or
tray. Any ants escaping during this
process were sampled and preserved in vials of alcohol. The
mesh bag containing litter was
placed into a Winkler sack and
attached by clips to the top wire
frame of the sack. The sack was
then closed at the top, with a cup
of alcohol tied at the bottom, and
hung vertically for 48 hours. The
idea is to allow active arthropods
such as ants to move through the
litter and out of the mesh bag. They
then fall to the bottom of the
Winkler sack where they are preserved in the cup of alcohol (Fig.
1). After 48 hours, all ants we
trapped and found in the cup were
removed and preserved in vials of
alcohol.
The
richness
and
abundance
of grounddwelling
ants was
investigated in
the forest
near the
San
Martin-3
camp from
March
8-18, 1997.
Pitfall Traps
At the same corner of each 1m2 quadrat, a pitfall trap was set by
digging a small hole and then
placing a 10-oz plastic drinking cup
into the hole. Soil and leaf litter
were replaced around the cup so
that its top was even with the soil
surface. Approximately 3 cm of
alcohol were poured into each cup
as a preservative. A large leaf was
laid over the cup to prevent rain
133
3 cm diameter metal ring

string
cm
27
16x27 cm metal wire support
27x36 cm mesh bag,

0.4 mm grid size,
to hold leaf liter
41cm
16x27 cm metal wire support
7.5 cm diameter metal

ring to funnel cloth
100% cotton cover
6.35 cm diameter metal ring

plastic cup and
plastic bag attached
to metal ring
27cm
Figure 1. Diagram of the Winkler sack used for extracting ants from leaf litter.
from entering. Pitfall traps sample
active ants and other arthropods as
they move along the soil and litter
surface and then fall, unaware, into
the cup. The pitfall traps were left
out in the field for 48 hours, then
sampled. Any arthropods in the
cups were removed and preserved
in vials of alcohol.
Data Analysis and
Species Identification
Analyses of species diversity
are difficult for social insects such
as ants because the number of
individuals sampled does not
necessarily reflect the abundance of
species, which is needed for diversity indices. This occurs because
some ant species with large colony
sizes may recruit many workers to a
bait or send a lot of workers out
foraging, while other ant species
forage solitarily. For ants and other
social insects, abundance is best
134
measured as the number of nests

(the reproductive unit) of each
species in an area (as determined by
the intensive sampling method). If
nests are not sampled, the frequency of capture, as measured by
the number of samples or traps in
which a species is found, can be
used as an estimate of abundance.
The taxonomy of neotropical
ant species is not well developed,
with many ant species yet to be
named and described in the literature. Therefore, the ants sampled in
this study have been identified to
genus and morphospecies only until
they can be compared with specimens in museum collections.
Results and
Discussion
The genus Pheidole was the

most speciose in this sample, with
17 species (Appendix 1). This genus
is also one of the most speciose
world wide, with more than 600
species (Wilson and Brown In
preparation). The genera
Paratrechina (six species, subfamily
Formicinae) and Solenopsis (five
species, subfamily Myrmicinae)
were also fairly diverse, as they are
throughout the neotropics. Eight
(44%) of the 18 genera were represented by only one species.
The 55 ant species were found
in a mean of 2.2 species per quadrat
(SD=3.4, median= 2, range 1-10
quadrats). While a few species were
fairly common, 27 (49%) of the 55
species were sampled in only one
quadrat (Fig. 2). This indicates that
either these species are not common
in the forest near San Martin-3 or
were inadequately sampled. Further
sampling in the area should provide
more information.
Pheidole sp. 1 was the most
widespread ant species, nesting in
10 of the 30 quadrats (Appendix 1).
Five ant species were found in five
or six quadrats: Adelomyrmex sp. 1,
Intensive Sampling
of
Ant Nests
Species richness. Nests of 55
ant species were sampled from the
30 1-m 2 quadrat using the intensive
sampling method. Of the six ant
subfamilies that occur in South
America, four were represented by
these 55 species (Appendix 1). Not
surprisingly, this method of sampling leaf-litter and soil-nesting ants
did not sample members of the
subfamily Ecitoninae, the nomadic
army ants, or the subfamily
Pseudomymecine, which are almost
exclusively arboreal nesters. The
subfamily Myrmecine was represented by the most species (36), the
Ponerinae by 11 species, the
Formicinae by seven species, and
the subfamily Dolichoderinae by
only one species (Appendix 1).
These results reflect the general
pattern of diversity within ant
subfamilies in the New World, with
Myrmicinae the most speciose
subfamily overall (Bolton 1996).
30
# of Species
25
20
15
10
5
0
1
10
# of Plot
Figure 2. Frequency of sampling (# 1-m 2 plots) of ant species

using an intensive sampling method at the biodiversity plot at San
Martin-3.
135
8
7
# of Plots
6
5
4
3
2
1
0
1
10
11
# of Ant Species
Figure 3. Density of ant nests in 30 1-m 2 quadrats at the

biodiversity plot at San Martin-3.
Crematogaster sp. 1, Gnamptogenys sp.
1, Paratrechina sp. 1, and Prionopelta
sp. 1. (Fig. 2, Appendix 1). Little is
known about the biology of these
species, but our results indicate that
they are fairly common leaf litter
and soil nesters in the area. As
predators and scavengers, these
species likely have a substantial
impact on the arthropod fauna in
this microhabitat.
Ant density. A mean of 4.3
(SD = 2.2, median = 4, range 2-11)
ant nests were found per 1-m 2
quadrat (Fig. 3) for a total of 120
nests in 30 m 2 . There was one nest
per species per quadrat, so the
density is equal to the richness per
quadrat. While 13 (43%) of the 30
quadrats had only two or three ant
nests, one quadrat had nine nests
and another had 11 nests (Fig. 3).
This ant density indicates there
is a fairly high density of ant nests
in the forest at San Martin-3. With a
mean of 4.3 nests per m2 , we can
estimate 43,000 ant nests per ha.
The number of worker ants per nest
varies greatly between species;
some species have nests of 10 ants,
while others have nests of over a
million individuals. Therefore, it is
136
difficult to estimate the number of

individual ants per area, but it is
likely that their numbers and biomass significantly exceed that of all
other animal species. Given their
importance in the ecosystem
through their various roles, ants are
certainly an essential element in the
functioning of the forest at San
Martin-3.
Estimates of species richness.
The 30 quadrats sampled under the
intensive sampling method were not
sufficient to sample all ant species
in the leaf litter and soil. With each
new quadrat sampled, more species
were added (Fig. 4). No new species
were added from quadrats 27
through 30, but this leveling off of
the species accumulation curve can
not assure us that no new species
will be added with additional
sampling. Furthermore, singleton
speciesthose found in only one
quadratwere continuously added
as more quadrats were sampled,
indicating that more rare species
will likely be sampled with each
additional quadrat. This type of
species accumulation curve (Fig. 4),
which does not level off, tells us
that there are more species in the
area than we were able to sample

and that more sampling is needed.
From these results, it is difficult to estimate the number of ant
species that are likely in the area of
San Martin-3 or in the biodiversity
plot. The intensive sampling
method samples a large portion of
the leaf-litter and soil-nesting ant
species, but it does not come close
to sampling all the ants in the area.
The species richness of ants in
other neotropical areas has been
found to be more than 600 species
in a 100-ha lowland rain forest in
Costa Rica (J. Longino personal
communication); 156 species were
found in lowland rain forest in
Brazil (Delabie and Majer 1996);
and more than 150 species were
found in the Tombopata region of
Per (S. Cover personal communication). One reason that we found
considerably fewer ant species in
this study is that we focused only
on the leaf-litter and soil ant fauna.
The tree canopies of tropical
forests contain a high diversity of
ant species not found on the forest
floor. Therefore, different sampling
methods should be used to sample

in different strata of the forest.
Results from the leaf-litter extraction with Winkler sacks and the
pitfall traps will likely add more
species and provide a more accurate
total of ground-dwelling ant fauna.
Future sampling using canopy
fogging and/or vegetation techniques will also provide a more
complete faunal list for the forest.
We wish to thank Shell Prospecting and Development (Per)
B.V. for funding this research and
for their awareness of the need to
protect biodiversity and their
actions towards that end . We thank
T. Drisckell and G. Valencia for
helping carry out the standard
protocol. L. Alonsos participation
was partially supported by a
National Science Foundation grant
to M. E. Kaspari.
The tree
canopies
of tropical
forests
contain a
high diversity of ant
species not
found on
the forest
floor.
60
# of Plots
50
40
30
20
10
# of species
singletons
0
1
11
13
15
17
19
21
23
25
27
29
Sampling Efforts (Plots)
Figure 4. Species accumulation curve of ant species in 30 1-m 2 quadrats

using an intensive sampling method at the biodiversity plot at San Martin-3.
137
References
Andersen, A. N. 1990. The use of

ant communities to evaluate
change in Australian terrestrial
ecosystems: a review and a recipe.
Proceedings of the Ecological Society of
Australia 16: 347-357.
Basset, Y. 1991. The taxonomic
composition of the arthropod
fauna associated with an Australian rainforest tree. Australian
Journal of Zoology 39: 171-190.
Bestelmyer, B. T., and J. A. Wiens.
1996. The effects of land use on
the structure of ground-foraging
ant communities in the Argentine
Chaco. Ecological Applications 6(4):
1225-1240.
Bolton, B. 1996. A Catalogue of the
Ant Genera of the World. Harvard
University Press, Cambridge.
Delabie, J. H. C., and J. D. Majer.
1996. Comparison of the ant
communities of annually inundated and terre firme forests at
Trombetas in the Brazilian Amazon. Insectes Sociaux 41: 343-359.
Fittkau, E. J., and H. Klinge. 1973.
On biomass and trophic structure
of the Central Amazonian rain
forest ecosystem. Biotropica 5:
2-14.
Gaston, K .J. 1991. The magnitude
of global insect species richness.
Conservation Biology 5(3): 283-296.
Holldobler, B., and E.O. Wilson.
1990. The Ants. Harvard University Press, Cambridge.
Kremen, C., R. K. Colwell, T. L.
Erwin, D. D. Murphy, R. F. Noss,
and M. A. Sanjayan. 1993. Terrestrial arthropod assemblages: their
use in conservation planning.
Conservation Biology 7: 796-808.
138
Majer, J. D. 1983. Ants: bioindicators of minesite rehabilitation,

land-use, and land conservation.
Environmental Management 7:
375-383.
Wilson, E. O. 1987. The little
things that run the world (the
importance and conservation of
invertebrates). Conservation Biology
1: 344-346.
Wilson, E. O. 1988. The current
state of biological diversity. In
Biodiversity. (E. O. Wilson and F.
M. Peter, eds.) National Academy
Press, Washington, DC.
Appendix 1. Ant species sampled by intensive sampling of leaf litter and

soil nests in biodiversity plot #1 at San Martin-3, Lower Urubamba region.
Sampling occurred March 8-24, 1997. The number of quadrats in which
each species was found is reported.
Genus
Subfamily Ponerinae
Gnamptogenys
Gnamptogenys
Gnamptogenys
Gnamptogenys
Hypoponera
Hypoponera
Hypoponera
Odontomachus
Pachycondyla
Pachycondyla
Prionopelta
Subfamily Myrmicinae
Apterostigma
Apterostigma
Apterostigma
Basciceros
Basciceros
Crematogaster
Crematogaster
Crematogaster
Cyphomyrmex
Cyphomyrmex
Hylomyrma
Megalomyrmex
Pheidole
Pheidole
Pheidole
Pheidole
Pheidole
Pheidole
Pheidole
Pheidole
Pheidole
Pheidole
Pheidole
Pheidole
Pheidole
Pheidole
Pheidole
Pheidole
Pheidole
Species
Quadrat
1
2
3
4
1
2
3
1
1
2
1
1
1
2
5
4
2
3
4
1
2
5
1
2
3
1
2
1
2
3
1
2
1
1
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
1
3
1
1
1
6
1
1
1
1
1
4
1
10
4
1
1
1
1
1
2
2
1
2
1
2
2
1
1
139
Appendix 1. Ant species sampled by intensive sampling (Cont.).

Genus
Species
Solenopsis
Solenopsis
Solenopsis
Solenopsis
Solenopsis
Wasmannia
Subfamily Dolichoderinae
Dolichoderus
Subfamily Formicinae
Brachmyrmex
Paratrechina
Paratrechina
Paratrechina
Paratrechina
Paratrechina
Paratrechina
140
Quadrat
1
2
3
4
5
1
2
3
2
4
1
3
1
1
2
3
4
5
6
2
5
1
1
3
2
1
Bees and Wasps

(Hymenoptera:
Aculeata):
Biodiversity
Assessment
in the Lower
Urubamba
Region
Manuel Laime
Mayor de San Marcos
Introduction
The eastern slopes of the

Peruvian Amazon contain regions
with very high estimates of species
diversity for all living groups (Wilson and Sandoval 1996, Udvardy
and Sandoval 1996). Among them,
the upper and lower regions of the
Urubamba River offer an intricate
landscape with great species diversity (Lamas 1984, Udvardy and
Sandoval 1996). Invertebrates have
relatively well studied in the upper
part of the Urubamba River, including groups such as butterflies,
beetles, dragonflies, flies, and bees
(Lamas 1984). However, the invertebrate fauna of the Lower
Urubamba is practically unknown.
This paper presents the results of a

study of bees and wasps in the
Lower Urubamba as part of the
general biodiversity assessment that
the Smithsonian Institution is
conducting in the region.
Bees and wasps, as with many
other insects, can fly to find food
and mates and to disperse. They are
extemely important in ecosystem
functioning since they are the
primary pollinators of hundreds of
flowering plants. Many species of
wasps parasite other species of
insects, serving as population
regulators of potential pest species,
and several species of bees have
evolved social traits that makes
them very competitive. The honey
that bees produce is used by indigenous peoples in remote areas as
an important source of sugars and
proteins. The biodiversity assessment conducted during this study
identified which species of bees and
wasps are present in the study area,
information that can be used in
management decisions.
141
Methods
The study was conducted at the

site of the San Martin-3 (Geographic Positioning Systems coordinates 114709.8"S, and
724205.3"W), where the characteristic vegetation is primary forest
dominated by the bamboo Guadua
sarcocarpa subsp. sarcocarpa Londoo
& P.M. Peterson. Several standardized methods were used to sample
arthropods by my collegues at the
camp. I used their samples of
wasps and bees to determine the
biodiversity of the sites. The
protocols used to sample bees and
wasps are described in the following
paragraphs.
Pitfall Traps and
Winkler Sacks
The pitfall traps consisted of
plastic cups approximately 6 cm in
mouth diameter and 12 cm deep.
These traps were filled with about
15 milliliters of ethanol and placed
10 m apart in four 100-m transects;
two inside and two outside the
biodiversity plot #1 (Alonso et al.
this volume). The traps were left on
the field for 48 hours. At each
station, the leaf litter accumulated
in 1m 2 was sifted for extraction
using the Winkler method; that is,
the leaf litter is placed in a thin bag
for drying. As the sample dehydrates, the insects crawl out the bag
and fall into a plastic cup containing ethanol as a preservative.
Malaise Trap
This tent-like structure intercepts insects as they fly upward, as
most do when they are disturbed. A
cup containing ethanol as a preservative is placed in the highest
corner of the roof of the trap. Six
malaise traps were placed in the
biodiversity plots along the streams,
142
where they aided in sampling

diurnal and nocturnal flying insects.
Sweep Sampling
This method facilitated the
opportunistic sampling of bees and
wasps encountered along trails and
streams. The insects were netted
with a circular white silk bag, 60 cm
in diameter by 90 cm deep, set on a
pole 1 m in length. Because many
species of bees and wasps sting, the
use of a killing jar filled with
ethyl acetate is recommended.
Baited Pitfall Traps
These traps consist of a halfliter plastic cup that contains a
smaller cup inside and filled with
bait. I used rotten fruit and fish,
human feces, and pieces of a dead
snake for bait. The smaller cup
containing the bait was covered
with a fine mesh to reduce contact
with the specimens. At the bottom
of the larger cup, a preservative
was placed to reduce decomposition
of the captured specimens. Traps
were left out for 48 hours, but they
were checked every 12 hours to
distinguish between diurnal and
nocturnal faunal activity.
Light Traps
Most nocturnal insects are
attracted to sources of light. This
principle is used to attract insects
by placing a light with an ultraviolet
bulb on a white sheet, a source of
light that competes with the light of
the moon used by the insects for
orientation. The insects that come
to the light do not leave until then
light is turned off.
General Observations
I used 7 x 35 binoculars to
make direct observations of bee
and wasp nests.
Specimen identification was

done by using the appropriate
taxonomic keys and by direct
comparison with museum specimens in the Hymenoptera collection
of the Department of Entomology,
Museo de Historia Natural de la
Marcos, Lima, Per. Ordination to
the level of superfamilies and
families followed Goulet and Huber
(1993). For the Spheciformes, I
followed Bohart and Menke (1976).
For the Apiformes families, I took
as reference the proposed changes
published by Michener et al. (1994)
by Roig-Alsina and Michener.
Actual ordination of the genera
followed Van der Vecht and Carpenter (1990). Albert Finnamore, a
hymenoptera specialist, confirmed
identification of many species.
Results and
Discussion
I registered a total of 54 species of wasp and bees, most of

which have been determined to a
morphological species label. The
most diverse groups are within the
superfamilies Vespoidea and
Apoidea. These results, however,
are just a fraction of the diversity
of wasps and bees that live at San
Martin-3. More intensive studies
will help determine just how diverse
in wasps and bees the study area is.
Twenty of the wasp species and
four species of bees live on the low
stratum near the ground and the
leaf litter, while 24 wasp and six
bee species are of the medium and
high strata. These data show that
the fauna in the different strata is
comprised of different species. It is
also likely that more specific microhabitats are located within the three
strata, as is the case for other
groups of insects (e.g., Carabidae;

Erwin 1990).
Even though the sample of the
hymenoptera species was good, it
can be improved by using the
malaise traps more intensively and
by setting canopy traps where the
fauna seems to be different. The
most effective protocols to sample
hymenoptera included baited traps,
light traps, and general observations. In the future, ground and
canopy yellow pan traps should be
used because they are effective in
attracting bees and wasps. I also
propose to use trap nesting in the
future, a protocol that has not been
tested in Per. It consists of setting
out the types of materials that the
insects will use to make their nests.
Sampling by this method allows one
to gather valuable information
about a number of factors, including complete life cycles, changes
with seasonality, and population
dynamics. Learning about nest
designs can also help distinguish
species.
Here, I also report a new supra
generic taxa for the entomofauna of
Perthe Sclerogibbidae and
Scolebithydae. This finding enhances our understanding of the
biogeography of the area since the
Scolebythidae is also reported in
tropical Africa and Brasil
(Humphries and Parenti 1986,
Goulet and Huber 1993), and it
confirms how poorly known the
hymenoptera fauna is even in this
areaone of most biologically rich
places in the world (Quintero and
Cambra 1996). I also observed that
Trigona spp. was collecting minerals
from pipes at the well sites. Further
study of this observation may prove
helpful in determining whether it is
useful as an indicator of pollutant
levels. Genetic studies of these
The most
diverse
groups are
within the
superfamilies
Vespoidea
and
Apoidea.
We found a
new supra
generic
taxa for the
entomofauna of
Perthe
Sclerogibbidae
and Scolebithydae.
143
bees can give us an idea of the

effects that the chemicals may have
on this group.
Genetic
studies of
these
bees can
give us an
idea of
the
effects
that the
chemicals
may have
on this
group.
General Observations
about the Wasps and Bees
of San Martin-3
Family Vespideae. Polybia sp. 1.
I observed two individuals of this
species. One flew into the ceiling of
our tent and landed on hundreds of
dead ant queens that perished the
night before. It is probable that it
selected some of the fresher specimens as prey, caught them by their
mandibles, and flew away with
them. The other was observed
hunting a light green adult
Trichoptera similar to a moth.
These two specimens were sampled
for future identification.
Family Pompilidae. Priochilus
sp. 2. A female wasp was captured
while hunting a wolf spider
(Lycosidae) that was carrying an egg
mass of the Aculeata. It has been
suggested that individuals in this
genus make their nests on cavities
of plants or inside the bark of trees
and use leaves of plants to build
nests.
Family Apidae. Trigona sp. 1.
Several specimens were observed
during my one-day visit to the
Cashiriari-2 well site. They were
taking minerals from a tube used to
dispose materials. I was told that
these bees have seen near the the
drilling site.
Other Bees and Wasps
Expected to Live
at San Martin-3.
Several genera of wasps and
bees that can be expected in the
area are within Chrysidoidea. The
Chrysididae could have more genera
within the Amiseginae and the
Chrysidinae. It is also probable that
144
they are more species of Bethylidae

and Drynidae. Among the
Vespoidea, the Tiphiida should be
represented by Tiphia spp. and many
genera of Thynninae. The Scoliidae
should be represented by Scoliinae
and the Pompilidae by more species
of Pepsis spp., Pompilocalus, and
other related genera (Roig-Alsina
1989).
The Mutillidae should be better
represented in the number of genera
and species (Quintero 1996). The
Vespidae, with the Eumeninae, can
be represented by Monobia spp.,
Montezumia spp., and Zethus spp.
(Willink 1982). The Polistinae
probably have more species within
Polybia spp., Polistes spp., Agelaia
spp., Mischocyttarus spp., and Sinoeca
spp.
The Apoidea, in the Sphecidae
can be represented by Sceliphron
spp., Sphex spp., Prionyx spp.,
Ammophila spp., Eremmophila spp.,
Trigonopsis spp., Podium spp., and
Penepodium spp. The
Pemphredonidae should be represented by Stigmus spp., Microstigmus
spp., Llagmastigmus spp., Pseneo spp.,
and Psen spp. (A. Finnamore, personal communication).
The Crabronidae could be
represented by more species of Liris
spp., Larra spp., Lyroda spp.,
Trypoxylon spp., Pisoxylon spp., and
Pison spp., as well as Tachytes spp.
and Tachysphex spp. The Nyssonidae
should include species of
Gorytinae; the Bembecinae the
genera of Stictia spp., Rubrica spp.,
Microbembex spp., Bycirtes spp., and
Editha spp. and many Nyssoninae.
The Apidae, the genera within the
Euglossinae Exaerete spp., Eulaema
spp., and Eufriesa spp., are expected
in the area as well as other species
within the Meliponinae, Halictidae,
and Megachilidae.
References
Bohart, R. M., and A. S. Menke.

1976. Sphecid Wasps of the
World: A Generic Revision. University of California Press, Berkeley and Los Angeles and London.
Erwin, T. L. 1990. Natural history
of the Carabid beetles at the
BIOLAT Biological Station, Rio
Manu, Pakitza, Per. Revista
Goulet, H., and I. T. Huber. 1993.
Hymenoptera of the World: An
Identification Guide to Families.
Publication 1894/E. Research
Branch Agriculture, Canada.
Humphries, C. J., and R. Parenti.
1986. Cladistic Biogeography.
Clarendon Press, Oxford.
Lamas, G. 1984. La Entomofauna
de la Zona Reservada de
Tambopata, Madre de Dios, Per.
1st Part. Revista Peruana de
Michener, C. D., R. McGlingley, and
B. Danforth. 1994. The Bee
Genera of North and Central
America.(Hymenoptera: Apoidea).
Washington, DC.
Quintero, D., and R. A. Cambra.
1996. Contribucion a la
sistematica de las Mutilidas (Hymenoptera) del Per, en especial
las de la Estacion Biologica
BIOLAT, Rio Manu, Pakitza.
(E. Wilson and A. Sandoval, eds.)
Roig-Alsina, A. 1989. La posicion

sistematica de los grupos hasta
ahora incluidos en Chirodamus
Haliday sensu lato y revision del
genero Pompilocalus gen. nov.
(Hymenoptera, Pompilidae).
Revista Sociedad Entomologica de
Argentina 47: 3-73.
Vecht, J., Van der, and J. M. Carpenter. 1990. A catalogue of the
genera of the Vespidae (Hymenoptera). Zool. Verhandl. Leiden.
260: 1-62.
Proceedings from the Workshop
on Biological and Cultural Diversity of the Lower Urubamba River.
Washington, DC.
Manu: The Biodiversity of Southeastern Per. Ed. Horizonte,
Lima.
Willink, A. 1982. Revision de los
Generos Montezumia Saussure, y
Monobia Saussure (Hymenoptera:
Eumenidae). Boletin Academia
Nacional de Ciencias (Cordova,
Argentina) 55: 1-321.
145
Appendix 1. Classification of bees and wasps sampled at San Martin-3,

Lower Urubamba region, Per (5 superfamilies cited of the series
Parasitica: Proctotrupoidea, Cynipoidea, Ceraphronoidea, Chalcidoidea and
Ichneumonoidea; among the Aculeata, three superfamilies were registered:
Chrysidoidea, Vespoidea and Apoidea; the Chrysidoidea is represented by 6
species, the Vespoidea by 27, and the Apoidea 22; the parasite series and
the Formicidae [ants] are only mentioned as a reference).
SUPERFAMILY
Family
TENTHREDINOIDEA
Pergidae
Tenthredinidae
PROCTOTRUPOIDEA
Proctotrupidae
Diapriidae
Platigasteridae
Evaniidae
Cynipidae
Ceraphronidae
Encyrtidae
Agaonidae
Chrysididae
Subfamily Amiseginae
CYNIPOIDEA
CERAPHRONOIDEA
CHALCIDOIDEA
CHRYSIDOIDEA
Bethylidae
Subfamily Epyrinae
Sclerogibbidae
VESPOIDEA
Scolebythidae
Tiphiidae
Subfamily Methocinae
Mutillidae
Subfamily
Sphaerofthalminae
Pompilidae
Subfamily Pepsinae
Subfamily Pompilinae
Vespidae
Subfamily Polistinae
146
Species
Pergidae sp. 1
Tenthredinidae sp. 1
Tenthredinidae sp. 2
Amiseginae sp. 1
Amiseginae sp. 2
Epyrinae sp. 1
Sclerogibbidae sp. 1
Sclerogibbidae sp. 2
Sclerogibbidae sp. 3?
Scolebythidae sp. 1
Methocinae sp. 1
Sphaerofthalminae sp. 1
Sphaerofthalminae sp. 2
cf Auplopus
Pepsinae sp. 1
Pepsis sp. 1
cf Aporus
Anoplius sp. 1
Priochilus sp. 1
Priochilus sp. 2
Polybia
Polybia
Polybia
Polybia
Polybia
sp. 1
sp. 2
sp. 3
sp. 4
sp. 5
Appendix 1. Classification of bees and wasps sampled at San Martin-3 (Cont.).

SUPERFAMILY
APOIDEA
SPHECIFORMES
APIFORMES
Family
Formicidae
Species
Polybia sp. 6
Polybia sp. 7
Polybia jurinei Saussure 1853
Polistes sp. 1
Agelaia testacea Fabricius 1804
Brachygastra sp. 1
Apoica pallida Olivier 1791
Apoica pallens Fabricius 1805
Pemphredonidae
Subfamily Pemphredoninae Incastigmus sp. 1
Crabronidae
Subfamily Larrinae
Larra sp. 1
Trypoxylon sp. 1
Trypoxylon sp. 2
Trypoxylon sp. 3
Pisoxylon sp. 1
cf Nitela sp. 1
Subfamily Crabroninae
Enoplolindenius sp. 1
Enoplolindenius sp. 1
Quexua sp. 1
Halictidae
Apidae
Subfamily Anthophorinae
Subfamily Meliponinae
Halictidae sp. 1
Halictidae sp. 2
Halictidae sp. 3
Xylocopa sp. 1
Trigona sp. 1
Trigona sp. 2
Melipona sp. 1
Apidae sp. 1
Apidae sp. 2
Apidae sp. 3
Apidae sp. 4
Euglossa sp. 1
Euglossa sp. 2
Apis mellifera Linnaeus 1758
147
148
Dragonflies and
Damselflies
(Odonata:
Anisoptera and
Zygoptera):
Biodiversity
Assessment
in the Lower
Urubamba
Region
Jerry A. Louton
Introduction
Dragonflies and damselflies

(Odonata: Anisoptera and
Zygoptera) are excellent subjects
for biodiversity studies. They are
relatively well known compared to
other groups of tropical insects,
they are easily sampled and studied,
and they are important in benthic
communities that are routinely used
to assess the quality of surface
water. Adults are large, colorful,
day-flying insects that can often be
identified to genus on the wing.
Larvae are aquatic and often en-
countered in benthic studies. Both

adults and larvae are large predators
near the tops of their respective
food chains. Sampling is relatively
non-disruptive because local populations of these insects generally
have high numbers of individuals,
and only a small proportion is
taken. In most of the tropical areas
of the American Continent (i.e.,
neotropical areas), much work
remains to be done in describing
new species, defining genealogical
relationships among species (i.e.,
producing phylogenies), determining species distributions, and understanding conservation needs
(Louton et al. 1996a, b).
The immediate objectives of
this study were to determine the
local biodiversity of the dragonflies
and damselflies (Odonata) during
the wet season at the Lower
Urubamba River, to compare these
baseline data with other neotropical
sites (e.g., Pakitza and Gueppi), to
develop further the Identification
Guide to Genera of New World
Odonata, and to test rapid sampling protocols for neotropical
Odonata. Subsequent studies
149
should include semi-quantitative

sampling of general aquatic insect
communities in streams, ponds, and
other aquatic habitats.
Methods
The area around the study site

is characterized as terra firme forest
with terraces and hills. The immediate area is pacal, a forest dominated by the spiny bamboos Guadua
sarcocarpa (see Acevedo et al. this
volume). The presence of the
bamboo species afforded excellent
study habitat for damselflies since
their immature stages are aquatic.
Various invertebrate and vertebrate
species make lateral perforations in
the bamboo stems. These become
filled with water pumped from the
roots and are used by several species of aquatic organisms to complete their life cycle.
I concentrated on aerial netting
of adult specimens. I used a typical
butterfly net to intercept dragonflies
and damselflies as they flew, and I
used the available trails as transects
through the bamboo-dominated
forests. I also included stream
crossings and the camp clearing (see
Appendix 1). Because the base
camp was located on a ridge top,
only a few small- and one
moderate-sized streams were available. Water that collected in nearby
depressions after hard rains drained
away in only a couple of hours.
During unfavorable weather (i.e.,
constant rain) for adult sampling, I
switched to sampling of aquatic
immature stages of bamboodwelling damselflies and their prey
community.
150
Results
The bamboo internodes were

sampled briefly by cutting internodes and examining by hand the
contents in enameled pans. It was
confirmed that possibly two species
of helicopter damselflies occupy
the top predator niche in the system
and that a diverse fauna of mosquitoes, other flies, and coleoptera are
present. The amount of water
available in the bamboo internodes
is estimated to be to be about
10,000 l/ha ([1 culm/m x 5 l/culm
x 10,000] / 0.2 [rate of stem perforation]).
Because of inclement weather
(more than seven hours of rainfall
daily), limited (three hours per day)
sunlight (Appendix 2), lack of
adequate sampling transects, and
inability to visit the site at
Cashiriari-2, we sampled only about
150 adult specimens. Habitat types
available included first- and
second-order streams and
phytotlemata, mostly water-filled
bamboo internodes (bromeliads
were present but not abundant).
Common Odonata habitats not
available at San Martin-3 included
larger streams, rivers, oxbow ponds,
swamps, and marshes. A
cross-tabulation of the raw data
yielded a listing of about 20
morphospecies (mostly genera) by
nine habitat types (Appendix 1). We
estimate that this will represent
about 40 species when preparation
(genetalia dissection) is complete.
Conclusions
In spite of the difficult circumstances and small sample, some

basic comparisons with the nearby
Smithsonian study site at Pakitza,
Rio Manu, Per, can be made if the
data are classified by species or
habitat. For example, the helicopter

damselfly fauna was identical with
Pakitza except for the absence at
San Martin-3 of one species,
Mecistogaster linearis, whose larvae
are known to inhabit water-filled
holes in large trees. This species
probably occurs locally, but is
uncommon because of the lack of
large mature trees with water-filled
holes. This genus is exceptional;
most other obligate under-canopy
genera were less frequently represented at San Martin-3 by a ratio of
about 1:2. Excluding phytotelmbreeders, the richness of forest
species at San Martin-3 may be
about one-half that of Pakitza.
Exact comparisons must await
exact identification of the taxa.
Presumably, if the study-area at San
Martin-3 were expanded to include
an area comparable in size and
floral diversity to Pakitza, the fauna
would be as diverse. This could
easily be accomplished via
trail-transects from San Martin-3 to
the Urubamba River.
A visit during the dry season
and expansion of the study area are
necessary to a high-quality evaluation of the Odonata fauna at San
Martin-3. The ability to ship ahead
is necessary for movement of
materials that are difficult to obtain
locally and difficult to carry as
luggage on domestic airlines. Finally, the mosquito fauna appears
very diverse in the water-filled
bamboo internodes and should be
given priority status for study.
References Cited
Louton, J. A., R. W. Garrison, and

O. S. Flint. 1996. The Odonata of
Parque Nacional Manu, Madre de
Dios, Per; natural history, species
richness and comparisons with
other Peruvian sites. Pages
431-449 in Manu: Biodiversity of
Southeastern Per (La
Biodiversidad del Sureste del
Per). (D. E. Wilson and A.
Sandoval, eds.) Editorial
Horizonte, Lima.
Louton, J., J. Gelhaus, and R.
Bouchard. 1996. The aquatic
macrofauna of water-filled bamboo (Poaceae: Bambusoideae:
Guadua) internodes in a Peruvian
lowland tropical forest. Biotropica
28: 228-242.
151
Appendix 1. The dragonflies and damselflies of the Lower

Urubamba region (species of dragonflies and damselflies sampled at
San Martin-3 in February and March, 1997, indicating the number
and the site where the specimens were found: A= camp clearing; B=
large stream; C= trail to large stream; D= stream at biodiversity plot
1; E= trail to biodiversity plot 1; F= stream at biodiversity plot 2;
G= trail to biodiversity plot 2; H= stream to waterfall; I= trail to
waterfall; J= grand total).
Order Odonata
Suborder Zygoptera
Family Polythoridae
Polythore manua?
Family Calopterygiidae
Hetaerina sp 1.
Mneserete? sp. 1
Family Perilestidae
Perissolestes
Family Megapodagrionidae
Heteragrion sp. 1
Family Pseudostigmatidae
Mecistogaster spp. (mostly jocaste)
Mecistogaster sp. nov.
Microstigma sp. 1
Family Protoneuridae
Protoneurid
Family Coenagrionidae
Argia black w. blue
Argia black w. olive
Argia black w. purple
Argia cuprea?
Argia females
Suborden Anisoptera
Family Aeshinidae
Gynacantha membranalis
Family Libellulidae
Libellulid (hairy torax)
Macrothemis? sp. 1
Orthemis sp. 1
Pantala flavescens
Grand Total
152
B C D E F G H I
0 0
0 1
0
0
4
2
0
0
0
2
0
0
2 0
16 0
0
0
0 6
0 20
5 0
0 5
20 0
0 25
1
0
0
0
0
0
16 0
4 0
0 0
2
1
0
0 0
0 0
0 1
0
0
0
3 22
0 5
1 2
11 0
0 0
0 11
0
0
0
0
0
8
1
13
6
4
1
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
4
0
0
0
0
0
0
0
1
0
0
0
1
0
0
0
0
0
0 0
0 7
0
0
8
2
18
2
0
0
0
52
0
0
0
0
22
0
1
0
0
3
0
0
0
0
4
0
0
0
0
47
0
0
0
0
6
0
0
0
0
4
0
0
0
0
1
10
1
17
6
6
2
1
8
2
157
Appendix 2. Temperature and hours of sun and rain recorded at San

Martin-3 during study period (maximum and minimum temperatures
in degrees celsius and the number of sun and rain hours; time of rain
= morning (AM), noon (N), and afternoon-night (PM); high temperature for February 16 and 18 not included in the the calculation of the
average because the thermomether was inside the tent).
Date
16-Feb-97
17-Feb-97
18-Feb-97
19-Feb-97
20-Feb-97
21-Feb-97
22-Feb-97
23-Feb-97
24-Feb-97
25-Feb-97
26-Feb-97
27-Feb-97
28-Feb-97
1-Mar-97
2-Mar-97
Average
Max temp Min temp Sun (hrs.) Rain (hrs.) Time of rain
27.8
36.7
38.3
30.6
28.3
20.6
22.2
21.1
21.7
20
29.4
26.7
28.3
34.4
34.4
27.2
32.2
27.8
31.1
30.1
20
20.6
23.3
20.6
22.2
20.6
20
20.6
20.6
21.0
0
4
3
1
0
0
3
0.5
4
8
8
0
4
0.2
3
2.6
12
10
9
6
4
14
0
6
8
0
4
10
11
13
0
7.1
AM, PM
N, AM
N, AM
PM
N
N, AM
PM
N
N
N, AM, PM
N, AM, PM
N, AM, PM
153
154
Spiders
(Arthropoda:
Arachnida):
Biodiversity
Assessment
in the Lower
Urubamba
Region
Saida Cordova and
Janine Duarez
Mayor de San Marcos
Introduction
Spiders are found worldwide in

all terrestrial environments. Members of this invertebrate group have
a great ability to find prey and to
disperse. Individuals are diurnal or
nocturnal predators, mostly of other
species of invertebrates. They have
developed a variety of ways to
capture their food. Some spiders are
orb-weavers, others are sheet-web
weavers, others are cursorial hunters, and still other are expert at
ambush (Silva and Coddington
1996); all play an important role in
controlling the populations of their
prey (Riechert 1974). Spiders are
particularly numerous in the tropics,
where the abundant, species-rich
vegetation supports a great

amount of their prey (Foelix
1982). Habitat complexity and
changes in temperature and
precipitation (i.e., seasonality)
have strong influences in the
distribution and abundance of
spider communities in ecosystems (Lubin 1978, Coddington
et al. 1991). Land-use practices
also affect the composition of
the species of spiders (Coyle
1981).
Spiders are very diverse, as
gathered from the fact that
34,000 spider species have been
reported around the world. At
least one-half of them are found
in neotropical America. By 1988,
some 1120 genera of spiders had
been recognized in neotropical
areas, and at least one-half of
those genera are found in Per
(Silva 1996), represented in 76
families (Coddington and Levi
1991). In this study, we conducted an assessment of spider
diversity in the Lower Urubamba
region. No spider studies had
occurred in this region before our
efforts.
155
Methods
For most
species of
spiders,
both sexes
are needed
for appropriate
classification to the
species
level.
To capture as many spiders as

possible at the study sites (San
Martin-3 and Cashiriari-2 well
sites), several sampling methods
described belowwere used. These
protocols were also helpful in
sampling other terrestrial invertebrates. Letters in parentheses
indicate the code used to label the
sample.
General Sollecting (G)
We sampled the spiders encountered as we walked the trails,
the drilling platform, and the camp.
During nocturnal expeditions, we
sampled spider that are active at
night, a very different assemblage
from the daytime spiders.
Winkler sack (W)
Ground leaf litter is sifted to
eliminate large particles. The
smaller material is placed in a bag
that is hung inside a larger bag
where the litter dries. Because all
invertebrates dislike desiccation,
the animals move in search of
better conditions as the litter dries.
As the invertebrates leave the bag,
they fall into the second bag, which
contains a flask at the bottom with
a preservative. Specimens are then
separated in different taxa groups
for classification. Solitary spiders
are commonly trapped with this
method.
Beating (B)
This protocol samples spiders
that live on vegetation. A framed,
white sheet is placed underneath a
branch, which is hit with a stake.
All living arthropods fall into the
sheet and are captured. The white
sheet is helpful in sampling very
small spiders, among other invertebrates.
156
All specimens were sorted to

morphospecies; i.e.,, individuals
with the same morphological characteristics were considered to
belong to the same species. Many of
the specimens could not be identified to species since the taxonomic
revisions are scarce, and many
neotropical species have not been
described. To make our task even
more challenging, many sampled
specimens were immature, or we
sampled only one sex. For most
species, both sexes are needed for
appropriate classification to the
species level. Thus, most of the
adult specimens were classified up
to the genus label.
Results and
Discussion
A total of 69 morphospecies
were identified from our samples.
Eighteen families and 59
morphospecies were found within
the Order Araneae. One family of
the infraorder Mygalomorpha was
represented by two species. Within
the Order Opiliones, six
morphospecies were found of two
infraorders. One morphospecies was
found on each of the Scorpions,
Pseudoscorpion, and Amblypigi
orders (Appendix 1).
Within the Order Aranea, the
Family Araneidae was represented
by 13 morphospecies. This family is
well represented in tropical areas;
its species can be found in a wide
variety of microhabitats. We
sampled seven adult individuals of
four morphospecies of Parawixia
spp., all sampled by beating the
vegetation. Parawixia spp. was
represented by nine described
species for Per (Levi 1992). Eight
morphospecies were recognized
within the Family Salticidae.
Twenty individuals, mostly juveniles, were sampled by beating the

vegetation. Three species were
found of the Family Lycosidae.
These spiders are active during the
day and can be found crawling on
the ground. We found 14 individuals, three of themtwo females
and one malebelonging to the
genus Pardosa spp. Members of this
family are nocturnal and can be
found in all forest strata, from the
ground (i.e., leaf litter) to the
canopy or nearby sources of water.
We found 34 individuals and recognized seven species, most of them
of them of the genus Ctenus spp.
Surprisingly, we did not capture any
members of the genus Phoneutria
spp., a common genus of medical
importance.
We found three species of the
Family Theridiidae, a worldwide
family with more than 2000 species.
These spiders can be solitary,
colonial, or kleptoparasites of other
spiders. Kleptoparasites are spiders
that live on webs made by other
spiders, allowing them to take the
food captured by the other spiders.
Members of this family can be
found from the ground to the
canopy. We found only five individualsone adult female Dipoena
spp., two adult females that have
not yet been identified, and two
juveniles. Dipoena spp. is found
throughout the tro-pics, and D. alta
is a species reported for Per (Levi
1963), but different than the one
sampled by us. Members of the
Family Corinnidae mimicboth
morphologically and behaviorally
ant species. They can be found
searching for prey on the ground or
at the forest canopy. Most of the
species are not common. Only one
adult male was sampled of the
genus Corinna spp. Spiders of the
family Deinopidae produce highly

specialized webs; in many cases the
web is smaller than the spider
(Coddington and Sobrevila 1987).
Two adult females were sampled of
the genus Deinopis spp.
References Cited
Coddington, J. A., C. E. Griswold,

D. Silva, E. Penaranda, and S. F.
Larcher. 1991. Designing and
testing sampling protocols to
estimate biodiversity in tropical
ecosystems. Pages 44-60 in The
Unity of Evolutionary: Proceedings of the Fourth International
Congress of Systematics and
Evolutionary Biology. (E. C.
Dudley, ed.) Dioscorides Press,
Portland, Oregon.
Coyle, F. A. 1981. Effects of clear
cutting on the spider community
of a Southern Appalachian forest.
Journal of Arachnology 9: 285-298.
Levi, H.W. 1963. American spiders
of the genera Audifa, Euryopis, and
Dipoena (Areneae: Theridiiae).
Bulletin of the Museum of Comparative Zoology 129: 121-185.
Levi, H. W. 1992. Spiders of the
orb-weaver genus Parawixia in
America (Araneae: Araneidae).
Bulletin of the Museum of Comparative Zoology 153: 1-46.
Lubin, Y. D. 1978. Seasonal abundance and diversity of
web-building spiders in relation to
habitat structure on Barro Colorado Island, Panama. Journal of
Arachnology 6: 31-51.
Riechert, S. E. 1974. Thoughts on
the ecological significance of
spiders. Bioscience 24: 352-356.
Silva, D. 1992. Observations on the
diversity and distribution of the
spiders of Peruvian montane
forests. Memorias del museo de
Historia Natural, UNMSM (Lima)
21: 31-37.
157
Silva, D. 1996. Determinacion de

areas importantes de diversidad
biologica en el Per: Situacion de
las aranas (Arthropoda: Araneae).
Pages 93-94 in Diversidad
Biologica del Per: Zonas
Prioritarias para su Conservacion.
(L. Rodriguez, ed.) Proyecto
FANPE GTZ-INRENA, Lima.
158
Silva, D., and J. A. Coddington.

1996. Spiders of Pakitza (Madre
de Dios, Per): Species richness
and notes on community structure.
Appendix 1. Spiders of the Lower Urubamba region (morphospecies recognized for each family are listed in the column).
Orden Araneae
Family Araneidae
Family Salticidae
Family Lycosidae
Family Ctenidae
Family Theridiidae
Family Corinnidae
Family Deinopidae
Family Scytodidae
Family Pisauridae
Family Oonopidae
Family Linyphidae
Family Anyphaenidae
Family Clubionidae
Family Heteropodidae
Family Pholcidae
Family Selenopidae
Family Tetragnathidae
Family Thomisidae
Genera
Morphospecies
Acacesia cf. benigna Simon 1895

Cyrtophora Simon 1864
Micrathena Sundevall 1833
Parawixia F. O. P.- Cambridge 1904
Pardosa C. L. Koch 1847
Ctenus Walckenaer 1805
Dipoena Thorell 1869
Corinna C. L. Koch 1841
Deinopis MacLeay 1839
Scytodes Latreille 1804
Ancylometes Bertkau 1880
INFRAORDEN
MYGALOMORPHA
Family Theraphosidae
Avicularia Lamarck 1818
Pamphobeteus Pocock 1901
ORDEN OPILIONES
SUBORDEN LANIATORES
Family Cranaidae
Family Gonyleptidae
SUBORDEN PALPATORES
13
4
8
3
7
7
3
1
1
1
2
1
2
2
2
1
1
1
1
2
2
2
1
1
1
ORDEN SCORPIONES
Family Scorpionidae
ORDEN PSEUDOSCORPIONES
ORDEN AMBLYPYGI
159
160
Snails (Mollusca:
Gasteropoda):
Biodiversity
Assessment
in the Lower
Urubamba
Region
Rina Ramrez and
Saida Crdova
Mayor de San Marcos
Introduction
Terrestrial mollusks include

snails, sloths, and pseudo-sloths
(veronicellidos). The estimated
number of these species in the
world is between 30,000 and
35,000, or more than the total
combined number of species of
mammals, birds, reptiles, and
amphibians (Solem 1984). The
largest portion of mollusk
biodiversity is found in the tropics.
However, large regions of the
tropics remain unexplored, and its
diversity is only predicted based on
a few better known sites (Solem
1984).
The biodiversity assessment
and monitoring program of the
Smithsonian Institution, Monitoring

Program (SI/MAB) for the Lower
Urubamba region provides an
excellent opportunity to complement the information known about
this group of invertebrates. In
addition, learning more about
mollusks should help increase
understanding of ecosystem functions. Mollusks are very dependent
on the type of substrate and vegetation structure on which they live.
Mollusks are also important as
biogeographic indicators of early
tectonic events, as keys to knowledge about morphological evolution, as digestors of large amounts
of cellulose, as calcium accumulators, as a source of food for many
animals, including humans, and as
producers of valuable pharmacological products (Emberton 1995a).
The objective of this investigation is to provide an assessment of
the diversity of mollusks at the San
Martin-3 well site.
Methods
The assessment of terrestrial

mollusks was conducted during the
first 27 days of April, 1997. An
161
35
# of Species
30
25
20
15
10
Alive
5
All
0
0
10
12
14
16
18
20
22
24
26
Time (Days in April)
Figure 1. Species accumulation curve for sampled terrestrial

mollusks at San Martin-3. The All curve represents terrestrial
snails that were found alive and those found only by shell. This
curve is higher because four species were never found alive. After
day 15 of sampling, all of the species had been found, but the
curve for the species found alive reached its maximum at day 24.
intensive survey was conducted in
the two biodiversity monitoring
plots at San Martin-3. Random
sampling was also conducted in the
trails and creeks near the
biodiversity plots. The authors, with
the assistance of Antonio Gomez,
worked intensively on direct observation and with 1 m2 soil and litter
samples to sample carefully all
micro-mollusks. Twenty nine
samples of litter and soil were
analyzed, 13 of which were taken
from the biodiversity monitoring
plots. Ten were taken from the same
quadrats at the two plots. Two
samples were taken from the quadrats 03, 08, 13, 18, and 23. The
samples were collected on April 1,
3, 15, and 18.
The litter and soil samples were
taken separately, the soil from
approximately 2 cm deep in the
ground. The litter was surveyed
under a stereoscope microscope to
162
sample adequately all the different

species of micro-mollusks. The soil
sample was screened through
different size mesh sifters and
checked under the microscope. The
analysis of the litter and soil
samples was done by two individuals with nearly 14 hours of work.
We created a species accumulation
curve and used the time of collection when the sample was taken,
not the time when it was processed
(Fig. 1).
The snails and veronicellids
sampled were fixed in 10% formalin
and preserved in a 70% alcohol
solution. The appropriate bibliographic references and the mollusk
reference collection of the Museo
de Historia Natural de la
Marcos provided great assistance in
identification. The classification of
species is from Vaugh (1989) and

for the family Systrophiidae from
Ramirez (1993).
Results and
Discussion
The terrestrial mollusks from

San Martin-3 were classified in 34
species33 snails (mollusks with
shell) and one veronicellid
(pseudobabosa with no shell; Table
1). For four species (Drymaeus cf.
catenae, Guestieria sp., Labyrinthus
diminutus, Subulina sp.), we found
only the shells. The remaining
species were found alive. Twelve
species were represented by less
than five live individuals; for others
such as Plekocheilus cf. floccosa, we
did not find adult individuals. Most
of the species were in reproductive
state as indicated by the presence
of eggs.
Snails can be classified by the

diameter of their shell (Table 1).
Following Emberton (1995b), we
found that 69.7% (15 to 18 species)
of our sample had minute or small
shells, 15.2% (five species) had
medium size shells, 12.1% (four
species) had large shells, and the
remaining 3% (one species, the
edible "congompe" Megalobulimus
popelairianus [Ramirez and Caceres
1991]) had giant shells.
The species accumulation
curve shows interesting patterns
(Fig. 1). It rises quickly during the
first few days of sampling and then
flattens as the samples increase.
The curve with all the species is
higher because four species were
never found alive. We found 22
species the first day through direct
sampling in the forest; the other
species were found as the leaf litter
and soil samples were processed.
2.5
The
terrestrial
mollusks
from San
Martin-3
were
classified
in 34
species
33 snails
(mollusks
with shell)
and one
veronicellid
(pseudobabosa)
with no
shell.
Abundance (Log
10 )
1.5
0.5
11
13
15
17
19
21
23
25
27
29
Species Rank
Figure 2. Species rank based on abundance of species of terrestrial mollusks at San Martin-3. The variation in species abundance is very high,
from species that are very abundant to four species that were found only
once. More than half (19) of the species were represented by fewer than
10 individuals.
163
Most of
the species
of
mollusks
sampled at
San
Martin-3
represent
new
species to
science.
164
After day 15 of sampling, all of the

species had been found, but the
curve for the species found alive
reached its maximum at day 24. The
most abundant species was the
micro-snail Zilchogyra aff. microhelix,
found in all our samples. The
variation in species abundance was
very high (Fig. 2), from very abundant species with 207 individuals to
four species that were found only
once. More than half (19) of the
species were represented by fewer
than 10 individuals.
We found 32 species in the
biodiversity plot #1. The only two
missing species were the micro-snail
Miradiscops sp. 1 and the mollusk
with no shell of the Family
Veronicellidae. We registered 29
species in the biodiversity plot #2.
The species missing were Drymaeus
sp.1, Obeliscus (Protobeliscus) sp.1,
Euglandina cf. striata, and Guestieria
sp. 1, and the veronicellid. Thus,
the diversity of terrestrial snails is
very high per ha sampled. More
than 70% of the species registered
at San Martin-3 live on the leaf
litter and the humus. We found only
three species of arboreal snails of
the genus Drymaeus (8.8%). The
other seven species live on the
ground but are frequently found in
the understory vegetation. These
species include Helicina sp. 1,
Wayampia sp. 1, and Bulimulus sp. 1.
species of mollusks at San
Martin-3, where the vegetation is
dominated by bamboo (Guadua
sarcocarpa, see Alonso et al. this
volume). Intensive sampling in a
lowland tropical rain forest with
more habitat diversity in the Department of Madre de Dios (Cusco
Amazonico) resulted in a similar
number of species, but with very
different composition (Ramirez
1991). San Martin-3 is characterized

by a large diversity of very small
snails, while 42.4% of the mollusk
fauna was represented by medium
and large species in Madre de Dios.
The mollusks species of Madagascar are similar in size to those at
San Martin-3; Madagascar is the site
of one of the highest recordings of
diversity for terrestrial snails52
species in the world (Emberton,
1995b).
Most of the species sampled at
San Martin-3 represent new species
to science. This is not surprising
since most tropical snail species
have not been described (Emberton
1995a, 1995b; Ramirez 1991,
1993). The high number of sympatric species (species that live in the
same area) of the Family
Systrophiidae is especially interesting. This family is represented by 10
species, the most reported to date.
Members of this family, distributed
primarily in South America, are
predators of other snails (Ramirez,
1996). It is also interesting to note
the high abundance of Zilchistrophia,
one of the five most abundant
species. This genus of the Family
Systrophiidae is endemic to Per
and has been sampled very rarely,
including in the forest at
Chanchamayo (Department of
Junin), Divisoria (Department of
Huanuco), and Contamana (Department of Loreto).
The biodiversity assessment of
terrestrial snails is nearly complete
at San Martin-3. It was also possible
to obtain natural history information from several species, and
additional research should improve
the state of knowledge of this
highly diverse tropical area.
References
Emberton, K. 1995a. On the

endangered biodiversity of
Madagascan land snails. Pages
69-89 in Biodiversity and Conservation of the Mollusca. (Bruggen,
Wells, and Kemperman, eds.)
Backhuys Publishers, Leiden, The
Netherlands.
Emberton, K. 1995b. Land-snail
community morphologies of the
highest-diversity sites of Madagascar, North America, and New
Zealand, with recommended
alternatives to height-diameter
plots. Malacologia 36(1-2): 43-66.
Ramirez, R. 1991. Molluscs of
Cusco Amazonico. Pages 59-61 in
Report on Biodiversity at Cusco
Amazonico, Per. (Duellman,
compiler) Center for Neotropical
Biological Division, Museum of
Natural History, University of
Kansas, Lawrence, Kansas.
Ramirez, R. 1993. A generic analysis of the family Systrophiidae
(Mollusca: Gastropoda): Taxonomy, phylogeny and biogeography. Masters thesis. University of
Kansas, Lawrence, Kansas.
Ramirez, R. 1996. Diversidad de

moluscos terrestres en el Per.
Pages 95-97 in Diversidad
biologica del Per. Zonas
prioritarias para su conservacion.
(L. Rodriguez, ed.) Proyecto de
Cooperacion Tecnica, Ayuda en la
Planificacion de una Estrategia
para el Sistema Nacional de Areas
Naturales Protegidas
GTZ-INRENA, Lima.
Ramirez, R., and S. Caceres. 1991.
Caracoles terrestres (Mollusca,
Gastropoda) comestibles en el
Per. Boletin de Lima (77): 67-74.
Solem, A. 1984. A world model for
land snail diversity and abundance.
Pages 6-22 in World-Wide Snails:
Biogeographical Studies on
Non-marine Mollusca. (Solem and
Van Bruggen, eds.) Brill &
Backhuy, Leiden, The Netherlands.
Vaugh. 1989. A Classification of
the Living Mollusca. American
Malacologist Inc., Florida.
165
Appendix 1. Species of terrestrial mollusks recorded at San Martin-3 April

10-27, 1997 (shell size refers to the shell diameter, intervals after Emberton
(1995b): 0.50-5.00 mm (minute), 5.01-10.00 mm (small), 10.01-20.00 mm
(medium), 20.01-40.00 mm (large), and 40.01+ mm (giant); habitat: (1)
ground, (2) trees, (3) ground + vegetation away from the ground; rank:
species ranked by abundance [see Fig.2]; asterisk = recorded only by shell).
Mollusks
SUBCLASS PROSOBRANCHIA
Family Helicinidae
Helicina cf.
Helicina sp. 2
Helicina aff. bourguignatiana
Family Megalomastomidae
Aperosotma (Aperostoma) peruvianum
SUBCLASS GYMNOMORPHA
Family Veronicellidae
Species 1
SUBCLASS PULMONATA
Family Bulimulidae
Bulimulus sp. 1
Drymaeus sp. 1
Drymaeus rustrigatus
Drymaeus cf. catenae
Plecocheilus cf. floccosa
Family Subulinidae
Beckianum beckianum
Lamellaxis (Lamellaxis) sp. 1
Lamellaxis (Leptopeas) sp. 1
Leptinaria lamellata
Obeliscus (Ischnocion) aff. triptyx
Obeliscus (Protobeliscus) sp. 1
Opeas pumilum
Subulina sp. 1
Family Spiraxidae
Euglandina cf. striata
Family Megalobulimidae
Megalobulimus popelairianus
Family Systrophiidae
Drepanostomella aff. ammonoceras
Drepanostomella aff. excisa
Drepanostomella sp. 1
Guestieria sp. 1
Happia sp. 1
Miradiscops sp. 1
Systrophia aff. eatoni
Tamayoa sp. 1
Wayampia sp. 1
166
Habitat
Rank
small
minute
small
3
3
3
7
26
5
large
30
small
small
medium
medium
large
3
2
2
2
1
10
20
14
*
27
minute
minute
minute
small
minute
minute
minute
minute
1
1
1
1
1
1
1
1
11
18
29
15
28
24
25
*
medium
23
giant
12
minute
minute
small
minute
medium
minute
medium
minute
small
1
1
1
1
1
1
1
1
3
16
22
8
*
6
21
13
9
2
Shell Size
Appendix 1. Species of terrestrial mollusks recorded at San Martin-3 (Cont.).

Mollusks
Zilchistrophia sp. 1
Family Helicodiscidae
Zilchogyra aff. microhelix
Family Euconulidae
Habroconus cf. cassiquiensis
Family Camaenidae
Labyrinthus diminutus
Family Solaropsidae
Psadara aff. monile
Shell Size
Habitat
Rank
small
minute
minute
19
large
large
17
167
168
Beetles
(Coleoptera:
Scarabaeidae):
Biodiversity
Assessment
in the Lower
Urubamba
Region
Gorky Valencia
Abad del Cusco
Alfonso Alonso
Program (SI/MAB)
Introduction
Tropical forests are very complex in structure and function. They

have been rapidly transformed in
the last decades, and forest exploitation and reforestation with exotic
species have changed entire ecosystems. To help understand these
changes, one can study the roles
that animals play in ecosystem
stability and functioning. The
beetles are an invertebrate group
that has great potential for
biodiversity monitoring because of
the multiple roles they play in

ecosystems and because they are
influenced by environmental
changes (Lobo and Moron 1993).
Among the beetles are the
Scarabaeidae, small animals with
wide, strong bodies about 1.5 cm in
length; some can be up to 5 cm in
length, e.g. Megaphanaeus ensifer,
while others can be just a few
millimeters long. Most Scarabaeidae
are dark in color, but others can
have bright metallic colors such as
species of the genus Canthon and
the green metallic Oxysternon
conspicillatum (Fig. 1). Their morphology is well adapted for excavation; that is, their legs are very
strong to help them move large
quantities of soil and food. They
feed mostly on animal dung and
decomposing animals and plants.
These organisms are very important
to nutrient recycling, leading to a
great deal of interest in studying
them (Howden and Nealis1975).
Their ability to recycle animal dung
has been used to improve the
cultivation of grasses (Bornemissza
1960, Woodruff 1972 in Howden
169
Beetles
are an
important
component
of the diet
of many
insectivores.
Figure 1. This is a male of
Oxysternon conspicillatum Weber
1801, the most conspicuous beetle
species of the Scarabaeidae of the
Lower Urubamba Region; scale = 1
cm (drawing by G. Valencia).
and Nealis 1975) and has been
connected to disease transmission
because they move eggs of parasites that affect human health to the
soil surface, thus enhancing parasite
transmission to hosts (Miller 1954
in Howden and Nealis 1975).
These beetles are an important
component of the diet of many
insectivores. For example, birds
consumed up to 85.4% of these
animals in the terra firme in central
Amazonia (Bierregard 1990) and
49% in Manu, Madre de Dios, Per
(Karr et al. 1990). Fluctuations in
beetle populations have been
correlated to behavioral changes in
several species of mammals (Janson
and Emmons 1990, Audaga and
Halffter 1991). The Scarabaeidae
are important food items for
quiroptera, strigiforms, procionids,
170
lemerids, and amphibians (Moron

1981).
According to Halffter (1991),
the Scarabaeidae is a monophyletic
group (i.e. all members of this
group have a common ancestor),
with approximately 6,000 species in
200 genera. Since their principal
source of food is animal dung,
particularly from herbivores, most
of their behavioral, ecological, and
morphological characteristics are
associated with specialized food
gathering.
We studied the Scarabaeidae of
the Lower Urubamba region as part
of the biodiversity assessment that
the SI/MAB is conducting. We also
tested some methodologies to trap
these beetles.
Methods
We sampled the fauna of

Scarabaeidae in the biodiversity
plots at San Martin-3 and
Cashiriari-2. The areas that we
sampled are described as follows:
the bamboo (Guadua sarcocarpa)dominated biodiversity plot #1 at
San Martin-3, characterized by a
hilly terrain crossed by many
streams; the flatter biodiversity plot
#2 at SanMartin-3 , which contain a
more dense understory that plot #1;
and biodiversity plot #3 at
Cashiriari-2, where the vegetation is
dominated by mature tropical forest
with many emergent trees (see
Alonso et al. this volume).
Our sampling followed the
methodology suggested by M. A.
Moron (G. Valencia personal communication). The first author
modified the baited traps NTP-80
from Moron (1984) that were used
in the study (Fig. 2). This trap
attracts individuals with the odor
released by the bait. Attraction
success depends on wind direction,

temperature, humidity, distance,
and the position (i.e., ground or tree
canopy) of the bait. The
Scarabaeidae crawl or fly towards
the bait by using their antenna.
Traps are made of 1000-ml white
plastic containers with hermetic
tops. We cut three 2 x 3-cm openings 5 mm from the top of each trap
and hung a smaller (225 ml) cup
inside the container with a 14-cm
long wire about 5 mm from the top.
We placed the bait inside the
smaller cup and covered it with a
sealed top (in biodiversity plot #1)
or mesh screen (biodiversity plots
#2 and #3) to prevent the animals
from contacting the bait. We made
holes in the cup to facilitate escape
of the bait odor and placed 100 ml
of 30% alcohol in the larger container. Insects drawn to the traps
were preserved in the alcohol.
At biodiversity plot #1, we set
traps on the ground and 1 to 2 m
above the ground. The traps on the
ground were half buried to facilitate
access for beetles that crawled to
the traps. We placed some bait

juices within 15 cm of the traps and
prevented surface rainwater from
entering the traps by digging circular ditches. Leaf litter was scattered
on top of the traps as camouflage.
We marked each trap with a flag
containing the date, location, and
type of bait used. An equal number
of traps were hung from adjacent
vegetation (Fig. 3).
The traps were baited with
different materials. Some contained
a mixture of available fermented
fruits, including bananas, pineapples, and papayas in addition to
one tablespoon of live yeast, water,
and a few spoonfuls of alcohol all
of which were placed in a sealed
container and allowed to sit for five
days. Other traps were set with
human feces, which contain high
quantities of proteins and are
attractive to several groups of
insects. These traps were effective
for two days. We also used ocelot
(Felis pardalis) feces that we found
in the forest. Still other traps were
baited with parts of dead animals,
Figure 2. Perspective and longitudinal views of the traps used in the study. Scale =
5 cm (drawing by G. Valencia).
171
Figure 3. Schematic representation of the forest plot with the position of

the traps at San Martin-3 (drawing by G. Valencia).
including pieces of rotten fish that
we allowed to decompose for 24
hours. We tested one fish-baited
trap in the center of biodiversity
plot 1; given its efficiency, we
increased the number of traps
baited with dead animals at plot #2,
using parts of a dead snake
(Boidae) that we found in the
forest. At plot #3, we used the
internal organs and meat of
chicken.
All traps were placed systematically in pairs (ground and air) at the
sides and centers of the biodiversity
plots (Fig. 4). We also used water
saturated with salt (NaCl) as a
preservative, adding a few drops of
shampoo to break the surface
tension of the water and help
ensure that the organisms would be
drawn into the solution. These traps
were serviced every 48 hours for
eight days. Each time we serviced
172
the traps, we stored all sampled

materials inside whirl-packs and
labeled all bags with the appropriate
sampling information, including
date, type of bait, and sampling
site. At biodiversity plot #2, we
modified the placement of the traps
by setting four sets of four traps
with different baits, each separated
by 10 meters. We serviced each trap
every 12 hours to detect changes in
diurnal and nocturnal beetle fauna.
Biodiversity plot #3 had the same
design as plot #1.
All sampled material was
determined to genus and classified
to morphospecies (i.e., specimens
that had similar morphology were
classified in the same
morphospecies). We used the
following studies for classification:
Halffter and Halffter (1977),
Halffter and Martinez (1977),
Moron (1979), Blackwelder (1982),

Kohlmann (1984), Moron and
Terron (1984), and Moron et al.
(1988).
Results and
Discussion
We detected 47 morphospecies
distributed in 13 genera in three
subfamilies. The most common
genus was Onthophagus sp. with nine
species, followed by Canthon and
Eurysternus with six species. We
found a larger number (36) of
species at San Martin-3 than at
Cashiriari-2 (23 species). We had,
however, a higher trapping effort at
San Martin-3, where biodiversity
plots #1 and #2 had 19 and 30
species, respectively.
Traps placed in the ground
were more effective that those hung
from vegetation. We captured only
three individuals with the aerial
traps, possibly because of the
natural way in which these beetles
find their food. The most effective
bait was the dead snake. It is a
nutrient rich substrate that attracted

many groups of insects, including
beetles, wasps, flies, and ants. The
second most effective traps were
the baits with feces, followed by the
fruit baits.
References
Audaga, S., and G. Halffter. 1991.

Escarabajos asociados a
madrigueras de roedores (Coleoptera: Scarabaeidae,
Scarabaeinae). Folia Entomolgica
Mexicana 81: 185-197.
Bierregarard, R. 1990. Species
composition and trophic organization of the understory bird community in a central Amazonian
terra firme forest. Pages 217-236 in
Four Neotropical Rrainforests. (A.
H. Gentry, ed.) Yale University
Press, New Haven.
Blackwelder, R. E. 1982. Checklist
of the coleopterous insects of
Mxico, Central America the West
Indies, and South America. Pages
197-265 in Smithsonian Institution, Washington, DC.
6 5
Ground
10
9
Air
8
7
20m
1 2
Base Line
Figure 4. Diagram of the biodiveristy plot #1 at San Martin-3

indicating the position of the traps (drawing by G. Valencia).
173
Halffter, G. 1991. Historical and

ecological factors determining the
geographic distribution of beetles
(Coleoptera: Scarabaeidae:
Mexicana 82: 195-238.
Halftter, G., and W. Edmonds.
1982. The nesting behavior of
dung beetles (Scarabaeinae): An
ecological and evolutionary approach. Page 184 in Instituto de
Ecologa. Mxico.
Halffter, G., and V. Halffter. 1977.
Notas sobre Eurysternus (Coleoptera, Scarabaeidae,
Mexicana 37: 43-86.
Halffter, G., and A. Martinez. 1977.
Revisin Monogrfica de
Canthonina Americanos, IV parte.
Clave para gneros y subgneros.
Folia Entomolgica Mexicana 38:
29-107.
Howden, H., and V. Nealis. 1975.
Effets of clearing in a tropical rain
forest on the composition of the
coprophagous scarab beetle fauna
(Coleoptera). Biotropica 7: 77-83.
Janson, C., and L. Emmons. 1990.
Ecological estructure of the
nonflyng mammal community at
Cocha Cashu Biological Station ,
Manu Natural Park, Per. Pages
314-338 in Four Neotropical
Rainforests. (A. H. Gentry, ed.)
Yale University Press, New Haven.
Karr, J., S. Robinson, J. Blake, and
R. Bierregarard. 1990. Birds of
four neotropical forest. Pages 237269 in Four Neotropical
Rainforests. (A. H. Gentry, ed.)
174
Kohlmann, B. 1984. Biosistematica

de las especies norteamericanas del
genero Ateuchus (Coleoptera:
Scarabaeidae: Scarabaeinae). Folia
Entomolgica Mexicana 60: 3-81.
Lobo, J., and M. Morn. 1993. La
modificacin de las comunidades
de coleopteros Melolonthidae y
Scarabaeidae en dos reas
protegidas mexicanas tras dos
dcadas de estudios faunsticos.
Giornale Italiano di Entomologia. 6:
391-406.
Morn, M. 1979. Fauna de
Coleopteros Lamelicornios de la
Estacin de Biologa Tropical,
Los Tuxtlas, Veracruz, UNAM.
Mxico. Anales del Instituto
Biologa, Universidad Nacional
Autnoma de Mxico (Serie
Zoologa) 50: 375-454.
Morn, M. 1981. Fauna de
Coleopteros Melolonthidae de la
Reserva de Biosfera La Michilia.
Durango Mexico. Folia Entomolgica
Mexicana 50: 3-69.
Morn, M., C. Deloya, and L.
Delgado-Castillo. 1988. Fauna de
Coleopteros Melolonthidae,
Scarabaeidae y Trogidae de la
regin de Chamela, Jalisco, Mxico.
Folia Entomolgica Mexicana 77:
313-378.
Morn, M., and R. Terron. 1984.
Distribucin altitudinal y
estacional de los insectos
necrfilos en la Sierra Norte de
Hidalgo, Mxico. Acta Zoolgica
Mexicana, Nmero 3. Instituto de
Ecologa, Mxico.
Appendix 1. The Scarabaeidae beetle fauna (Coleoptera) at San Martin-3

and Cashiriari-2 in the Lower Urubamba region (the biodiversity plot in
which the specimens were sampled is indicated).
Orden Coleoptera
Family Scarabaeidae
Biodiversity Plot
Subfamily Scarabaeinae
Tribe Coprini
Series Pinotides
Pinotus Erichson 1847
Pinotus sp. 1
Pinotus sp. 2
Pinotus sp. 3
Series Phanaeina
Coprophanaeus dOlsoufieff 1824
Coprophanaeus sp. 1
Coprophanaeus sp. 2
Coprophanaeus sp. 3
Oxysternon Laporte 1847
Oxysternon sp. 1
Series Dichotomina
Ateuchus Weber 1801
Ateuchus sp. 1
Ateuchus sp. 2
Dichotomius Hope 1838
Dichotomius sp. 1
Tribe Scarabaeini
Series Canthonina
Canthon Hoffmannsegg 1817
Canthon sp. 1
Canthon sp. 2
Canthon sp. 3
Canthon sp. 4
Canthon sp. 5
Canthon sp. 6
Canthonidia Paulian 1939
Canthonidia sp. 1
Deltochilum Eschscholtz 1822
Deltochilumsp. 1
Deltochilumsp. 2
Deltochilum sp. 3
San Martin - 3 San Martin - 3 Cashiriari - 2

2
1
3
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
175
Appendix 1. The Scarabaeidae beetle fauna at San Martin-3 (Cont.).

Orden Coleoptera
Family Scarabaeidae
Biodiversity Plot
Pseudocanthon Bates 1887
Pseudocanthon sp. 1
Pseudocanthon sp. 2
Series Eurysternina
Eurysternus Dalman 1824
Eurysternus sp. 1
Eurysternus sp. 2
Eurysternus sp. 3
Eurysternus sp. 4
Eurysternus sp. 5
Eurysternus sp. 6
Series Onthophagini
Onthophagus Latreille 1802
Onthophagus sp. 1
Onthophagus sp. 2
Onthophagus sp. 3
Onthophagus sp. 4
Onthophagus sp. 5
Onthophagus sp. 6
Onthophagus sp. 7
Onthophagus sp. 8
Onthophagus sp. 9
Subfamily Ceratocanthinae
Ceratocanthus White 1842
Ceratocanthus sp. 1
Subfamily Hybosorinae
Anaides Westwood 1841
Anaides sp. 1
Anaides sp. 2
Anaides sp. 3
To be determined (Scarabaeinae)
Scarabaeinae sp. 1
Scarabaeinae sp. 2
Scarabaeinae sp. 3
Scarabaeinae sp. 4
176
San Martin - 3 San Martin - 3 Cashiriari - 2

1
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Long-term
Monitoring
of Arthropod
Fauna in
the Lower
Urubamba
Region
Albert Finnamore
Provincial Museum of Alberta
Introduction
Arthropods (insects, spiders,

mites, and relatives) are the most
biologically diverse group of organisms in terrestrial ecosystems,
constituting about 64% of all
known global biodiversity (Fig. 1).
Recent estimates place the global
arthropod fauna between 10 and 30
million species (Erwin 1982).
Biological data acquired for environmental monitoring and management have generally been obtained
from the megafauna and megaflora
(Finnamore 1996a), visible living
things such as vascular plants and
vertebrates that comprise between
2% and 6% of estimated global
biodiversity (Hawksworth and
Mound 1991, Hammond 1992). The
invertebrates, including the arthropod fauna, the micro-flora, and the
micro-fauna (bacteria, algae, fungi,
protozoa, etc.) account for about

95% of biodiversity and collectively form the invisible infrastructure that drives ecosystem
dynamics (Fig. 2). Invertebrates
and microorganisms are crucial
to the maintenance of
biodiversity (Hawksworth and
Ritchie 1993). Not only does
species richness of arthropods
vastly exceed that of vascular
plants and vertebrates taken
together, but biomass of
arthropods alone within natural
ecosystems can exceed that of
vertebrates (Lauenroth and
Other
19%
Vertebrates
3%
Plants
14%
Arthropods
64%
Figure 1. The known global biota
177
5%
Visible biota:
vertebrates and vascular plants
95%
Invisible biota:
arthropods, bacteria, algae, fungus, protozoa, etc.
Figure 2. Biota contributing to ecosystem managment.

Milchunas 1992, Wilson 1987). It
follows that biotic information
derived solely from the megabiota
presents a skewed view of ecosystem dynamics that can contribute to
poor management of biodiversity
resources.
Arthropods can provide information on virtually all macro- and
micro-habitats within an ecosystem.
They cover several size classes (i.e.,
micro-, meso-, and macro-fauna);
exhibit a range of ecosystem requirements (highly specific to
generalist), dispersal abilities, life
cycles, and development times;
assist in mediating ecosystem
functions such as decomposition
and in maintaining soil structure
and soil fertility; regulate populations of other organisms (including
arthropods, vertebrates, and plants);
respond quickly to environmental
changes; and act as mobile links
essential to the reproduction of
many flowering plants (Kremen et
al. 1993). Information derived from
arthropod species assemblages can
178
be used to characterize accurately

almost any aspect of an ecosystem.
The central issue in monitoring
and assessment of biotic shifts in
ecosystems is our ability to provide
spatial-temporal referencing for
biotic data. To analyze biotic shifts
associated with climate or anthropogenic change, biotic data must be
comparable in space and time and
placed within the context of natural
ecosystems (i.e., data from modified
systems must be compared with
data from the appropriate natural
counterpart). Both sampling protocols and the methods used to record
and structure abiotic data associated with the protocols must be
standardized (Appendix 1 and 2).
The ecosystem framework used to
implement a sampling and monitoring program must also be standardized (Finnamore 1996b).
Vegetation structure, considered as multiple-scale canopies,
provides the ecosystem framework
that can be used to organize the
monitoring effort from the perspective of arthropods. Data acquired
using appropriate protocols within

this framework may then be compared to data from communities
across gradients in and among
ecosystems or to analogous communities in highly disparate ecosystems. The recognition that ecosystems possess several scales of
canopies has implications both for
the design of a sampling program
and for the protocols used to conduct it. For instance, canopy sampling protocols should be applied
vertically along the aridity gradient
within sampling sites (i.e., low, midand high canopy of trees and
herbaceous vegetation) and horizontally along the hydrological
gradient represented by the transition from riparian zone to high
elevation points. The pulse fauna
(locally transient fauna), the most
diverse arthropod community
assemblage, can then be characterized using specific protocols and
not be confused with the arthropod
fauna endemic to the litter, soil,
canopy, and aquatic communities.
The latter groups can also be characterized with appropriate protocols
used in association with the respective vegetation canopy.
species or to assess changes in a
few species without a baseline
reference to the appropriate natural
ecosystem. The acquisition of
biodiversity baselines from unmodified sites is the single most important component of a monitoring
strategy. Baselines are data sets
against which similar, usually
smaller, collections of data can be
viewed in perspective to provide an
interpretation that reflects ecosystem reality. The existence of
baselines is a prerequisite for the
analysis of change. The inventory
of species, or baseline, sets the
stage for development of monitoring technology while acquiring a

database that functions as a powerful tool in the analysis of ecosystem
change. Implementation of monitoring programs can then be based
on a broad and detailed knowledge
of the species.
The acquisition of baseline
inventories is costly and limited by
the availability of skilled human
resources. For instance, the Costa
Rican All Taxa Biotic Inventory
cost well in excess of $100 million
for arthropods alone and would
have, had it proceeded, consumed
the time of nearly every trained
systematist in the world for five
years.
The Arthropod
Monitoring Strategy
for the Lower
Urubamba Project
The Camisea natural gas field
in the Lower Urubamba region is
only about 100 km west of the
Manu Biosphere Reserve. Recent
investigations (Wilson and Sandoval
1996) have found Manu to be one
of the richest regions in relation to
biodiversity known on earth. Similar
richness can be expected in the
Lower Urubamba.
For the Lower Urubamba
project, I have recommended that
the arthropod baseline be restricted
to the well sites and consider only
those groups that are taxonomically
feasible. The arthropod baseline for
each site should consist of at least
two biodiversity plots; one situated
in the forest type most likely to be
affected by gas well operations; the
other, a control plot, situated in the
same forest type in a nearby area
unlikely to be affected by gas well
Vegetation
structure,
considered
as multiple-scale
canopies,
provides
the ecosystem framework that
can be
used to
organize
the monitoring
effort for
arthropods.
179
The
information
obtained
from the
species
baselines
should be
used to
develop the
monitoring
program.
180
operations or pipeline construction.

Subsequent monitoring programs
must use the same biodiversity plots
from which the baselines were
acquired. Each baseline must also
be associated with concurrent
monitoring of the abiotic factors
that structure the ecosystem (e.g.,
climate, soil, water quality, etc.),
and tailored to accommodate
available expertise from the international systematics community
(Appendix 3). Cooperation from
systematists is essential for expert
identification of species in target
groups. The baseline and monitoring
program should be aimed strictly at
the species level. Acquisition and
processing of baselines should be
coordinated by a central national
agency (e.g., Museo de Historia
Mayor de San Marcos or another
national institution) in cooperation
with the SI/MAB Program.
The information obtained from
the species baselines should be used
to develop the monitoring program.
To be cost-effective, arthropod
biodiversity monitoring must use a
subset of species as a proxy for
total biodiversity in an ecosystem.
Species may be selected for monitoring for a number of reasons (e.g.,
threatened species, importance to
native communities, etc.), but a
central suite of species should also
be selected from the baseline study
to provide representation of arthropod size classes, dispersal abilities,
and trophic levels in the soil, litter,
and canopy communities existing
across major physical and chemical
gradients in the biodiversity plots.
The baseline acquisition and
monitoring program should provide
data that are comparable over space
and time so that fauna can be
compared spatially among sites and
temporally from year to year at a

single site. To accomplish these
objectives, the baseline must encompass the entire year at each site.
Some research is necessary to
determine the overriding environmental cues for species in the
Lower Urubamba and in particular
the climatic influence of El Nio.
Attention to seasonal standardization is necessary so that subsequent
monitoring efforts will match the
biological time periods to those of
the relevant baseline at each
biodiversity site. Biological communities seldom follow the Julian
calendar. Differences in seasonal
activity from year to year may be
standardized by recording and using
equivalent precipitation, degree-day
accumulations, flower phenology, or
a combination of such factors.
The Arthropod
Assessment and
Monitoring Program
The arthropod assessment and
monitoring program for the Lower
Urubamba is underway, according
to the basic precepts described
above. The objectives are to:
* acquire arthropod faunal
baselines for at least two
biodiversity plots including one
control plot at each of three gas
well sites to be considered for
biodiversity monitoring;
* monitor in subsequent years
selected components of the arthropod fauna representative of size
classes, dispersal abilities, and
trophic levels in the soil, litter, and
canopy communities existing across
major physical and chemical gradients in the biodiversity
* quantify the rate of
biodiversity change in arthropod
species assemblages that can be
associated with natural gas extraction;

* assess the impact of changes
in arthropod species assemblages in
the study area; and
* develop the arthropod faunal
inventory of the region using data
obtained from the baselines and the
monitoring program.
Substantial progress has been
made in acquiring initial data for
the baselines. During the next phase
of the project (Phase III), additional
work on baseline acquisition and
the main thrust of the monitoring
program will take place, as outlined
in the three steps discussed below.
Step 1. Staff Training
Training a group of competent
Peruvian counterparts is essential
to the success of the Lower
Urubamba project. When properly
executed, training ensures redundancy of expertise and provides a
measure of quality assurance.
Therefore, it must encompass all
aspects of specimen sampling,
processing, curation, development
of parataxonomists, electronic data
management, and data analysis.
Suggested training methods for the
Lower Urubamba project include
workshops in Per and Washington,
D.C. in cooperation with qualified
Peruvian specialists and training
sessions for parataxonomists conducted by appropriate specialists
the institutions where the specialists work or on site in the Lower
Urubamba.
Step 2. Acquiring the
Arthropod Baseline
A number of protocols (see
Table 1) are designed to elicit
information about arthropod community structure and species assemblages in the major habitats (soil,
litter, canopy levels, etc.) of the

Lower Urubamba biodiversity plots.
These protocols can provide a
combination of qualitative (species
richness) and quantitative (absolute
abundance and relative susceptibility to trapping) measures that
permit quantification of arthropod
species within a biodiversity plot
and comparison of species assemblages among biodiversity plots or
over time within a single plot.
The protocols should be coordinated by a field operations manager and several (five) assistants
over a three-year period (i.e., one
site per year). The field staff will
acquire the arthropod samples with
associated data, preserve the
samples, and ship them to the
processing center. The protocols
provide a methodology for the
accumulation of more than 17,800
arthropod samples in the one-year
window available for baseline
acquisition at each of three gas well
sites. That means about 50 samples
per day must be sampled, catalogued, and preserved. All sorting
will take place at the processing
center.
The processing center is responsible for sorting samples to
target groups, sorting target groups
to morphospecies, selecting several
specimens of each morphospecies
(macro-fauna, some micro-fauna)
for mounting or preparing an entire
lot (micro-fauna) when requested by
a specialist, archiving residues,
mounting selected specimens,
labelling mounted specimens,
sending specimens to systematists,
tracking loans of specimens, comparing identified specimens with
sorted morphospecies, and entering
number of specimens of each
species in the database.
One of the
objectives is
to quantify
the rate of
biodiversity
change in
arthropod
species assemblages
that can be
associated
with natural
gas
extraction.
181
Table 1. Summary of arthropod sampling protocols. (calculation of total samples is for one
gas well site containing two biodiversity plots; n.a. = not applicable)
Protocol
Code Community
sampled
Operating
time
Service
interval
Total
samples
1. BerleseTullgren
Bt, Bb
soil
12 (1)
72
6/year
7 days (2)
432
2. Sifting
Sft
litter
108 (3)
6/year
7 days (2)
216
3. Winkler
ants in litter
48
6/year
2 days (2)
288
4. Pitfall traps
PFT
ground surface 6
60
continuous 5 days
5. Baited
pitfall traps
BPFT
decay
6 (4)
48
6/year
6. Pan traps
PT
surface/pulse
70
continuous 5 days
5,110
pulse, photo +
pulse, photo -
6
6
3
3
continuous 5 days
36 MTs
108 MPTs continuous 5 days
2,628
2,628
7. Malaise traps MT
MPT
Malaise
pan traps
Habitats Replicates Total traps

per plot per habitat extractors
1 week
4,380
288
780
8. Canopy
Malaise traps
CMT
canopy
30
continuous 14 days
9. Fogging
Fog
canopy
n.a.
n.a.
30
nocturnal,
aquatic
vegetation
n.a.
12/year
n.a.
96
12/year
n.a.
360
10. Light traps BLT

11. Sweep
samples
12. Special
habitats
write
out
anywhere
30 cages
6/year
n.a.
180
13. Spot
collections
anywhere
n.a.
n.a.
n.a.
anytime
n.a.
n.a.
14. Point &

transect
PTC
butterflies
n.a.
n.a.
12/year
n.a.
192
15. Kick
sweeps
AKS
benthic aquatic
4 nets
12/year
n.a.
216
(1) Samples from six habitats are divided into top (Bt) and bottom (Bb) core sections for arthropod extraction.
(2) Extraction time.
(3) Uses same extractors as preceding protocol.
(4) Two replicates of each of three baits.
182
Sixty-three groups are proposed

for monitoring in the study area at
the well sites. The groups are
classed as aquatics and terrestrials.
Specialists have been selected to
identify specimens in each of the
groups.
Step 3. Monitoring
The arthropod baseline is
necessary to determine which
species and unique habitats should
be used for monitoring. The specieslevel data obtained from the
baselines should be used to develop
the monitoring program. To be cost
effective, arthropod biodiversity
monitoring must use a subset of
species as a proxy for total
biodiversity in an ecosystem.
Selection of individual species
in a taxonomic group should be
avoided for monitoring because of
the likelihood of year-to-year
fluctuations in population levels.
Preference should be given to
selecting a cross-section of genera
(covering the ecosystem components) where the component species
are relatively rare in the study area
baselines. It is likely that different
sets of genera will be appropriate
for different sites. This approach
not only provides better environmental partitioning for the analysis,
but it also avoids most of the
boom-bust cycles that may be
associated with relatively common
species. A change in species richness or relative abundance among
one or more groups of relatively
rare species can be used as evidence
that these groups are likely responding to a change in conditions,
human caused or otherwise, at the
well sites.
Detailed costing for the monitoring program is dependant on the
number of taxa selected for moni-
toring. However, the field program

is identical to that required for the
baseline acquisition since all sampling protocols (abiotic and biotic)
must be precisely repeated at the
biodiversity plots where the
baseline was collected. The only
difference in cost is that fixed assets
(e.g., field microscopes) that are
one-time expenditures do not have
to be repurchased for the monitoring program. Personnel costs remain
the same for field sampling in the
monitoring program.
Substantial cost reductions are
realized in the sample processing
for the monitoring program. Fixed
assets are not included in the
calculation for sample processing,
and a reduced number of target
groups are processed, thereby
saving further on labor. The difficulty of specimen identification,
especially for the physically smaller
specimens, means that a portion of
the budget must be allocated for
expert identification services.
To be cost
effective,
arthropod
biodiversity
monitoring
must use a
subset of
species as a
proxy for
total
biodiversity
in an
ecosystem.
Conclusion
Arthropods are the most diverse group of organisms in terrestrial ecosystems. Many species are
highly sensitive to ecosystem
changes and can provide valuable
advance warning that change is
occurring. The assessment and
monitoring strategy presented in
this proposal, if implemented, can
be expected to generate information
on most levels of the ecosystems
surrounding the gas well sites in the
Lower Urubamba region. Information will be gathered on the
biodiversity in the soil, litter,
surface, vegetation, and canopy
communities at the sites. An estimated 15,000 species of arthropods
will be used in the biodiversity
183
analysis at the site in the most costeffective manner possible.
References
Alonso, L. and A. Alonso. In press.

Recommended Protocols for
Measuring and Monitoring
Ground-dwelling and Aerial
Arthropods in Taiwan. In Young C.
and F. Dallmeier (eds.). Proceedings of the ROC/SI Biodiversity
Symposium. September 1996.
Anderson, R. S. 1996. Sifting and
Berlese protocols. Pages 52-53 in
The SAGE Project - A workshop
report on arthropod sampling
protocols for graminoid ecosystems (A.T. Finnamore, ed.) EMAN
Occasional Paper Series Report
No. 7.
Behan-Pelletier, V. M., A. Tomlin,
N. Winchester, and C. Fox. 1996.
Sampling protocols for
microarthropods. Pages 47-52 in
The SAGE Project - A workshop
report on arthropod sampling
protocols for graminoid ecosystems (A.T. Finnamore, ed.) EMAN
Occasional Paper Series Report
No. 7.
Briggs, J. B. 1961. A comparison of
pitfall trapping and soil sampling
in assessing populations of two
species of ground beetles (Col.:
Carabidae). Report of the East
Malling Research Station 1960:
108-112.
Crossley, D. A., Jr., and J. M. Blair.
1991. A high-efficiency,
low-technology Tullgren-type
extractor for soil microarthropods.
Agriculture, Ecosystems and Environment 34: 187-192.
Edwards, C. A. 1991. The assessment of populations of
soil-inhabiting invertebrates.
Agriculture, Ecosystems and Environment 34: 145-176.
184
Erwin, T. L. 1982. Tropical forests:

their richness in Coleoptera and
other arthropod species. The Coleopterists Bulletin 36: 74-75.
Finnamore, A. T. 1996a. The
advantages of using arthropods in
ecosystem management. A brief
prepared on behalf of the Biological Survey of Canada, terrestrial
arthropods.
Finnamore, A. T. 1996b. Summary
and recommendations. Pages 3-10
in The SAGE Project: A Workshop
Report on Arthropod Sampling
Protocols for Graminoid
Ecosystems.(A. T. Finnamore, ed.)
EMAN Occasional Paper Series
Report No. 7.
Greenslade, P. 1973. Sampling ants
with pitfall traps: digging-in effects. Insectes Sociaux 20:343-353.
Greenslade, P., and P. J. M.
Greenslade. 1971. The use of baits
and preservatives in pitfall traps.
Journal of the Australian Entomological Society 10: 253-260.
Greenslade, P. J. M. 1964. Pitfall
trapping as a method for studying
populations of Carabidae (Coleoptera). Journal of Animal Ecology
33: 301-310.
Hammond, P.M. 1992. Species
inventory. Pages 7-39 in Global
Biodiversity: Status of the Earths
Living Resources. (B. Groombridge, ed.) Chapman and Hall,
London.
Hawksworth, D. L., and L. A.
Mound. 1991. Biodiversity databases: the crucial significance of
collections. Pages 17-29 in The
Biodiversity of Microorganisms
and Invertebrates: Its Role In
Sustainable Agriculture (D.L.
Hawksworth, ed.) C.A.B International, Wallingford, England.
Hawksworth, D. L., and J. M.
Ritchie. 1993. Biodiversity and
Biosystematic Priorities: Microorganisms and Invertebrates. CAB

International, Wallingford,
England.
Krantz, G. W. 1978. A Manual of
Acarology, 2nd edition. Oregon
State University Book Stores, Inc.,
Corvallis, Oregon.
Kremen, C., R. K. Colwell, T. L.
Erwin, D. D. Murphy, R. F. Noss,
and M. A. Sanjayan. 1993. Terrestrial arthropod assemblages: their
use in conservation planning.
Conservation Biology 7(4): 796-808.
Lauenroth, W. K., and D. G.
Milchunas. 1992. Short-grass
steppe. Pages 183-226 in Natural
Grasslands Introduction and
Western Hemisphere. Ecosystems
of the World 8A. (R. T. Coupland
ed., D. W. Goodall, ed. in chief)
Elsevier, Amsterdam.
Marshall, S. A. 1996. Corpse, dung
and mammal burrow protocols.
Pages 59-63 in The SAGE Project
- A workshop report on arthropod
sampling protocols for graminoid
ecosystems (A. T. Finnamore, ed.)
Report No. 7.
Martin, J. E. H. 1977. Collecting,
preparing, and preserving insects,
mites, and spiders. The insects and
arachnids of Canada Part 1. Research Branch Canada Department
of Agriculture, Publication 1643.
Masner, L. 1996. Pan traps. Pages
57-59 in The SAGE Project - A
workshop report on arthropod
ecosystems (A. T. Finnamore, ed.),
Report No. 7.
Masner, L., and G. A. P. Gibson.
1979. The separation bag - a new
device to aid in collecting insects.
The Canadian Entomologist 111:
1197-1198.
Merchant, V. A., and D. A. Crossley,

Jr. 1970. An inexpensive,
high-efficiency Tullgren extractor
for soil microarthropods. Journal of
the Georgia Entomological Society 5:
83-87.
Mitchell, B. 1963. Ecology of two
carabid beetles, Bembidion
lampros (Herbst) and Trechus
quadristriatus (Schrank). II. Studies on populations of adults in the
field, with special reference to the
technique of pitfall trapping.
Journal of Animal Ecology 32:
377-392.
Niemel, J., E. Halme, and Y. Haila.
1990. Balancing sampling effort in
pitfall trapping of carabid beetles.
Entomologica Fennica 1: 233-238.
Norton, R. A. 1986. A variation of
the Merchant-Crossley soil
microarthropod extractor. Quaestiones Entomologicae 21: 669-671.
Pollard, E., and T. Yates. 1993.
Monitoring butterflies for ecology
and conservation. Chapman and
Hall, London.
Scudder, G. G. E. 1996. Pitfall
trapping. Pages 53-56 in The
SAGE Project - A workshop report
on arthropod sampling protocols
for graminoid ecosystems (A. T.
Finnamore, ed.) EMAN Occasional Paper Series Report No. 7.
Seastedt, T. R., and D. A. Crossley,
Jr. 1978. Further investigations of
microarthropod populations using
the Merchant-Crossley high gradient extractor. Journal of the Georgia
Entomological Society 13: 338-344.
Sharkey, M.J. 1996. Malaise/flight
intercept sampling protocols. Pages
65-67 in The SAGE Project - A
workshop report on arthropod
ecosystems (A. T. Finnamore, ed.)
Report No. 7.
185
Southwood, T.R.E. 1978. Ecological methods with particular reference to the study of insect populations. 2nd edition. Chapman &
Hall, London.
Spence, J. R., and J. K. Niemel.
1994. Sampling carabid assemblages with pitfall traps: the madness and the method. Canadian
Entomologist 126: 881-894.
Townes, H. 1972. A light-weight
Malaise trap. Entomological News 83:
239-247.
186
Wilson, D. E., and A. Sandoval,

eds. 1996. Manu: The Biodiversity
of Southeastern Per. Smithsonian
Institution, Washington, DC
Wilson, E. O. 1987. The little
things that run the world (the
importance and conservation of
invertebrates). Conservation Biology
1: 344-346.
Appendix 1. Protocols for abiotic measurements.

General
1. Create data sheet for data acquisition.
2. Georeferencing (data need to be
acquired only once):
* global positioning reference
preferably differentially corrected
latitude, longitude, altitude for each
trap or sample site.
* slope.
* aspect.
* height of trap above surface
or depth extracted below surface.
* trap or sample number.
3. Temperature: 24 hour minimum/
maximum (automatic recorders are
available).
4. Precipitation.
Pitfall, pan, and Malaise protocols

1. Vegetation structure measure
using vertical grids (vertical 1 m 2
with 10 cm grid, and 10 m2 with 1
m grid).
2. Leaf Area Index.
Aquatic
1. Turbidity (depth of light penetration).
2. pH of water at sample sites.
3. Stream width.
4. Flow rate.
5. Particle size of substrate.
Soil samples for Berlese

extraction
(see Berlese protocol Appendix 2,
one sample is retained for analysis
per vegetation type in each
biodiversity plot)
1. pH of soil at sample sites.
2. Soil organic carbon.
3. Bulk density.
4. Biomass of carbon.
5. Particle size distribution.
6. Electrical conductivity.
7. Soil temperature (max. mins.) at
surface, 5cm and 10cm depths.
8. Exchangeable Calcium status.
187
Appendix 2. Arthropod sampling protocols.

Terrestrial Arthropod
Sampling Protocols
Biological diversity, considered
at the species level, is a function of
both species richness (number of
species) and abundance (number of
individuals). All of the protocols
provide some qualitative measure
(species richness) when applied at a
site, and if applied in a consistent
manner, can provide data suitable
for comparative analysis. Most of
the protocols can also provide
quantitative measures (abundance)
in the form of either scalable
population estimates or of relative
trapability.
The arthropod sampling protocols listed below remove individuals
in small quantities from their habitats (e.g., are destructive techniques). The effect of removal of
the small quantities of arthropods is
negligible because of the density of
resident populations or because of
the dispersal abilities of nearby
populations. Consideration should
be given to placement of the arthropod sampling devices within the
biodiversity plot so as not to disturb
sampling programs for other organisms and vice versa. In the case of
highly destructive techniques where
large amounts of fauna are removed
from the habitat, such as canopy
fogging, the procedure should be
conducted well away from the
biodiversity plot (i.e., at least 0.5
km away) so as not to unduly
influence the species assemblages
of any organisms in the biodiversity
plots.
Replicates at each sample site
for each of the sampling protocols
should be determined through use
of species accumulation curves for
a variety of trophic levels (e.g.,
188
herbivore, predator, parasitoid, etc.)

within the sample. The number of
traps needed to achieve the asymptote in species accumulation should
be viewed as the number of trap or
sample replicates to be used. The
number of replicates suggested in
the following text for each sampling
protocol is a suggested, untested,
minimum. A short test period (two
weeks) to determine the appropriate
replicates for each protocol by
species accumulation analysis
should be incorporated into the
sampling program at the initial site.
1. Berlese-Tullgren Soil
Arthropod Sampling
Protocol Summary
* applicability: any soil, but
especially soils under the different
vegetation types found in the
biodiversity plots (e.g., palms,
bamboo, canopy emergent trees,
ferns, herbaceous vegetation,
riparian zone, etc.).
* placement: to depth of A
horizon.
* suggested minimum samples
or replicates per vegetation type:
three.
* total extractors (simultaneous
extraction): 72.
* operating time: procedure
should be performed a minimum of
six times per year (once every two
months) - wet and dry seasons:
early, mid and late seasons.
* extraction time: seven days.
Samples are extracted in field lab
using power from the project generator (about 1100 watts requirement).
* total samples - for one well
site containing two biodiversity
plots with six vegetation types
represented in each plot, minimum
Appendix 2. Arthropod sampling protocols (Cont.).

suggested replicates (three), division of each sample into upper and
lower subsamples, and a one-year
collecting window: 432 samples.
* sample tracking code followed by sample number
(subsamples from the same sample
should have the same numbers):
* top subsample - Bt001
* bottom subsample - Bb001
The following is modified from
Behan-Pelletier et al. (1996). The
small size of microarthropods
makes hand collection of specimens
from the substrate impractical. Thus
it is usual to collect samples of the
habitat and separate
microarthropods from the substrate
using behavioral extractors. The
sample unit can be a core or cube,
ranging from 2.5 cm to 10 cm in
diameter, or can be a specific
volume of substrate. The most
common core size used in recent
publications on quantitative sampling for soil arthropods in
agroecosystems is 5 cm diameter, to
a depth of 15 cm in soil. However,
the core size used will depend on
the habitat (Edwards 1991).
A modified Tullgren apparatus,
based on the Berlese funnel (thus
often called Berlese-Tullgren funnel), and its various modifications,
is the most commonly used method
for separating arthropods from soil
and litter. The essential component
of these extractors (see Krantz
1978) is a sample container with
wire mesh or screening on the
bottom, a metal or plastic funnel in
which, or over which, the sample
container is placed and a collecting
vessel below the funnel which
usually contains a liquid preservative, generally 70 to 80% ethanol. A
source of heat and desiccation
(light bulb, electric resistance wire)

is placed above the sample. The
objective is to create a steep gradient of temperature and moisture
throughout the sample (Edwards
1991). Microarthropods react to the
heat and desiccation by moving
downward (away from the heat) and
eventually fall through the screen at
the bottom into the preservative.
Cheesecloth below the sample and/
or a baffle in the funnel can reduce
debris from the sample falling into
the preservative as the sample dries
out or is agitated by the movement
of larger organisms. The wattage of
light bulb used in the BerleseTullgren funnel depends on the size
and water content of the sample
and on the distance of the bulb
from the sample surface. A 15-watt
bulb is recommended for soil
arthropod extraction in the Camisea
samples.
Behavioral extractors extract
only active stages; eggs, prelarva,
and quiescent premolt stages remain
in the sample. Taxa sampled include
mites, Collembola, soil Diptera, soil
Coleoptera, flightless Hymenoptera,
and numerous larval stages of a
wide variety of orders. Processing
using behavioral extractors should
be carried out as soon as possible
after field collection of samples
(Edwards 1991). At a minimum,
any study should incorporate behavioral extraction of soil arthropods.
Soil Sampling
If necessary, gently remove
growing vegetation from sample
plot using scissors or knife. Use
standard 5 cm diameter corer (bulk
density corer, or equivalent) to
remove soil sample to depth of A
horizon (usually 5 to 20 cm depth).
189

Corer constructed from
case-hardened steel pipe with 5.0
cm internal diameter; pipe with Tee
handle with cross pedal and
anti-compression ring in coring pipe
(note: in humic soils and moss a 10
cm corer is preferable to reduce
compaction, but these samples
would require a larger extractor).
Divide sample into layers of 5 cm
depth and bag (brown wax paper
bags, cloth bags; or if using plastic,
if possible leave top open) and
label separately. Keep samples cool
until they can be extracted
preferably the same day as sampled.
If samples cannot be extracted on
the same day, they should be maintained at 4 to-10 C. Arthropod
extraction techniques follow. Three
to five replicates should be taken
from each habitat within a site, and
from each plot in a management
regime. Samples should be collected
a minimum of three times in the
wet season and three times in the
dry season (more frequent in early
characteri-zation stage; twice per
season when species richness and
population trends are established).
Behavioral Extraction
Use modified Tullgren, or
Macfadyen High Gradient (MHG),
or Kempson, Lloyd and Gelhardi
(KLG) to behaviorally extract
microarthropods from soil samples.
The most effective behavioral
extractor for microarthropods is the
Macfadyen High Gradient extractor
or its refinements by Merchant and
Crossley (1970), Seastedt and
Crossley (1978), Norton (1986),
Crossley and Blair (1991). However, variations between different
modifications of MHG, KLG and
modified Tullgren are quantitative
190
rather than qualitative. For this

reason, any of the above extractors
is acceptable if the following
protocols are adhered to:
* Record wet weight of sample.
* Unless sample is very compact, place layer of cheese-cloth or
fine screen over screen of extractor
to prevent soil particles being
dislodged into the extractant.
* Invert 5 cm soil sub-samples
over screen of extractor, i.e., the
top of the sample should be closest
to the screen.
* Ensure an air-space between
sample and collecting funnel, to
avoid condensation in the funnel.
* Extract into 75% ethyl alcohol.
* Use a dimmer (rheostat)
attached to the heating source to
establish gradient between top and
bottom of sample; this gradient
should reach about 30 C by the
end of the extraction period.
NOTE: dimmers should be temperature calibrated for several days
to ensure uniformity between
funnels. Alternatively, place extractor in refrigerated cold room
(<15 C), and gradient will be
established.
* Extract for four to seven
days depending on moisture and
organic matter content of sample; if
in doubt, extract for seven days.
* Record dry weight of sample.
* Retain sample for determination of soil abiotic characteristics.
2. Sifting - Litter Arthropod
Sampling Protocol
Summary
* applicability: any litter, but
especially leaf/twig litter under the
different vegetation types found in
the biodiversity plots (e.g., under

palms, bamboo, canopy emergent
trees, ferns, herbaceous vegetation,
etc.) but also flower and fruit falls,
dead wood, and vertebrate nest
material.
* placement: 6 liters of sifted
litter from some or all of the communities indicated above. Record
number of square meters sifted to
obtain 6 liters.
* samples/replicate: three.
* total berlese extractors: 108
(use same extractors as Berlese
protocol above but employ protocols consecutively - total power
requirements: 1650 watts).
six times per year (once every two
* extraction time: seven days.
plots with six litter types represented in each plot, minimum
suggested replicates (three), and a
one-year collecting window: 216
samples.
* sample tracking code followed by sample number: Sft001.
Anderson (1996). Many small
arthropods live in various kinds of
decaying organic material. Collection of this material and the separation of the arthropods from it
reveals an assemblage of cryptic
taxa not generally collected in other
ways. For many taxa, species diversity in this assemblage is generally
high and species distributions are
often restricted, likely because
dispersal powers are often limited
by loss of wings and/or eyes.
Various forests have been sampled
for litter arthropods by sifting
general forest floor leaf litter

followed by extraction using berlese
funnels (see Berlese-Tullgren soil
protocol above and the Winkler
method below). General forest floor
leaf litter usually yields many
species, but attempts should also be
made to specialize on other focal
substrates such as flower and fruit
falls, vertebrate nest materials,
concentrations of fungi or twigs, or
similar concentrations of any kind
of organic material. Specialized
habitats in all areas should yield
valuable and perhaps unique collections. Selection of various types of
substrate debris will diversify the
sampling results but sampling effort
and diversity of substrates sampled
should be consistent across sites to
permit meaningful comparisons.
Field sampling should first
consist of selection of site and or
focal substrates. The selected
material should then be sifted using
a standard entomological sifter with
a 1-cm hardware cloth as a screen.
Litter, to the level of the soil only,
should be scraped into the sifter.
Approximately 6 liters of sifted
litter should be taken as one
sample. This sifted litter is placed in
a cotton pillowcase (or similar
receptacle), labelled and tied. Three
such samples are taken for each
selected sampling period in that
habitat or microhabitat. Samples
can be kept in the pillowcases in
the laboratory for up to four to five
days as long as they are turned
frequently and not exposed to
excess moisture or extreme temperatures. The construction of
sifters is described in Martin (1977).
Processing in the laboratory
consists of placement of 2 liters of
litter in each Berlese funnel and
191

heating, using light bulbs, for up to
seven days. Extracted insects
should fall into 80% ethanol. The
specimens extracted from three
funnels should represent a single
sample. General litter samples
should be taken once every two
months; special substrates should
be sampled when encountered. As
in all sampling protocols, feedback
should ensure the maximally efficient deployment of resources. Taxa
sampled will include small beetles,
ants, small flightless Hymenoptera
and Diptera, spiders, mites, millipedes, centipedes and numerous
larval stages of various taxa.
3. Winkler Method - Ant
Sampling Protocol
Summary
* applicability: any litter, but
especially leaf/twig litter under the
different vegetation types found in
the biodiversity plots (e.g., under
etc.) but also flower and fruit falls,
dead wood, and vertebrate nest
material.
* placement: one square meter
of litter is sifted from all the communities indicated above.
* samples/replicate: four.
* total winkler extractors: 48.
6 times per year (once every two
* extraction time: two days.
* total samples - for one gas
extraction site containing two
biodiversity plots with six litter
types represented in each plot,
minimum suggested replicates
192
(four), and a one-year collecting

window: 288 samples.
* sample tracking code followed by sample number: W001.
Alonso and Alonso (in press).
Winkler sacs are a light-weight,
portable method for extracting
arthropods from leaf litter in the
field. A 1-m 2 area of leaf litter
(including twigs, rotting branches,
etc.) is collected at each sampling
point in order to collect arthropods
that are living or passing through
the litter. The leaf litter is first
sifted through a wire mesh screen
(with 0.5 -1.0 cm openings) and
collected in a bag for transportation
to the field camp or laboratory. The
main advantage of this technique is
that it does not require electricity.
The principle of the winkler extractor is to hang a thin layer of leaf
litter in mesh bags for at least 48
hours, during which time arthropods
move through the litter and out of
the mesh bag. The bags are hung
inside a larger sac so that the
arthropods cannot escape, but
instead fall to the bottom into a cup
of ethyl alcohol, which kills and
preserves them.
Winkler sacs are made of lightweight cotton or other fabric (ripstop nylon) that is hung around two
rectangular metal frames (29 x 15
cm) separated by 36 cm. The sac is
open at the top for insertion of the
litter sample, and is closed off with
a cloth tie during extraction. The
bottom of the sac is fitted with a
metal ring (5.5 cm diameter) into
which a plastic cup or bag is inserted. The cup should be 1/4 filled
with ethyl alcohol. A mesh bag (28
cm x 35 cm) is inserted into the sac

and attached to the sides of the sac
with two metal clips.
In preparation for extraction,
each leaf litter sample is placed in a
separate mesh bag. The mesh bags
have ties along their lengths to keep
the bags from becoming too thick
with litter. Since the extraction
works by arthropod movement from
within the litter to the outside, it is
important to keep a thin layer of
litter in the mesh bags so that the
arthropods reach the outside
quickly. Once the mesh bag is
inserted into the winkler sac, the
top and bottom of the sac are
closed off with their attached ties.
The winkler sac is then hung vertically. As the active arthropods leave
the mesh bags, they fall to the
bottom of the sac and into the cup
of alcohol where they are preserved.
At the end of the 48-hour
period, the arthropods should be
removed from the litter extractors
and sorted from the sample. This
process may take from two to four
hours for two people, depending on
the ability of the researchers to
distinguish arthropods in the bottom of muddy cups. This is an
important step, and should be done
with care so as not to miss any
arthropods, some of which are
microscopic. Specimens should be
placed in vials of alcohol and
completely labelled with collection
method, sample number, date, and
collectors name.
The Winkler method samples
arthropods that are active in the soil
and leaf litter. It is especially
effective for Formicidae (ants),
Araneae (spiders), Coleoptera
(beetles), Collembola, and some
Orthoptera (e.g. grasshoppers).
4. Pitfall Traps
Summary
* applicability: ground level
terrestrial habitats, but especially
under the different vegetation types
found in the biodiversity plots (e.g.,
* placement: in ground under
canopy.
* suggested minimum trap
replicates per vegetation type: five.
* total pitfall traps (two
biodiversity plots): 60.
* operating time: continuous.
* service interval maximum:
five days with salt in traps as a
preservative, 24 hours without salt.
plots with six vegetation types
suggested replicates (five), a fiveday service interval, and a one-year
collecting window (73 sample
times): 4,380 samples.
* sample tracking code followed by sample number: PFT001.
The following is adapted and
modified from Scudder (1996).
Pitfall trapping is a good method of
sampling arthropods because of its
simplicity and ease of operation
(Greenslade and Greenslade 1971).
It is an effective and cheap way of
surveying the ground surface-active
arthropods in terrestrial ecosystems,
and allows for comparison of
assemblages in different habitats.
This method of trapping must
be used with discretion (Greenslade
1964). It must be viewed cautiously
if used to provide comparative
estimates of species abundance
across habitats (Greenslade 1964,
193

Niemel et al. 1990, Spence and
Niemel 1994). Unless it is correlated with independent measures of
population numbers, pitfall trapping
should not be used for population
estimates. However, abundance
measures obtained from pitfall
samples can be compared on the
basis of the relative trapability of
species within or between sites.
Pitfall trapping, as the name
implies, involves the capture of
ground surface-active arthropods
that fall into a pit-like trap sunk
into the ground. It is recommended
that plastic 450 ml beer mugs be
used as traps. Disturbance of the
surroundings should be minimized
when first inserting these into the
ground, and traps should not be
removed for servicing. Although
pitfall traps can be emptied with a
hand-operated or mechanical suction apparatus (Southwood 1978),
ground disturbance can also be
avoided by using two beer mugs
nested one inside the other. The
beer mugs from different manufacturers are often of different
dimensions, and should be selected
so as to fit snugly inside each other,
with rims touching. The outer
container can then be left in the
ground permanently, and the inner
container removed for collection
extraction.
If traps are to collect only
ground-active arthropods, then
opaque plastic square covers (15-30
cm) should be used, raised 2.5 cm
off the ground over the traps. These
covers should be weighted with a
stone and supported by 7.5-15 cm
square wooden or hollow aluminum
rods, set at right angles around the
trap mouth to form an X. These
support rods should touch the
194
mouth of the trap for efficient

capture of specimens. Support rods
arranged in this fashion not only
support the cover firmly, but also
intercept animals moving near the
trap and funnel them into the trap.
The covers serve to keep out debris
and prevent the entry of animals
and precipitation from above.
If pitfall traps are used for live
trapping, or are baited, they must be
serviced at frequent intervals to
avoid spoilage or predation within
the trap. Sampling over periods
longer than one or two days requires
use of a preservative in the trap. If
traps are serviced at weekly intervals, a saturated saltwater solution
with a few drops of soap can be
used. The soap acts as a surfactant
destroying the surface tension
thereby drowning any arthropods
that fall into the trap. Soap should
be clear with as little odour as
possible so as not to repel
arthropods (e.g., liquid dishwasher
detergent).
At least five traps should be
installed at each sampling site.
Number each trap separately with a
unique identifier on a plastic stake
placed near the trap and always
record the number of the trap with
each sample. Statistical analyses
usually require that arthropods at a
site have an equal chance of entering each trap. To accommodate this
requirement, traps should be placed
fairly close together. Five individual
traps set up in a circle of 10 m
radius to form a trap circle meet
these requirements.
Content of traps can be collected by straining the preservative
through a fine mesh sieve and
rinsing the contents of the strainer
into a specimen container with

ethanol. Each sample should be
given a unique number to identify
the trap and trapping session. The
plastic traps should be kept clean,
because dirty traps may aid escape.
In order to fully sample any site
or habitat, trapping should be
continued through the whole of the
active season, the entire year in
tropical ecosystems. Such an extended trapping period is needed to
accommodate the different phenologies exhibited by the various
taxa. Catches obtained immediately
after a pitfall trap is placed in
position are sometimes found to be
higher than samples obtained
subsequently. These digging-in
effects (Greenslade 1973) are
evidently owing to local initial
depletion of individuals.
Pitfall traps set up as described
above are effective in capturing
terrestrial isopods and amphipods,
pseudoscorpions, scorpions,
solpugids, spiders, harvestmen,
ground dwelling crickets, true bugs,
carabid beetles, ants, and many
other taxa.
The rate of capture of
arthropods by pitfall traps depends
on the activity and population
density of each species captured
(Mitchell 1963, Greenslade 1964).
Temperature and humidity also
affect activity and locomotion, and
hence capture rate (Briggs 1961,
Greenslade 1964). Vegetation
structure can also greatly influence
trap catches (Greenslade 1964,
Baars 1979), by influencing locomotion ability especially close to the
traps. Nevertheless, pitfall trapping
remains the only realistic way to
survey large areas where qualitative
inventory and comparison of species assemblages of ground-active
arthropods is required (Spence and

Niemel 1994), and cheap and easy
to operate methods are needed
(Greenslade and Greenslade 1971).
5. Baited Pitfall Traps
Summary
under the different major vegetation
types and elevations found in the
biodiversity plots (e.g., bamboo,
herbaceous vegetation, high elevations, riparian zone, etc.).
canopy at various elevations.
replicates per vegetation type or
elevation: two of each bait (fish or
chicken, human feces, and fruit).
* total baited pitfall traps (two
* operating time: six times per
yearone week in early, mid-, and
late wet and dry seasons.
one week.
plots with two major vegetation
types and two elevational extremes
suggested replicates (two of each
bait), and a one-year collecting
window (six sample times): 288
samples.
* sample tracking code followed by sample number:
BPFT001.
Marshall (1996). Communityspecific collecting techniques can
be used to get a very quick snapshot
of the discrete faunas associated
with carrion, faeces, and fruit.
Techniques for dung, fruit and
195

carrion sampling are generally
similar, and rely on attraction to the
substrate, emergence from the
substrate, or a combination of
attraction and emergence. Simply
adding bait to an existing sampling
device such as a Malaise trap,
intercept trap, or pan trap can draw
the desired fauna, but most studies
of dung and carrion insects utilize
traps specifically designed to
sample insects associated with
specific substrates. Attraction traps
are of various designs but can be
loosely divided into pitfall-type
traps which take insects which fall
into a container below a bait, and
aerial traps which take insects as
they fly up from a bait. Because
standardization is important to our
study, careful consideration must be
given not only to the type of trap
but also to the size, age, and source
of dung, carrion, or fruit. A dead
fish will not attract the same fauna
as a dead mouse, and baits made
from deer dung and tapir dung will
attract different insects in identical
habitats. Even such minor differences as the degree of shade can
change the species taken in a trap.
Pitfall trapping is by far the
most widely used approach to
sampling dung and carrion insects.
The most effective method is a
simple uncovered pitfall trap. While
a great variety of sizes and type of
bait would be required to effectively sample all dung and carrion
insects, small muslin-wrapped balls
of carrion (fish or chicken wings are
popular choices), fruit or human
dung suspended within the pitfall
trap will attract a large proportion
of each respective arthropod community. Pig dung is almost as
attractive as human dung, and bait
196
balls can be made up in bulk then

frozen in appropriate sized lots for
the planned sampling program. The
trapping fluid should consist of salt
water and soap for sampling periods
of up to one week. Carrion-baited
traps can be enclosed in chicken
wire or hardware cloth to prevent
destruction of the traps by scavengers. Number each trap separately
with a unique identifier on a plastic
stake placed near the trap and
always record the number of the
trap with each sample.
Baited pitfall traps sample a
narrower range of insects, in carrion
and dung traps principally carrion
beetles (Scarabaeidae,
Staphylinidae, Silphidae), Diptera
and crickets; but other groups are
attracted to fruit-baited traps.
6. Pan Traps
Summary
under the different vegetation types
elevational high points, riparian
zone, etc.).
canopy.
elevation: five.
* total pan traps (two
five days with salt in traps as a
preservative, 24 hours without salt.
plots with five vegetation types and
two elevations represented in each

plot, minimum suggested replicates
(five), a five-day service interval,
and a one-year collecting window
(73 sample times): 5,110 samples.
* sample tracking code followed by sample number: PT001.
The Following is modified from
Masner (1996). Many insects are
attracted to the colour yellow,
especially bright yellow. Thus,
yellow containers filled with water
can be used to trap them. Detergent
is added as a surfactant to break the
surface tension of the water causing
the insects to drown, and salt is
added as a preservative.
Pan traps may be made from a
variety of sources including yellow
garbage bags, yellow vinyl sheets,
transparent food trays spray-painted
with acrylic yellow on the outside,
aluminum roasting pans sprayed
with yellow enamel on the inside.
The recommended type for the
biodiversity plots are yellow plastic
bowls (270 mm diameter x 78 mm
depth). Depth of the trap can vary
depending on regional aridity
(deeper to contain more water in
dryer regions or for longer servicing
intervals) but the surface area
(diameter) should remain constant.
The color should be bright yellow
(e.g., Safety Yellow). The water can
either be from a tap or from natural
sources, the latter should be filtered
through the aquarium net to remove
insects and mites. Detergent should
be clear and unscented so as not to
act as an insect repellant (e.g.,
liquid dishwasher detergent).
Equipment consists of the pan
traps - yellow plastic bowls, water,
pure detergent, salt, aquarium net,
sample bags, alcohol, soft-lead
pencil, quality label paper or card
stock, and a container for carrying
water. Self-sealing plastic bags (i.e.,

whirlpak type) are most convenient
for field specimen storage. Alcohol
is preferably 90% ethanol but 70%
isopropyl will work and is available
in most countries.
The pans should be sunk into
the ground with their rims level to
the surface. Water, salt and detergent are then added to the trap. The
traps are filled about 3/4 with
water and enough salt to make a
saturated solution. About five drops
of detergent are added as a surfactant. If traps are to be emptied
every two days or more frequently
then no salt is needed but with
longer servicing intervals (to a
maximum of seven days), salt is
essential. Number each trap separately with a unique identifier on a
plastic stake placed near the trap
and always record the number of
the trap with each sample.
Traps are serviced by scooping
out the contents with an aquarium
net and then either reusing the old
solution or replacing it with fresh
water, salt and detergent. Before
scooping remove large objects such
as leaves, twigs, small animals, etc.
that may have fallen into the trap.
The net should be dragged gently
near the bottom in one direction
several times until the contents are
recovered. Specimens can be processed for each trap or pooled
depending on the objective of the
sampling program. Check trap to
see if it is still level, top up with
water (if necessary), and add more
salt and detergent at each servicing.
Most importantly, ensure that the
solution is transparent enough for
the yellow background to show
through.
197

Specimens are transferred to
plastic bags from the net using a
squeeze bottle of water to remove
last specimens if each trap is to be
processed separately (label in each
bag), or place the net in the bag
with specimens to prevent drying if
sample is pooled. Specimens are
processed the same day in a lab or
base camp by rinsing the contents
of each bag in the net under a
gentle stream of water for several
minutes to remove dirt, salt and
detergent. Pick out any remaining
large debris. Fresh water washing is
essential because the detergent and
dirt will form a film on the insects
if they are placed directly in alcohol. This film is difficult or impossible to remove once specimens are
in alcohol, thus rendering the
specimens much less useful. Invert
net contents into whirlpak and
using a squeeze bottle filled with
alcohol gently wash contents out of
net and down sides of whirlpak to
bottom. Cover sample in the
whirlpak with alcohol to at least
level of sample volume.
Sample labels must be placed
inside the bag with the specimens.
Labels should be printed in softlead pencil or India ink on a small
piece of good quality paper or card
stock. Poor quality paper will
disintegrate or printing will slough
off.
Store specimens in a cool dark
place (refrigerate if possible), but
not in direct sunlight or near a heat
source. Replace alcohol with fresh
80% alcohol after one or two
weeks. The sooner the catch is
processed the better. For long-term
storage (several months to years) it
is essential to replace alcohol (to
prevent freezing) and to keep the
198
catch below freezing, preferably in a

deep freeze at -10 C or lower to
prevent specimen deterioration.
Material to be sent through the mail
should be packed in hard protective
containers such as mailing tubes or
strong cardboard boxes. Bulk and
weight can be greatly reduced by
carefully draining out most of the
alcohol from each whirlpak and
replacing it with a small quantity of
fresh 95% alcohol and washing the
contents to the bottom of the
whirlpak. Squeeze out air from the
whirlpak and close by gently rolling
down the end with twist-tie towards
the other end and clamping it.
Removal of air and addition of only
a little alcohol to the whirlpak is
essential to prevent drying, sloshing
and breakage of material during
shipment.
The most frequent causes of
damage to collected material are: 1)
weak salt solution in traps; 2)
drying out of material; 3) failure to
rinse in fresh water before adding
alcohol; and 4) failure to prevent
sloshing of specimens during
transportation.
Pan traps sample a variety of
surface-active taxa including spiders, Hemiptera, Orthoptera,
Diptera, Coleoptera, and
Hymenoptera among others.
7. Malaise Traps
Summary
* applicability: terrestrial
habitats, but especially in the
different herbaceous vegetation
types found in the biodiversity plots
(e.g., palms, bamboo, ferns, other
herbaceous vegetation, elevational
high points, riparian zone, etc.).
* placement: ground level.

elevation: three.
* total Malaise traps (two
* total pan traps (grey) under
Malaise: 108.
* operating time: continuous
five days with salt water in trap
heads as a preservative, 24 hours
without salt.
plots with four herbaceous vegetation types and two elevations
suggested replicates (3), a five-day
service interval, and a one-year
times): 2,628 Malaise head samples
and 2,628 pan trap or trough
samples. Total samples = 5,256.
* sample tracking code followed by sample number: MT001
for Malaise heads and MPT001
(sample numbers should be identical in associated samples) for pans
or trough under Malaise.
The following is modified after
Sharkey (1996). The Malaise trap is
constructed according to the specifications of Townes (1972). In that
article Townes provides detailed
measurements for construction of a
Malaise trap. Those measurements
will not be repeated here.
Malaise traps catch flying
insects. Most flying insects that hit
the central panel of the Malaise
trap respond by flying upward,
presumably in an attempt to fly over
the barrier. The slanted roof directs
insects toward the high point of the
trap and a bottle of salt water (with
a drop of soap) into which they are
collected. There are two modifica-
tions that should be applied to the

Townes style trap. The first is a
strip of black material about 2 m
long and 15 cm wide that is placed
along each edge of the roof of the
trap. This acts as a barrier to insects
that try to fly out of the trap. The
result is that many more fast flying
insects with good eyesight are
captured. The second modification
is the size of the mesh that is used.
Instead of 10 meshes per cm as
suggested by Townes (1972), a
mesh size of at least 24 per cm has
been shown to increase the diversity
of the catch albeit at the expense
of some of the larger specimens
with better visual acumen.
Those insects that drop and
those that fly down (e.g., many
Pompilidae) when encountering the
central panel of the Malaise trap are
not sampled efficiently by Malaise
traps. To capture this component of
the insect fauna pan traps are
placed on the ground under the
central panel of the trap. The
trough (or pan traps) runs the
length of the trap (about 2 m), is
about 7 cm deep and 60 cm wide.
The trough or pan traps should be
painted grey or the trough lined
with thick, grey vinyl that is sealed
at the corners. It can be filled with a
variety of fluids. Minimally the fluid
consists of water saturated with
salt, and a small amount of detergent (20 ml) as a surfactant. The
trough should be flat and almost
full so as to maximize the time
required for evaporation. If pan
traps are used along the bottom of
the central panel they should be
treated as a single sample and
associated with the concurrently
collected Malaise trap head sample.
Number each trap separately with a
199

unique identifier on a plastic stake
placed near the trap. Indicate on the
number whether the sample is from
the Malaise head or pans and always
record the number of the trap with
each sample.
This combination trap now
functions in two different ways: 1) a
Malaise trap to capture flying
insects; and 2) a flight intercept trap
to capture insects that drop when
encountering the central panel
(mostly beetles). Sinking the trough
or pans is not recommended if
surface-active fauna is to be analyzed separately from the actively
flying community. Use of yellow
trough or pan traps is also not
recommended if the faunal component that drops downward is to be
analyzed separately from the component attracted to yellow.
Perturbations by large mammals
(especially cattle, deer, and possibly
tapir) can be a serious problem with
Malaise traps. Avoid placement of
traps on or near game trails and
construct exclosure fencing for
cattle if necessary. Fencing should
be constructed of materials that do
not present a barrier to insect flight
so as not to cause insects to fly
upwards over the fence and over
the trap.
It may be sufficient to place
one of these traps in each community sampled if species richness is
the only parameter under consideration. Malaise trap replicates should
be considered to avoid problems
associated with trap damage or loss
and to demonstrate that existing
richness has been adequately
sampled. Three replicates are
suggested as the minimum in richness studies; depending on the
200
statistical design additional replicates may be required in quantitative studies.

Malaise traps sample the following groups in order of greatest
to least proportion of catch:
Diptera, Hymenoptera, Homoptera,
Coleoptera, followed by a small
amounts of a wide variety of other
groups.
8. Canopy Malaise Traps
Summary
* applicability: dominant tree
species found in the biodiversity
plots (e.g., palms, bamboo, canopy
emergent trees, elevational high
points, riparian zone, etc.).
* placement: mid-upper canopy.
* suggested minimum replicates: three.
* total canopy Malaise traps
(two biodiversity plots): 30
* service interval maximum: 14
days with alcohol and six drops
propylene glycol in trap heads as a
preservative.
plots with three herbaceous vegetation types and two elevations
suggested replicates (three), a 14day service interval, and a one-year
times): 780 samples.
* sample tracking code followed by sample number: CMT001.
Canopy Malaise traps are
similar in construction to the
Malaise trap but oriented on a
vertical axis with the collecting
head located at the highest point.
The four vertical central panels
intercept flying insects most of
which are eventually collected in

the collecting head at the top of the
trap. The body of the canopy
Malaise trap is constructed with a
coarse mesh instead of a fine mesh
that is used in the Malaise trap.
Most of the mesh is dyed black but
the top of the trap is white, thereby
encouraging a positive phototaxic
response by the insects and concentrating them in the head region of
the trap. The central panels are kept
open by light weight structural
supports at each end.
The canopy Malaise trap is
suspended with twine from a branch
about two-thirds of the distance to
canopy height. Several techniques
can be employed to position the
traps in the canopy. These include
climbing the tree, throwing a stone
attached to a light-weight twine,
use of a slingshot, a bow and arrow,
or fishing rod and spinner, among
others. Once the line has been
secured to a suitable branch it is
attached to the top of the canopy
Malaise trap. Another line is attached to the bottom of the trap.
The two lines can now be used to
raise and lower the trap or to maneuver around other branches.
The difficulty of moving these
traps through the canopy means
that longer servicing intervals are
more convenient for these traps.
The preservative used in the collecting head is alcohol with a few
drops of propylene glycol. The
servicing interval is once every 14
days. Number each trap separately
with a unique identifier on a plastic
tag placed near the trap and record
the number of the trap on the label
in each sample.
Canopy Malaise traps sample
insect groups similar to the Malaise
trap at ground level with Diptera,
Hymenoptera, Homoptera and

Coleoptera prominent in the
samples. However, the species
found in the mid-upper canopy
differ from most of those collected
at ground level.
9. Fogging
Summary
* applicability: dominant tree
species found outside the
biodiversity plots (e.g., palms,
bamboo, canopy emergent trees).
* placement: canopy.
* suggested minimum replicates: three.
* total traps: not applicable.
* operating time: once in wet
season, once in dry season.
not applicable.
site with five tree species (selected
from biodiversity plots), minimum
suggested replicates (three), a oneyear sampling window (two sample
times): 30 samples.
* sample tracking code followed by sample number: Fog001.
Among the most visible of the
sampling techniques, fogging
employs a contact insecticide to kill
all insects in the canopy of a tree.
The technique is among the most
effective known for building faunal
inventories particularly of the larger
arthropods. Unfortunately, fogging
is highly destructive and would
affect information obtained with
other protocols if it were conducted
in the biodiversity plots. Fogging
should be conducted for inventory
purposes in forest types as similar
as possible to the biodiversity plots
but located at least half a kilometre
outside the plots.
201

10. Light traps
Summary
* applicability: nocturnal fauna
at high and low elevational points
in the biodiversity plots.
* placement: high and low
elevations.
* samples/replicate: na.
* total light traps (two
biodiversity plots): four.
* operating time: once/month
over two consecutive moonless
nights; operate for three hours
commencing at dusk.
* service interval: na.
plots with two elevations represented in each plot, run over two
consecutive nights, and a one-year
times): 96 samples.
* sample tracking code followed by sample number: BLT001.
Many nocturnal insects use
light reflected from the moon or the
horizon as a navigational aid. As a
result they can be attracted to local
artificial sources of light when the
local light source masks the distant
light source. Light trapping is one
of the most effective means of
collecting large numbers of nocturnal and adult aquatic insects. A
number of light traps are commercially available but almost any light
will work. The black light trap is
among the most effective. Fixtures
and filtered ultra violet tubes are
available at larger commercial
hardware outlets or the entire light
trapping system can be purchased
from entomological supply companies. The light can also be placed in
front of a vertical white sheet.
Insects attracted to the light will
land on the sheet where they can be
202
collected. Commercially available

light trap systems incorporate a
collection chamber as part of the
trap. These systems can be left
unattended until the following day
when the insects can be collected.
For best results light traps should be
operated on a warm moonless night
away from competing light sources.
In many field conditions batteries or generators are used as a
source of power. A timer switch can
be used to automatically start and
stop the lights when they are placed
in remote, rugged terrain. Because
of changing night to night conditions (cloud cover, precipitation,
relative humidity, temperature, etc.)
lights in different plots should be
operated simultaneously in order to
produce comparable samples. The
climatic conditions under which the
sample was collected should be
recorded and associated with the
sample. Subsequent monitoring
should attempt to duplicate as
much as possible the climate conditions under which the baseline
collection was obtained.
Light traps are most effective
for nocturnal members of the
following groups: Lepidoptera,
Diptera, aquatic Hemiptera, Coleoptera, and Hymenoptera.
11. Sweep Samples
Summary
* applicability: anywhere but
especially herbaceous communities
bamboo, ferns, other herbaceous
vegetation, elevational high points,
* placement: anywhere.
* suggested minimum replicates
per vegetation type or elevation:

five minute sweeps repeated three
times.
* total sweep nets: six.
* operating time: once per
month.
not applicable.
plots with three vegetation types
and two elevations represented in
each plot, minimum suggested
replicates (three), conducted once
per month: 360 samples.
* sample tracking code followed by sample number: S001.
Sweeping is one of the most
effective methods for quickly
capturing large quantities of insects
in vegetation. A standard or reinforced insect net (for construction
see Martin 1977) is employed in a
sweeping motion on vegetation to
dislodge the arthropods. Arthropod
samples collected with this technique are affected by time of day,
moisture, temperature, mesh size of
the net, and subjectivity of the
collector. In the latter case different
collectors focus their collecting
efforts on different aspects of the
vegetation. Because of the difficulty in standardizing these variables sweep samples, particularly
for the meso- and microfauna
should be used only in qualitative
comparisons. The macrofauna (> 2
cm) is relatively less affected and
can be used (with caution) in
quantitative comparisons.
The sweeping motion concentrates arthropods and debris in the
apex of the net. Arthropods can be
separated from debris using a
separation bag (Masner and Gibson
1979). Separation bags use the
positive response to light exhibited
by many arthropods to separate

them from debris. Specimens can be
killed in water with a few drops of
soap added as a surfactant to break
the surface tension. They are then
transferred to whirlpacks with
alcohol added as a preservative.
Specimens can then be stored and
later mounted, labelled, identified
and added to the database. Sweep
nets are inexpensive and commercially available. A sturdy wide-mesh
net with canvas reinforced mouth is
recommended.
12. Special habitats
Summary
* applicability: derived soils,
fungi, logs, bamboo internodes,
fermenting bamboo shoots, etc.
* placement: not applicable.
* total extractors: use existing
Berlese-Tullgren and six emergence
chambers.
* suggested minimum samples:
opportunistic, three if possible.
* operating time: as special
habitats are discovered, or once
every 2 months if easily accessible.
* extraction time: seven days
for Berlese-Tullgren.
plots with five special habitat types
suggested replicates (three), a twomonth sampling interval (12/year):
180 samples.
* sample tracking code followed by sample number: none
write out habitat followed by
sample number.
A number of arthropod species
are associated with discrete communities within an ecosystem. These
species assemblages are generally
not adequately sampled by other
203

means or if they are sampled, they
can not be analyzed separately from
the other arthropod fauna. In the
Camisea region particular attention
should be given to the derived soil
community where a large, unique
arthropod fauna can be expected.
Derived soils are arboreal, they are
formed when accumulations of
leaves and twigs decompose in tree
bowls or collect in branch bases.
They can also be found beneath the
epiphyte community in many
canopy emergent trees. These soils
should be collected and extracted in
Berlese-Tullgren funnels according
to the protocol set forth above.
Other communities that can be
extracted in the same manner are
fungi and rotting logs. In the latter
instances it is desirable to extract
arthropods independently from each
species of fungi or wood encountered and where possible record the
host species (species name, photograph, and preserve an example of
the specimen for fungi).
Bamboo internodes and fermenting bamboo shoots (i.e.,
broken young shoots) are examples
of other special habitats that contain specialized arthropod fauna.
Bamboo internodes that have been
punctured as a result of insect,
rodent, or human activity fill with
water and soon attract a resident
arthropod fauna. The water-filled
internodes can be collected and
placed in emergence chambers
where the arthropod fauna can be
allowed to develop to adult stages
when it can then be captured in the
emergence cage and preserved. The
water in the internodes should then
be strained through a fine mesh
aquarium net to collect any aquatic
mite fauna. Similar procedures can
204
be used for the fermenting bamboo

shoots and the fungi and wood
mentioned above.
13. Spot collections
Summary
* applicability: anywhere.
* placement: anywhere.
* samples/replicate: not applicable.
* total traps: not applicable.
* operating time: minimum
once/month.
not applicable.
* total samples: opportunistic few specimens.
* sample tracking code followed by sample number: G001.
Among the most traditional
methods for arthropod sampling,
hand collecting remains the technique of choice for many professionals. Vegetation, substrate, or
any habitat of interest can be
visually examined for arthropods.
For example rocks and logs can be
turned over (and subsequently
replaced to retain the microhabitat
for future use) and any arthropods
captured for identification. Species
obtained in this manner can be used
only to supplement the species list
for a site. Arthropod samples
collected with this technique, like
sweep samples, are affected by time
of day, moisture, temperature,
visual acumen of the collector, and
subjectivity of the collector. Specimens can be killed in water with a
few drops of soap added as a
surfactant to break surface tension.
Specimens can then be mounted
(Martin 1977), labelled, identified
and added to the collection.

14. Point Counts and
Transect Counts for
Butterflies
Summary
* applicability: along the grid
that divides each quadrat of the
biodiversity plots.
* placement: alternate transect
counts and point counts every other
day, for 8 days in the mornings and
in the afternoon.
* samples/replicate: two (one
researcher at each of two
biodiversity plots)
* total traps: na.
* service interval: na.
* total observations: for one
well site containing two biodiversity
plots, using two observation techniques, four replicates of each
technique, and a one-month observation interval (12/year): 192
samples.
* sample tracking code: PTC
Transect counts and point
counts for butterflies require good
field identification skills on the part
of the observer. Butterfly bait traps
and general net collecting are the
best ways to become familiar with
the butterfly fauna of an area.
Butterfly bait traps are made up of
a cylindrical net (0.90 m height,
0.20 m diameter) placed above a
platform containing a bait. Most
aerial arthropods have the tendency
to fly upward after feeding or when
they are startled. Thus, traps are
designed to allow the organisms
access to the bait, then capture
them as they fly upwards after
feeding. The baits usually consist of
rotten fruit, carrion, or dung.
The methodology for transect
counts consists of counting and
identifying individual butterflies as
they fly (Pollard and Yates 1993).

Thus, the researcher needs to be
familiar with the fauna of the area
under study. Visual identifications
are restricted to an area 5 m in front
and above the researcher as she/he
walks, at a slow pace, along a
transect (e.g., walking on the grid
and perimeter of the standard 1 ha
plot established by the biodiversity
program). The researcher should
take at least five minutes on each
100 m of the transect (20 meters/
minute). Two point counts should
be done during sampling days, in
the morning (8:00 to 11:00), and
early afternoon (13:00 - 16:00).
Sample periods should be done
under standard weather conditions
(e.g., temperature, cloud cover).
The starting point and direction of
the transect should be selected
using a random number table.
Another method that is recommended is a modification of the
point count method used by ornithologists. This method also consists of visual identifications of
individuals as they fly, thus requiring a knowledge of the fauna. Here,
the researcher stays for 30 minutes
at several selected points (such as
each of the corners of the biodiversity plot) and records observations
of all individuals and species as
they fly. Observations should be
taken in the morning and again
during the afternoon. Sample
periods should be also done during
standard weather conditions.
15. Aquatic kick sweeps
Summary
* applicability: aquatic habitats,
but especially in the different
stream and small pool substrates
205

sand, coarse sand, small stones,
etc.).
* placement: in different substrates.
* suggested minimum replicates
per stream substrate: three.
* total kick sweep samples per
sampling interval (two biodiversity
plots): 18.
* operating time: once per
month.
not applicable.
plots with three stream substrates
suggested replicates (three), conducted once per month (12/year):
216 samples.
* sample tracking code followed by sample number - AKS001.
Aquatic kick sweeps are an
efficient method of sampling the
bottom dwelling arthropod fauna in
shallow water bodies. The technique works by disturbing the
stream or pond substrate and
collecting the debris with a net. A
fine mesh net is attached to a
standard butterfly net handle. The
collector, using the feet to agitate
the substrate causes the fauna to
swirl upwards into the stream
current or water column in a pool.
The net is passed continuously
through the dislodged material
collecting any suspended
arthropods. The procedure should
be continued for five minutes in any
one substrate and then be replicated
twice (minimum) more in similar
substrates and flow rates.
The sampled material can be
bottled with some stream or pond
water for sorting later the same day.
Care should be taken to keep the
206
sample cool, and out of direct

sunlight. Sorting is done live by
pouring small portions of the
sample into white trays, waiting
several minutes for debris to settle,
then picking out any arthropod that
moves. Mites should be placed
directly into Koenikes solution to
preserve colour and to prevent
collapse or shrinkage of soft tissues.
Other arthropod material may be
preserved in 70% alcohol to await
further processing.
Baseline Sample Processing
In arthropod sampling operations of this scale, acquisition of
samples and sample processing are
conducted as two separate processes. Separation of the processes
achieves a substantial reduction in
the cost of field operations
(amounting to about 10% of entire
project budget) while adding only
slightly to the cost of processing.
Furthermore, sample tracking can
be streamlined and quality control
and assurance is more dependable
both in the field and in the processing centre.
Choice of a suitable processing
centre (museum) is critical. Processing the specimens in Per transfers
control of specimen deposition to
the host country and assures that
the opportunity to develop a specimen-based national biodiversity
database is maintained in Per. It
also demands a long-term commitment by foreign institutions to the
museum selected as the processing
centre. The selection of a Peruvian
museum to act as a processing
centre for the Camisea biodiversity
samples should be guided by the
following considerations: 1) the
museum should have suitable space

both to process the material and for
subsequent storage of specimens; 2)
the museum management should be
receptive to large scale biodiversity
initiatives; 3) the amount of overhead, if any, charged by the museum; and 4) the loan policy of the
museum which should, if necessary,
be amended to accommodate the
Camisea project. If a suitable
museum cannot be identified then
consideration should be given to
establishing a new Peruvian entity
to deal with large-scale specimenbased collections (see Appendix 2
for further suggestions).
Once the samples have been
collected, preserved, and shipped to
the processing centre (e.g., Museo
de Historia Natural, Universidad
Nacional Mayor de San Marcos, or
another national institution) species
must then be sorted, mounted,
labelled, identified, and entered into
the database for the project. The
sampling protocols can be expected
to accumulate about 18,000 arthropod samples in the one year window
available for baseline acquisition at
each of the Camisea gas well sites.
All samples should be processed to
the point of species identification
within 10 months of arrival at the
processing center, and identified to
species within 12 months of arrival
at the center. To meet the 12-month
deadline for results, the sample
processing should be coordinated by
an experienced Processing Manager
assisted by 10 parataxonomists.
Expert identification should be
carried out by contract with the
appropriate systematists to ensure
results within a two-month time
frame. To save time and money,
only the taxa listed below should be
sorted for processing. The residues
from the samples should be

archived for future use.
The processing centre is responsible for the following duties:
* Sorting of samples to target
groups (Orders and Families listed
below).
* Sorting of target groups to
morphospecies.
* Selection of several specimens of each morphospecies
(macrofauna, some microfauna) for
mounting or prepare entire lot
(microfauna) when requested by a
specialist.
* Archiving residues.
* Mounting of selected specimens.
* Labelling of mounted specimens.
* Send specimens to systematists.
* Track loans of specimens.
* Compare identified specimens with sorted morphospecies.
* Enter number of specimens
of each species in database
Target Taxa
The following groups are
proposed for monitoring at the
Camisea well sites. The group name
is followed by the specialist proposed for species identifications,
and the city, country, or institution
of residence.
Aquatics
1 Ephemeroptera (mayflies):
Domingues (Argentina).
2* Trichoptera (caddisflies):
Flint (Washington), Curry
(Toronto).
3* Hydracarina (water mites):
Smith (Ottawa).
4 Coleoptera (water beetles):
Spangler (Washington).
207

5 Dytiscidae (predacious diving
beetles): Roughley (University of
Manitoba).
6* Odonata (dragonflies &
damselflies): Louton (Washington).
Terrestrial
7 Diplopoda (millipedes):
Hoffman (University of Virginia).
8 Chilopoda (centipedes)
Shelly (North Carolina).
Acari (mites)
9* Oribatids: Behan-Pelletier
(Ottawa).
10 Parasitformes: Lindquist
(Ottawa).
11*Araneae (spiders): Silva
(NewYork).
12 Mygalomorphs (tarantulas):
Goloboff.
13 Collembola:
14 Embioptera: Zsumick
(Argentina).
15 Neuroptera (lacewings):
Oswald (Arizona).
16*Orthopteroids (grasshoppers): Otte (Philadelphia); Vickery
(Montreal).
17 Mantodea (mantids):
Strazanac.
Homoptera:
18*Cicadelloidea (leafhoppers):
Lozada (Lima).
19 Fulgorids:
Hemiptera (true bugs):
20 Lygaeidae: Scudder
(Vancouver).
21 Reduviidae:
Coleoptera (beetles):
22*Carabidae (ground beetles):
Will (Cornell)/Erwin (Washington).
23 Anthribidae: Chandler
(Durham NH).
24*Curculionidae (weevils):
Lanteri/Anderson/Howden (Ottawa).
208
25 Mycophilous beetles:
Wheeler (Cornell)/ McHugh (Georgia).
26*Staphylinids: Campbell
(Kentucky), Ash (Kansas).
27 Lampyrids & Phangalids:
Bojorquez (Lima).
28 Buprestids (metallic woodboring beetles): Bellamy.
29 Elateriforma (click beetles):
Fuller (Edmonton).
30*Scarabaeids (scarab
beetles): Valencia (Cusco) &
Howden (Ottawa).
31 Passalids: Marshall (Cornell).
32*Chrysomeloidea (incl.
Cerambycids): Santisteban (Per) +
others.
Diptera (flies):
33 Tabanidae (horse and deer
flies: ?
34*Tachinidae: Wood (Ottawa).
35*Phoridae: Brown (Los
Angeles).
36 Phaerocerids: Marshall
(Guelph).
37 Culicidae (mosquitos): ?
38 Syrphidae (flower flies):
Vockeroth (Ottawa).
39 Asilidae (robber flies):
Fisher.
Hymenoptera (sawflies,
wasps, ants & bees):
40 Symphyta (sawflies): Smith
(Washington), Goulet (Ottawa).
41 Chalcidoidea: Woolley
(Texas A & M), Gibson (Ottawa).
42*Proctotrupoidea,
Scelionidae: Masner (Ottawa).
43 Cynipoidea: Rondqvist
(Sweden).
44*Ichneumonoidea:
Braconidae: Sharkey (Kentucky).
Ichneumonidae: Wahl
(Gainesville).

Chrysidoidea:
45*Bethylids, Dryinids:
Finnamore (Edmonton).
46 Chrysididae: Kimsey
(Davis).
* Vespoidea:
47 Thinnidae: Kimsey (Davis).
48 Tiphiidae: Finnamore
(Edmonton).
49 Vespidae (social wasps):
50 (Eumeninae): Carpenter.
51 Formicidae: Cober (MCZ
Harvard) and L. Alonso (Conservation International).
52 Pompilidae (spider wasps):
53 Mutillidae: Quintero
(Panama).
54*Apoidea: Sphecidae (sand &
twig-nesting wasps): Finnamore
(Edmonton).
Apiformes (bees):
55 Megachilidae (leafcutter
bees): Griswald (Utah).
56*Halictidae: Danforth
(Cornell).
57 Anthophoridae:
Lepidoptera (butterflies &
moths):
58*Rhopalocera (butterflies):
Lamas (Lima), Robbins (Washington).
59 Saturnidae (silk moths):
Lamas (Lima).
60*Noctuidae: Lafontaine
(Ottawa).
61 Sphingidae (hawk moths):
Lamas (Lima).
62 Ctenuchidae: Grados (Lima).
63 Hesperidae (skippers):
Lamas (Lima).
* minimum groups (22) to be
selected for monitoring. Species
may be selected for monitoring for a
number of reasons but a central
suite of species should be also be
selected, from the baseline study, so

as to provide representation of
arthropod size classes, dispersal
abilities, and trophic levels; in the
soil, litter, and canopy communities
existing across major physical and
chemical gradients in the
biodiversity plots at the well sites or
in biodiversity plots in forest communities along a pipeline.
There is a nearly global need to
improve the capacity of third world
countries to document their
biodiversity. The development of
in-country infrastructure for processing and storing specimens
resulting from large-scale
biodiversity projects should be
viewed as a priority by the international community. Processing
specimens in Per accomplishes
several tasks: 1) it transfers control
of specimen deposition to the host
country; 2) it assures that the
opportunity to develop a specimenbased national biodiversity database
is maintained in Per; 3) it develops
the infrastructure and human
resources to ensure a Peruvian
biodiversity capability over the
long-term; and 4) it demands a
long-term commitment by foreign
institutions to the museum selected
as the processing centre.
The first choice for an arthropod processing centre for the
Camisea project is the museum
which logically functions as the
national museum in Per. The
Marcos in Lima is one such institution. Another choice is the Museo
de Historia Natural, Universidad
Nacional San Antonio Abad in
Cusco. Unfortunately, neither of
these institutions has sufficient
209

space to house a processing centre
(11 persons) let alone deal with the
subsequent expansion of its collection.
A short-term solution to this
problem would entail rental of a
facility that could function as a
processing centre and as museum
storage until a long-term resolution
can be reached. An appropriate
long-term solution would require an
upgrading of Peruvian museum
infrastructure to process and house
such collections. That could mean
improved capacity for an existing
institution or establishment of a
new entity. In either case, a facility
with office, processing laboratory,
and substantial storage capacity is
necessary if the Peruvian museum
community is to appreciably participate in biodiversity initiatives on a
long-term basis.
Consideration should be given
to one or a combination of the
following funding options for a
long-term solution to the Peruvian
Biodiversity infrastructure. The
Smithsonian Biodiversity Program
should act as the lead agency in
brokering discussions with the
stakeholders and in development of
a viable proposal for the upgrading
of Peruvian infrastructure. The
potential stakeholders are summarized on the following page:
1. Shell Prospecting and Development (Per) B. V. has already
directed resources to examining and
managing the impact on biodiversity
its natural gas extraction operations
have in Per. There is a commitment to the sustainable management of biodiversity from the
highest levels of the Shell executive. Shell is a large and influential
stakeholder, and positioned as the
210
key player in development of any

proposal to enhance the Peruvian
biodiversity capacity. Members of
the Shell Board of Directors have
visited the San Martin site in the
Camisea Region and have stressed
the importance of the biodiversity
project to Shell Prospecting and
Development (Per) B. V. operations in the Region.
2. The Instituto Nacional de
Recursos Naturales (INRENA) acts
as a clearing house for information
on biodiversity in Per. They may,
in the absence of a suitable museum, be in a position to fund a
management structure and the
curatorial positions to operate a
facility of this nature, given that the
funding for the facility is in place.
3. International development
aid may be of some assistance in
the upgrading of the Peruvian
museum infrastructure and in the
training of staff. Canada has an
International Development Research Centre (IDRC) which has
funded museum enhancement
projects in Indonesia. In light of
Canadas wish to expand the
NAFTA to other Latin American
countries the timing may be appropriate for a submission of behalf of
Per to the IDRC.
Other possible sources of
assistance in Canada include the
Canadian International Development Agency (CIDA). The Canadian
Institute for Petroleum Industry
Development (CIPID) has a proposal pending before CIDA which
aims at strengthening agencies and
institutions involved in regulating
and monitoring natural gas extraction in Per. The Camisea Region is
among the target areas of the
proposal. The CIPID proposal

contains an environmental monitoring component (physical, chemical,
and biotic) that may offer an avenue
for funding. A decision on the
proposal is expected in October
1997 at which point the project will
go to tender. A proposal at that
point could be prepared for the
project environmental consultant,
Swiss Environment and Safety Ltd.
of Calgary. The Biodiversity Science Board of Canada which is
developing biodiversity monitoring
standards for Canada would be
prepared to undertake the preparation of a submission to the environmental consultant.
Several Canadian Government
departments have Matching Investment Initiatives (MII) that could be
of assistance to the project. Money

or in-kind contributions from
Canadian companies can be
matched for research projects that
fall within the departments mandate. Agriculture and Agri-Food
Canada may be interested in an MII
that develops arthropod monitoring
technology. The Nexus Ecological
Research Institute (NERI) is a nonprofit company that is prepared to
act as a broker for Canadian systematists. Contracts for species identifications by Canadian specialists
could be administered by NERI and
used to develop an MII with Agriculture and Agri-Food Canada.
211
212
Amphibians
and Reptiles I:
Biodiversity
Assessment
in the Lower
Urubamba
Region
Robert Reynolds, Thomas
Fritts, Steve Gotte
Biological Resources Division,
US Geological Survey, National
Museum of Natural History
Javier Icochea
Mayor de San Marcos
Guillermo Tello
Justo Vigil 469
Introduction
Several studies documenting

the amphibians and reptiles in
tropical lowland forests of southeastern Per have been published in
recent years. Morales and
McDiarmid (1996) reviewed the
published herpetofaunal surveys
that have been completed in the
department of Madre de Dios since
1980 and suggest that as a result of
these studies, the herpetofauna of
lowland Madre de Dios may be
better known than any other

similar lowland area in tropical
America. Work in the adjacent
areas of the Lower Urubamba
could complement this baseline
knowledge and provide comparative data. The lowland tropical
forested area of the Lower
Urubamba region has not been
studied to the degree that
biodiversity has been and is
being investigated in Madre de
Dios. Morales (1997) recently
published the results of a rapid
assessment of the herpetofauna
in the Rio Camisea area in which
he recorded 24 species of amphibians and seven species of
reptiles.
The SI/MAB studies at the
San Martin-3 and Cashiriari-2
well sites in the Lower
Urubamba focus on faunal and
floral assessments and the establishment of permanent
biodiversity test plots. The next
phase of the project will focus
on monitoring at the plots to
detect diversity, habitat use,
213
Amphibians
and reptiles
comprise an
important
component
in vertebrate faunas
of tropical
forested
regions.
relative abundances, and ecological

changes.
Amphibians and reptiles comprise an important component of
vertebrate faunas in tropical forested regions. They exhibit high
diversity and extreme ecological
specialization and habitat preferences, and they are generally amenable to quantitative sampling. The
primary objective of our participation in this project was to begin
establishing baseline data on the
herpetofauna present at the two
well sites, which can be incorporated into the ultimate task of
creating a multi-taxa monitoring
program to help detect changes in
the ecosystem and assist in management of natural resources in the
area. Selected specimens of amphibians and reptiles encountered
during the survey were sampled and
preserved as vouchers to document
individual species occurrence and to
begin examining and clarifying
distributions and taxonomy of the
herpetofauna in the region.
Methods
Field work at the primary site,

San Martin-3, was in non-flooded
forest habitat dominated by bamboo
(Guadua sarcocarpa), (see Alonso et
al. this volume). The site has extremely steep and irregular topography lacking swamps, major riverine
habitats, and other situations with
the standing water typical of many
tropical lowland sites in Per. The
tall-growing bamboo, which can
have stolons up to several meters
long allowing it to become quite
invasive, is very dense at this site.
Standard nocturnal and diurnal
herpetofaunal surveys and sampling
for amphibian larvae were conducted March 10 to April 9, 1997.
214
In addition to the diurnal and

nocturnal surveys and larval sampling, specimens were sampled at
San Martin-3 using three relatively
experimental quantitative methods:
1) mouse glue boards set on the
forest floor in canopy gaps, 2) from
a 10 X 10-m unfenced plot in which
all ground litter was examined and
displaced, and 3) from an experimental 10 X 10-m fenced plot from
which all above-ground vegetative
matter was removed.
The unfenced 10 X 10-m litter
plots have been commonly used in
New World tropics and elsewhere as
a quantitative measure of forest
floor herpetological communities.
The fenced experimental 10 X 10-m
plot allowed for a complete and
quantitative sample of the amphibians and reptiles present. In this
plot, all vegetation extending
upwards from ground level through
the canopy was excluded along the
perimeter of the plot, and a fence
of greased aluminum flashing with
walls 0.5 m in height was erected at
the plot perimeter to serve as a
barrier to amphibian and reptile
movement into or out of the plot
during the examination period. The
vegetation in the plot was then
inspected and removed down to the
soil substrate, and all amphibians
and reptiles encountered were
recorded and sampled. Other
studies have variably controlled
immigration and emigration of
specimens from plots (i.e., using
perimeter barriers, canopy separation, or isolation of vertical tree
trunks), but none to our knowledge
have addressed all three at the same
time as we attempted in this study.
Additional survey work was
done at the Cashiriari-2 well site on
the evening of March 26. This area
is primary lowland wet forest
without the extensive bamboo

stands that are present at San
Martin-3. The accommodations for
scientific personnel at Cashiriari-2
were not completed before the end
of our scheduled field time, preventing further work at that site
(but see Icochea and Mitchell, this
volume). As a result, we were only
able to survey amphibians and
reptiles for the one evening, and we
have minimal data from the site.
Results and
Discussion
In contrast to more typical

tropical lowland sites where bamboo thickets are only a fraction of
the total habitat, nearly all of our
efforts at San Martin-3 were conducted inor in close proximity
toprofuse bamboo stands. This
habitat feature comprised a significant visual complexity and furnished a myriad of potential perch
sites, which undoubtedly affected
our sampling results. Exclusive of
glue board or removal plot sampling, most surveys of amphibians
and reptiles depend on visual
searches for animals as well as
detecting frog vocalizations. The
density and visual complexity of the
bamboo forest limited our ability to
see even the calling frogs known to
be present and undoubtedly reduced
sightings of terrestrial and arboreal
reptiles.
The timing of the field work
coincided with the late part of the
wet season and the initial transition
period into the dry season. The
well-drained soils and steep terrain
at San Martin-3 were entirely
devoid of standing above-surface
ground water suitable for frog
breeding and subsequent larval
development. However, a myriad of
small water-filled sites were available in the form of broken off or

otherwise opened bamboo internodes. These water-filled bamboo
sites included both standing and
fallen stalks, stalks with openings
gnawed by arboreal rodents foraging
for insect larvae, and stalks that
had been opened during camp
development, trail maintenance,
and natural tree falls. No evidence
of frog reproduction (eggs or
larvae) was found during examination of more than 200 standing or
fallen water-filled bamboo internodes, although the majority of these
aquatic sites did contain insect
(primarily mosquito) larvae and
other organic material.
A single Phyllomedusa egg mass
was found on the underside of a
leaf overhanging a small stream in
the biodiversity plot #2. Despite
extensive sampling in and along the
streams, this was the only evidence
of anuran reproduction found.
Approximately 100 dip net samples
were taken at two second-order
streams and five seeps feeding into
the streams near San Martin-3. No
larvae or other indications of
amphibian reproduction were found
in the stream samples or observed
during the sampling process. Because of the steep topography, most
streams at the site were subject to
major rising of water levels (up to 4
m during our visit) and dramatic
flooding during heavy rains that
would easily carry away eggs and
larvae from the stream channels.
The most common anurans at
San Martin-3 were members of the
genus Eleutherodactylus. Frogs of this
genus develop directly from an egg
to the adult stage without a freeliving aquatic tadpole stage and are
thus not necessarily dependent on
standing water for development.
The most
common
anurans at
San
Martin-3
were
members
of the
genus
Eleutherodactylus.
215
Members of the typically abundant

tropical frog family Hylidae, although present, were remarkably
uncommon at San Martin-3. In
contrast to the Eleutherodactylus,
most hylids have a free-living
aquatic larval stage and are dependent on standing or flowing water
for development. We also encountered Hemiphractus probocideus and H.
scutatus, hylids in which the unpigmented eggs develop on the
females back; four species of
Dendrobatids that have the ability
to transport larvae over land after
the eggs hatch; and Osteocephalus

leprieurii, a hylid known to deposit
eggs in cavities such as bamboo or
other isolated water receptacles.
A total of 504 specimens were
sampled, documenting more than 80
from the San Martin-3 and
Cashiriari-2 well sites (Table 1 and
Appendix 1). We recorded 16
genera in seven families of amphibians, and 31 genera in 10 families
of reptiles. There were substantially
more specimens per taxa of amphibians, but more families and
Table 1. Number of genera, species and specimens of the amphibians and reptiles sampled at San Martin-3 and Cashiriari-2 in March
and April, 1997, during the biodiversity assessment in the Lower
Urubamba region
Species
Specimens
CLASS AMPHIBIA
16
ORDER ANURA
14
Family Buonidae
1
Family Dendrobatidae
2
Family Hylidae
5
Family Leptodactylidae
5
Family Microhylidae
1
unidentified eggs/larvae
ORDER CAUDATA
Family Plethodontidae
1
ORDER GYMNOPHIONA
Family Caediliidae
1
40+
38+
2
4
11
20+
1
2
410
394
52
28
89
223
5
CLASS REPTILIA
Suborder Sauria
Family Gekkonidae
Family Gymnophthalmidae
Family Polychrotidae
Family Scincidae
Family Teiidae
Family Tropiduridae
Suborder Serpentes
Family Boidae
Family Colubridae
Family Elapidae
Family Viperidae
Genera
216
31
12
2
5
2
1
1
1
19
2
14
1
2
40
15
2
5
5
1
1
1
25
2
20
1
2
94
54
3
11
14
20
3
3
40
2
34
2
2
genera of reptiles even though

reptiles constituted only 18% of the
total number of specimens collected (Table 1). The species designations for the Eleutherodactylus are
tentative and some will undoubtedly change as specimen identifications are further refined through
additional comparisons with museum collections and the systematic
literature.
In addition to the specimens
sampled, we also recorded four
additional taxa at San Martin-3 that
were not sampled. We captured and
released one caiman (Paleosuchus
palpebrosus) from a small creek east
of camp and positively identified
the call of the tree frog Hyla boans
one evening from the large
quebrada east of camp. That vocalization constituted our only record
for this species. A single Plica plica
was observed out of reach high on
a tree trunk in the vicinity of our
experimental removal plot. Finally,
the botanists reported that they
encountered a tortoise (Geochelone
denticulata) while mapping trees in
the biodiversity plot #1.
The difference in numbers of
amphibians encountered using
traditional plot methodology (i.e.,
unbounded plots in which litter is
moved toward the outside perimeter) versus the barrier plot (where
all vegetation was examined and
removed after the plot was isolated
using a 0.5-m aluminum fence and
canopy separation achieved by
pruning) was strikingly notable. The
traditional plot produced three
anurans (one dendrobatid and two
Eleutherodactylus), whereas the
barrier plot produced 29 anurans
(including five microhylids not
encountered during any other
sampling effort) and one snake
(Corallus hortulanus). In addition to
the five Syncope antenori, the amphibians captured in the litter plot
included Eleutherodactylus and Bufo.
Additional replicates of this methodology are needed to properly
evaluate its utility in determining
better estimates of absolute abundance relative to the unbounded
plots (which are less labor and time
intensive), but this initial attempt
produced extremely encouraging
results.
Study of lizards at San
Martin-3 was greatly facilitated by
the use of mouse glue boards
placed on the ground as well as on
low branches and bamboo stems in
areas of broken sunlight and shade.
The glue boards were successful in
capturing four gymnophthalmid
lizards (Arthrosaura, Cercosaura,
Iphisa, and Prionodactylus) that were
either never or only rarely seen
during visual surveys. Glue boards
also facilitated capture of some
larger lizards (Kentropyx pelviceps,
Anolis punctatus, Mabuya bistriata) in
densely vege-tated areas where
hand capture was hindered and less
successful. The glue boards were
also used on tree trunks and
branches to monitor potential
movement of animals out of the
fenced plot during the removal
experiment.
Other Important
Activities
We took the opportunity to

work with our samples for a few
days at the Museo de Historia
Natural at the Universidad Nacional
Mayor de San Marcos in Lima.
While there, we spent considerable
time assessing and discussing
herpetology collection management
issues, data standards, and electronic databases with the curator of
217
We were
successful
in initiating
the surveys
and
sampling
necessary
for establishing the
baseline
data on the
herpetofauna
present at
San
Martin-3.
218
the herpetology department. As a

result of these discussions, we have
already begun providing information and guidelines for database
structure and management to our
Peruvian counterparts based on our
collection management activities at
the U.S. National Museum. Our
recent involvement and commitment to the collection management
capabilities at the Museo de
Historia Natural is a first step
toward a cooperative venture to
help improve the status of the
herpetology collection there, and
hence, the research potential of the
specimens. Half of the voucher
specimens collected during our field
work will be deposited at the
Museo de Historia Natural, as have
collections from previous work by
Smithsonian Institution and other
foreign investigators working in
Per. Any effort, therefore, to help
enhance the collection capabilities
of the museum for conservation
and curation is in the best interest
of the specimens, the research they
support, and their ultimate use by
national and international scientists
in the future.
We were successful in initiating
the surveys and sampling necessary
for establishing the baseline data on
the herpetofauna present at San
Martin-3, but because of circumstances beyond our control, comparable data are lacking for Cashiriari2 (but see Icochea and Mitchell,
this volume). Additional herpetological survey work spanning the
wet and dry seasons is necessary at
both sites to build on the inventory
started during this initial effort. The
potential exists for much amphibian
reproduction to be restricted to
periods of low or intermediate
levels of rainfall when pools in
streams would be more suitable for
reproduction by hylid and

microhylid frog species. Tests of
new sampling strategies and techniques were encouraging, and our
Peruvian counterparts expressed
interest in future amplification of
quantitative sampling efforts at
additional sites in Per.
References
Morales, V. R., and R. W.

McDiarmid. 1996. Annotated
checklist of the amphibians and
reptiles of Pakitza, Manu National
Park Reserve Zone, with comments on the herpetofauna of
Madre de Dios, Per. Pages 503522 in Manu: The Biodiversity of
Southeastern Per. (D. E. Wilson
and A. Sandoval, eds.) Smithsonian Institution Press,
Washington, DC.
Morales, V. R. 1997. La
herpetofauna en la zona del Rio
Camisea, Cusco, Per: Una
evaluacion rapida. Pages 76-79 in
of the Lower Urubamba, Per.
(S. Udvardy and A. Sandoval, eds.)
Biodiversity Programs,
Washington, DC.
Appendix 1. Species list of the amphibians and reptiles

sampled at San Martin-3 and Cashiriari-2 in March and April,
1997, during the biodiversity assessment in the Lower
Urubamba region, Cusco, Per.
CLASS AMPHIBIA
ORDER ANURA
Family Bufonidae
Bufo marinus
Bufo cf. typhonius
Dendrobates biolat
Dendrobates cf. ventrimaculatus
Epipedobates femoralis
Epipedobates macero
Family Hylidae
Hemiphractus proboscideus
Hemiphractus scutatus
Hyla geographica
Hyla lanciformis
Hyla marmorata
Hyla parviceps
Osteocephalus leprieurii
Phyllomedusa cf. tarsius
Phyllomedusa tomopterna
Phyllomedusa vaillanti
Scinax garbei
Adenomera cf. andreae
Eleutherodactylus cf. altamazonicus
Eleutherodactylus cf. carvaloi
Eleutherodactylus cf. conspicillatus
Eleutherodactylus cf. croceoinguinis
Eleutherodactylus cf. diadematus
Eleutherodactylus cf. fenestratus
Eleutherodactylus cf. martae
Eleutherodactylus cf. ockendeni
Eleutherodactylus Peruvianus
Eleutherodactylus sulcatus
Eleutherodactylus cf. toftae
Eleutherodactylus cf. ventrimarmoratus
Ischnocnema quixensis
Leptodactylus leptodactyloides
Leptodactylus pentadactylus
Leptodactylus petersii
Leptodactylus rhodonotus
Leptodactylus stenodema
Lithodytes lineatus
Family Microhylidae
Syncope antenori
ORDER CAUDATA
Bolitoglossa altamazonica
ORDER GYMNOPHIONA
Family Caeciliidae
Caecilia tentaculata
San Martin-3
Cashiriari-2
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
219
Appendix 1. Species list of the amphibians and reptiles (Cont.).

San Martin-3
CLASS REPTILIA
ORDER SQUAMATA
Suborder Sauria
Family Gekkonidae
Pseudogonatodes guianensis
Thecadactylus rapicauda
Alopoglossus cf. andeanus
Cercosaura ocellata
Iphisa elegans
Prionodactylus argulus
Neusticurus ecpleopus
Anolis fuscoauratus
Anolis punctatus
Anolis trachyderma
Anolis transversalis
Polychrus liogaster
Family Scincidae
Mabuya bistriata
Family Teiidae
Kentropyx pelviceps
Family Tropiduridae
Plica umbra
ORDER SQUAMATA
Suborder Serpentes
Family Boidae
Corallus hortulanus
Epicrates cenchria
Family Colubridae
Atractus elaps
Atractus cf. badius
Atractus major
Chironius cf. monticola
Clelia clelia
Dipsas catesbyi
Dipsas indica
Imantodes cenchoa
Imantodes lentiferus
Leptodeira annulata
Liophis cobellus
Liophis reginae
Oxyrhopus doliatus
Oxyrhopus petola
Pseustes sulphureus
Rhadinaea brevirostris
Tantilla melanocephala
Tripanurgos compressus
Xenodon rabdocephalus
Xenopholis scalaris
Family Elapidae
Leptomicrurus narducci
220
Cashiriari-2
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Appendix 1. Species list of the amphibians and reptiles (Cont.).

San Martin-3
CLASS REPTILIA
ORDER SQUAMATA
Suborder Serpentes
Family Viperidae
Bothrops cf. brazili
Lachesis muta
LARVAL SAMPLES
Epipedobates macero
Phyllomedusa sp.
Phyllomedusa cf. tarsius
Cashiriari-2
x
x
x
x
x
221
222
Amphibians
and Reptiles II:
Biodiversity
Assessment in
the Lower
Urubamba
Region
Javier Icochea
Mayor de San Marcos
Joseph Mitchell
Department of Biology,
Univerisity of Richmond
Introduction
While the study of biological

diversity is receiving increased
attention, especially in tropical
areas, the study of amphibians in
particular has been accelerated,
most likely because of the extinction of some amphibian species
even in protected areas. Habitat
loss caused by destruction of the
tropical rain forest, thinning of the
ozone layer that protects organisms
from the radiation of ultraviolet
rays, and greater pollution of the air
and water are considered important
factors in the widespread decrease
of amphibian populations (Blastin
and Wake 1995). The extent of this

phenomenon is a matter of current
debate, which makes it all the more
important to establish amphibian
monitoring programs so that information is available to answer the
questions concerning amphibian
declines.
As noted by Reynolds et al (this
volume) amphibians are extremely
helpful to the broader study of
ecosystem functioning. Their life
cycles involve aquatic and terrestrial habitats, their permeable skins
and mostly cutaneous respiration
make them sensitive to pollutants
while exchanging materials with
their environments, their nutrition
level is high, and they constitute an
appreciable portion of the biomass
in any given ecosystem (Wake
1991).
Few biodiversity studies have
been conducted in the Lower
Urubamba region, considered by
Rodriguez (1996) as a priority area
for the conservation of biodiversity
in Per. The only published records
of amphibians and reptiles24
amphibian and seven reptile
223
speciesfor the Camisea River

Basin were made by Morales
(1996), who participated in the
initial evaluation conducted by SI/
MAB for this project (see also
Reynolds et al. in this volume for
additional information on amphibian and reptile species in the area).
Methods
The second assessment of

amphibians and reptiles was conducted May 2-15, 1997. We spent
five days at the San Martin-3 well
site at the beginning of the field trip
while accommodations were prepared at the Cashiriari-2 well site.
San Martin-3 is a lowland forest
dominated by bamboo (Guadua
sarcocarpa), while Cashiriari-2 is
characterized by large emergent
trees typical of tropical rainforest.
We took observation and
sampling trips at both study sites,
walking on existing trails, in
streams, and through opened areas
where the well sites are found. We
also sampled inside the biodiversity
plots established by participants in
this study (see Alonso et al. this
volume). Sampling methods are
described as follows:
Visual encounter. We conducted night searches using existing
trails and streams as transects and
carefully examining both sides of
the transects from the ground to the
tops of trees. Once an animal was
spotted, it was captured by hand.
Locating by song. When we
heard a frog call, we directed our
attention toward the sound to
locate the animal. We then either
identified the animal by its call or
captured the individual once it was
located. The study was completed
at the end of the rainy season, so
numerous species were not repro224
ducing and their singing was at a

minimum.
Sticky (glue) traps. We placed
these traps in opened areas of the
forest on tree branches, fallen tree
trunks, bamboo shoots, and the
ground. Since Reynolds et al. (this
volume) sampled a goodly number
of species in San Martin-3 in March
and April, our work consisted of
trapping the organisms, measuring
and weighing them in the laboratory, and releasing them at the same
place they were sampled. Some
specimens were preserved in 10%
formaldehyde as museum specimens. We took photographs of all
live specimens trapped.
We used the collection of
amphibians and reptiles at the
Museo de Historia Natural de San
Marcos (MUSM) to determine the
specimens sampled, assisted by the
following literature to aid with
identification: Avila-Pires (1995),
Brame and Wake (1963), Duellman
(1978), Rodriguez (1994),
Rodriguez and Duellman (1994),
Peters and Donoso-Barros (1970),
Peters and Orejas- Miranda (1970),
Vanzolini (1986), and Vitt and De
la Torre (1996).
Results and
Discussion
We registered 23 species of
amphibians and 19 species of
reptiles within the following groups:
salamanders (one), caecilia (one),
frogs (21), lizards (nine), and snakes
(10) (Appendix 1). Added to our
visual and auditory records, we
sampled 94 individuals within the
following groups: salamanders
(one), caecilia (one), frogs (50),
larvae series (two), lizards (27), and
snakes (13). The specimens
sampled have been numbered
within the series USNM-FH

(211051-211130). The specimens
sampled during May by well construction site workers have been
numbered within the series JIM
(14478-14488). Specimen number
14489 is a snake (Pseustes
sulphureus) from Cashiriari-3; it was
not included in the list because it
was not registered in the localities
before. The material associated with
the data in Table 1 (USNM-FH) has
been deposited at Departamento de
Herpetologa del Museo de Historia
Natural de la Universidad Nacional
Mayor de San Marcos (MUSM).
Visual encounters during
nocturnal transects helped us locate
a larger number of individuals.
Several species of lizards were only
seen and captured with the stiky
traps (number of individuals in
parenthesis): Gonatodes hasemanni
(one), Thecadactylus rapicauda (one),
Prionodactylus argulus (two), Anolis
fuscoauratus (three), Anolis
trachyderma (two), Mabuya bistriata
(five), and Ameiva ameiva (13). We
also sampled a tree snake, Imantodes
cenchoa, with the trap and, unfortunately, several birds species, so the
use of this trap should be evaluated. A lizard (Pseudogonatodes
guianensis) within the gekkonidae
that lives on the leaf litter was
caught in a pitfall trap set by the
entomologists. The caecilia (Caecilia
cf. tentaculata) was found by construction workers as they were
removing top soil.
Several of the amphibian
species reported in this study have
been found at other lowland tropical sites in Per; 91% of the species
have been found in Cocha Cashu
and 78% in Pakitza. The reptile
fauna also overlaps with the other
two sites72% and 67%, respectively. We can not make fair com-
parisons of the species richness

between San Martin-3 and
Cashiriari-2 because the sampling
efforts were more extensive at San
Martin-3. We found additional
species to the ones reported by
Reynolds et al. (this volume):
Atelopus spumarius and Osteocephalus
taurinus, amphibians with large
distribution in the Amazon Basin,
Gonatodes hasemanni (gekonidae) also
registered at Cocha Cashu and
Pakitza, and the snakes Drymobius
rhombifer and Liophis epinephelus
fraseri which have not been registered for the Manu River.
Several of the species found
are not abundant at other sites.
Caecilia cf. tentaculata has been
reported for Per, but the Department was not given (Rodriguez et
al. 1993), and it has not been
reported at Cocha Cashu or Pakitza.
In general, caecilias are dificult to
find because they live underground.
It is also possible that this species
has been confused with Siphonops
annulatus, a species with larger
distribution.
We can not make estimates of
abundance with our current sampling effort. We can detect, however, some species that seem to be
more common; e.g., Bufo marinus (a
species of toad frequently found at
night near camps), Bufo gr. typhonius
(a species of toad found living on
the leaf litter on the forest floor),
Ameiva ameiva (a species of lizard
that is very active during the day
and that prefers open areas),
Mabuya bistriata (a species of lizard
that is very active and was frequently captured on the stiky traps
placed on the bamboo at San
Martin-3 but was not found at
Cashiriari-2). Several species of
frogs were also relatively common,
including Epipedobates macero (frogs
Visual
encounters
during
nocturnal
transects
helped us
locate a
larger
number of
individuals.
225
active during the day but easier to

trap at night while sleeping on the
foliage), Eleutherodactylus toftae
(small frogs frequently found at
night on foliage), and Leptodactylus
rhodonotus (found near the camps,
especially at Cashiriari-2). The
relative abundance of the species
of the genus Eleutherodactylus spp.
indicates that the forest has received minimum disturbance. The
relative low abundance of the tree
frogs of the Family Hailed is due to
the fact that our sampling was
conducted close to 500 m above sea
level. At this altitude, the most
appropriate reproductive sites for
tree frogs are not common.
We recommend further studies,
particularly a long-term monitoring
program to detect changes in the
population dynamics of the species.
The traditional use of these species
by the indigenous communities
should help researchers better
understand the natural history of
many species. It is also important to
monitor the aquatic immature life
stages of the amphibians, especially
in the areas where the treated water
from the drilling exploration is
delivered for storage.
We thank Dra. Nelly Carrillo de
Espinoza for checking the snake
identifications.
References
Avila-Pires, T. C. S. 1995. Lizards

of Brazilian Amazonia (Reptilia:
Squamata). Zool. Verh. Leiden 229:
1-706.
Blastin, A. R., and D. B. Wake.
1995. The puzzle of declining
amphibian populations. Scientific
American 272: 56-61.
Brame, A. H., Jr., and D. B. Wake.
1963. The salamanders of South
226
America. Contributions in Science 69:

1-72.
Carrillo de Espinoza, N., and J.
Icochea. 1995. Lista taxonomica
preliminar de los Reptiles vivientes
del Per. Publicaciones del Museo de
Historia Natural U.N.M.S.M.
(A)49: 1-27.
Duellman, W. E. 1978. The Biology
of an Equatorial Herpetofauna in
Amazonian Ecuador. Museum of
Natural History, Kansas University, Miscellaneous Publication 65:
1-352.
Morales, V. 1996. La Herpetofauna
en la Zona del Rio Camisea,
Cusco, Per: Una Evaluacion
Rapida. Pages 76-79, 167-168 in
Actas del Taller Sobre Diversidad
Biologica y Cultural del Bajo
Urubamba, Per. 2-4 de
Septiembre de 1996. (S. Udvardy
and A. Sandoval , eds.)
Washington, DC.
Morales, V., and R. McDiarmid.
1996. Annotated checklist of the
amphibians and reptiles of
Pakitza, Manu National Park
Reserve Zone with comments on
the herpetofauna of Madre de
Dios, Per. In Manu: The
Editorial Horizonte, Lima.
Peters, J. A., and R. Donso-Barros.
1970. Catalogue of the Neotropical Squamata: Part II. Lizards and
Amphisbaenians. Bulletin of the
United States National Museum 297:
1-293.
Peters, J. A., and Orejas-Miranda.
1970. Catalogue of the Neotropical Squamata: Part I. Snakes.
Bulletin of the United States National
Museum 297: 1-347.
Rodriguez, L. 1994. A new species
of the Eleutherodactylus conspicillatus
group (Leptodactylidae) from Per,
with comments on this call. Alytes

12(2): 49-63.
Rodriguez, L. 1996. Areas
Prioritarias y vacios de
informacion para la Conservacion
de la Biodiversidad en el Per.
Pages 29-34 in Actas del Taller
Sobre Diversidad Biologica y
Cultural del Bajo Urubamba, Per.
2-4 de Septiembre de 1996.
(S. Udvardy and A. Sandoval, eds.)
Washington, DC.
Rodriguez, L., and J. E. Cadle.
1990. A preliminary overview of
the herpetofauna of Cocha Cashu,
Manu National Park, Per. Pages
410-425 in Four Neotropical
Rainforest. (A. H. Gentry, ed.)
Rodriguez, L., J. H. Cordova, and J.
Icochea. 1993. Lista preliminar de
los Anfibios del Per. Publicaciones
del Museo de Historia Natural
U.N.M.S.M. (A) 45: 1-22.
Rodriguez, L., and W. Duellman.
1994. Guide to the Frogs of the
Iquitos Region, Amazonian Per.
Special Publication No 22.The
University of Kansas Natural
History Museum, Topeka.
Scott, N. J., Jr. 1994. Complete

species inventories. Pages 78-84 in
Measuring and Monitoring Biological Diversity: Standard Methods
for Amphibians. (W. R. Heyer, M.
A. Donnelly, R. W. McDiarmid, L.
A. C. Hayek, and M. S. Foster,
eds.) Smithsonian Institution Press,
Washington, DC.
Vanzolini, P. E. 1986. Addenda and
corrigenda to the Catalogue of
Neotropical Squamata. Smithsonian
Herpetological Information Service 70:
1-25.
Vitt, L. J., and S. de la Torre. 1996.
Guia para la investigacion de las
lagartijas de Cuyabeno. Museo de
Zoologia P.U.C.E., Monografia 1:
1-165.
Wake, D. B. 1991. Declining amphibian populations - a global
phenomenon? Findings and
recomendations. Alytes 9(2):
33-42.
227
Appendix 1. Amphibians and reptiles in the Lower Urubamba region (listed

specimens sampled in May 1997 at the San Martin-3 and Cashiriari-2 well
sites; number of specimens sampled at each site and the strata where each
of the species was foundarboreal (A), terrestrial (T), or fussorial (F); note
that (O) = observed, (V) = identified by vocalizations , and X = specimen
was detected at the site but not sampled).
San Martin-3
CLASS AMPHIBIA
ORDER ANURA
Family Bufonidae
Atelopus spumarius
Bufo marinus
Bufo gr. typhonius
Epipedobates femoralis
Epipedobates macero
Family Hylidae
Hyla geographica
Hyla marmorata
Osteocephalus leprieurii
Osteocephalus taurinus
Phrynohyas venulosa
Phyllomedusa vaillanti
Eleutherodactylus cf. conspicillatus
Eleutherodactylus diadematus
Eleutherodactylus cf. diadematus
Eleutherodactylus fenestratus
Eleutherodactylus ockendeni
Eleutherodactylus cf. Peruvianus
Eleutherodactylus toftae
Eleutherodactylus cf. ventrimarmoratus
Ischnocnema quixensis
Leptodactylus pentadactylus
Leptodactylus rhodonotus
ORDER CAUDATA
Bolitoglossa cf. altamazonica
228
Cashiriari-2
Strata
Record
1
3
1
5
3
T
T
T
S
S, O, V
S, O, V
x
x
T
T
O, V
S, O
A,T
A
A
A
A
A
A
A
S
S, O, V
V
S
S
S
S, O
O
A,T
A,T
A,T
T
A,T
A,T
A,T
A,T
T
T
T
T
S
S
S, O
S
S
S, O
S, O
S
S
O
S, V
S, O, V
S, O
1
4
x
x
1
3
2
2
1
2
1
2
1
x
1
x
1
1
1
1
1
2
1
1
4
Appendix 1. Amphibians and reptiles in the Lower Urubamba region (Cont.).

San Martin-3
ORDER GYMNOPHIONA
Family Caecilidae
Caecilia cf. tentaculata
CLASS REPTILA
ORDER SQUAMATA
Suborder Sauria
Family Gekkonidae
Gonatodes hasemanni
Pseudogonatodes guianensis
Thecadactylus rapicauda
Prionodactylus argulus
Anolis fuscoauratus
Anolis trachyderma
Family Scincidae
Mabuya bistriata
Family Teiidae
Ameiva ameiva
Family Tropiduridae
Plica umbra
ORDER SQUAMATA
Suborder Serpentes
Family Boidae
Epicrates cenchria
Family Colubridae
Atractus elaps
Drymobius rhombifer
Imantodes cenchoa
Leptodeira annulata
Liophis epinephelus fraseri
Liophis reginae
Family Viperidae
Lachesis muta
Cashiriari-2
Strata
Record
T
T
A
S
S
S
A, T
A
A
S, O
S, O
A, T
S, O
S, O
T
T
A
A
T
A, T
S
S
S
S
S
S
T
T
S
S
1
1
2
2
1
2
1
2
1
3
1
9
1
1
1
1
1
2
1
1
1
229
230
Genetic
Structure of
Amphibians
and
Reptiles:
Biodiversity
Assessment
in the Lower
Urubamba
Region
Jesus Cordova and
Cesar Aguilar
Mayor de San Marcos
Introduction
Biological assessments in
tropical areas have recently begun
incorporating studies of the genetic
diversity of several groups of
animals. Such studies help identify
species in an area, detect variability
within populations (another important aspect of biodiversity), and
determine changes in genetic
structure once a baseline is obtained. We participated in the
biological assessment in the Lower
Urubamba region of Per to gather

baseline data on several species of
amphibians to compare possible
changes in time. We are currently
analyzing the samples; thus this
paper presents a summary of the
information gathered to date.
Methods
We visited the well sites at San

Martin-3 and Cashiriari-2 from June
4 to June 20, 1997. Our samples
were obtained with the aid of
numerous collaborators working on
the overall biological assessment.
We used standardized procedures to
sample amphibian and reptile
specimens, including visual and
vocalization/calling surveys, sticky
traps, and tadpole netting. We
conducted daily diurnal and nocturnal surveys along existing trails, in
the biodiversity plots at the well
sites, and at the camp. Once we had
the specimen, we obtained tissue
samples from several sources,
including bone marrow, testicles,
blood, and livers. We also sampled
material from the paratyroid glands
within the Bufonidae (Table 1).
231
Careful attention was paid to

recording the sampling sites so that
we can correlate site collection and
animal status as we finish the
analysis.
We made permanent samples
(slides) in situ from the bone marrow, blood, and testicle tissues. We
are in the process of examining
those samples for micronucleus and
chromosomic data. To eliminate
bias in our data recording, the
person analyzing the sample
through the microscope does not
know where the sample is coming
from or the identity of the species
under study. Nearly all species were
photographed alive, and when
necessary, we photographed the
habitat where they were found.
232
We sampled 84 specimens of
amphibians and reptiles and took 50
blood samples (for the micronucleus
test) of 21 species. To obtain
reliable results, we are processing
3,000 nuclei per individual. To test
for chromosomic alterations, we
obtained 51 samples from 18 species and are analyzing at least 30
nuclei in metaphase by specimen.
We expect that this will generate at
least 102 slides, two per individual.
It takes about 15 minutes to analyze
each slide. Our findings will be
submitted to the final report concerning the overall project.
Appendix 1. List of amphibians and reptiles sampled at San Martin-3 and Cashiriari-2 for the
study of genetic diversity of these species in the Lower Urubamba region.
Number
20271
20272
20273
20274
20275
20276
20277
20278
20279
20280
20281
20282
20283
20284
20285
20286
20287
20288
20289
20290
20291
20292
20293
20294
20295
20296
20297
20298
20299
20300
20301
20302
20303
20304
20305
20306
20307
20308
20309
Species
Epipedobates macero
Bufo typhoniu
Leptodactylidae 1
Leptodactylus cf. rhodonotus
Bufo marinus
Atelopus spumarius
Atelopus spumarius
Alpoglossus sp. 1
Epipedobates cf. femoralis
Bufo typhonius
Bufo marinus
Bufo marinus
Bufo marinus
Atractus elaps
Cercosaura ocellata
Hyla lanciformis
Eleutherodactylus sp. 1
Epipedobates macero
Bolitoglossa altamazonica
Eleutherodactylus sp. 1
Bufo marinus
Bufo marinus
Bufo marinus
Bufo marinus
Bufo marinus
Bufo marinus
Ameiva ameiva
Atractus major
Status
adult
immature
immature
juvenile
adult
juvenile
juvenile
juvenile
adult
adult
adult
adult
adult
adult
adult
adult
juvenile
adult
adult
adult
juvenile
adult
adult
adult
adult
adult
adult
adult
adult
adult
adult
adult
adult
adult
adult
adult
adult
juvenile
adult
Sample Bone Testicles Blood Skin Liver Glands

x
*
*
x
x
x
x
x
x
x
x
x
x
x
x
x
x x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
233
Appendix 1. Amphibians and reptiles sampled at San Martin-3 and Cashiriari-2 (Cont.).
Number
20310
20311
20312
20313
20314
20315
20316
20317
20318
20319
20320
20321
20322
20323
20324
20325
20326
20327
20328
20329
20330
20331
20332
20333
20334
20335
20336
20337
20338
20339
20340
20341
20342
20343
20344
20345
20346
20347
20348
20349
20350
20351
20352
20353
20354
234
Species
Anolis trachiderma
Osteocephalus sp. 1
Dendrophidium dendrophis
Bufo typhonius
Epipedobates macero
Bufo typhonius
Eleutherodactylus diadematus
Atractus elaps
Amphisbaena fuliginosa
Bufo typhonius
Hylidae 1
Bufo typhonius
Anolis trachiderma
Bufo marinus
Bufo typhonius
Hylidae 1
Hylidae 1
Anura
Bufo typhonius
Anolis trachiderma
Epipedobates macero
Bufo marinus
Micrurus sp.
Ameiva ameiva
Ameiva ameiva
Atelopus spumarius
Atractus elaps
Bufo marinus
Bufo typhonius **
Bufo typhonius **
Bufo marinus
Bufo typhonius
Imantodes sp.
Status
Sample Bone Testicles Blood Skin Liver Glands
adult
adult
adult
adult
adult
adult
adult
juvenile
adult
adult
adult
adult
adult
adult
juvenile
immature
adult
adult
adult
adult
adult
immature
adult
immature
immature
adult
immature
adult
adult
adult
adult
adult
juvenile
juvenile
juvenile
juvenile
adult
adult
adult
adult
adult
adult
adult
adult
adult
x
x
x
x
x
x
?
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Long-term
Monitoring of
Amphibians
in the Lower
Urubamba
Region
Joseph C. Mitchell
University of Richmond
Introduction
Amphibians (frogs, salamanders, caecilians) are a diverse

group of predatory animals that
occur on all continents but reach
their greatest abundance in tropical
regions. Most have complex life
cycles, an aquatic larval stage, and
a terrestrial adult stage that is
unique among vertebrates. Each
individual is thus exposed to
aquatic and terrestrial (and atmospheric) perturbations during its life
time. This life history, combined
with a moist, sensitive skin used for
respiration (Duellman and Trueb
1986) and lack of long-distance
migratory abilities (Sinsch 1990),
renders individuals and populations
susceptible to a variety of environmental modifications at several
geographic and ecological scales.
They are usually the first verte-
brates to respond to environmental

change.
Amphibians world wide have
responded to large-scale perturbations by undergoing population
decline, range reduction, and extinction (Blaustein and Wake 1990,
Phillips 1990, Wyman 1990, Wake
1991, Blaustein et al. 1994c).
Causes of decline include habitat
loss and degradation (Lannoo et al.
1994, Delis et al. 1996), acid
precipitation (Wyman 1988,
Sadinski and Dunson 1992), UV-B
radiation (Blaustein et al. 1994a),
pathogens (Blaustein et al. 1994b,
Anderson 1995), introduction of
exotic species (Schwalbe and Rosen
1988, Fisher and Shaffer 1996),
harvesting by humans (Hayes and
Jennings 1986), and natural population fluctuations (Pechmann et al.
1991). The most common causes of
range reduction, population decline,
and alteration of community structure are habitat loss and degradation. The scientific community has
responded to the crisis by forming
the Declining Amphibians Populations Task Force (DAPTF) administered by the World Conservation
235
Monitoring
of amphibians in the
Lower
Urubamba
will help
determine
whether
changes in
species
distribution
have taken
place because of
habitat
alterations.
236
Union (IUCN). The DAPTF focuses

its efforts on determining the status
of amphibians in the regions of the
world and developing standardized
techniques for long-term monitoring. A primary result of the latter
effort is the publication of a book
on techniques by Heyer et al.
(1994).
The Lower
Urubamba
Amphibian
Monitoring Strategy
The primary perturbations
likely to occur at Shell Prospecting
extraction sites in the Lower
Urubamba region that may affect
amphibians are: 1) direct habitat
loss at each construction site and
on the roads cut through primary
forest in the 1980s, 2) changes in
the quality of rain-forest habitat
adjacent to each site (the edge
effect), and 3) siltation of streams
in the watershed from soil washing
from the construction sites. Since
most species are adapted to forest
conditions, the number of species
that occurred originally at each
construction site has already declined. However, at least two
species survive in the open areas
the marine toad and a leptodactylid
frog. Once the rainforest is opened
by a disturbance such as clearing for
a well site, the trees and other
vegetation that occurred inside the
forest are exposed to higher levels
of sunlight, temperature, humidity,
and wind. The composition of the
vegetation at the forest edge
changes in response to the changes
in environmental conditions. Because many rainforest amphibians
are adapted to interior forest condi-
tions, few will tolerate open areas

or forest edges. The few that can
tolerate such conditions live in the
ground and breed in open puddles
or are drawn from the edge vegetation by the lights that attract insects. Siltation does not appear to
be a current problem in the small
streams visited at San Martin-3 and
Cashiriari-2 (based on the presence
of tadpoles), although additional
land excavation and site operation
may have greater effects in the
future. Any such changes in habitat
affect the structure of the amphibian communities.
A monitoring program for
amphibians in the Lower Urubamba
will help determine whether
changes in species distribution have
taken place because of habitat
alterations. If conducted for several
years, it can also determine which
species, if any, have been impacted
by past operations and whether
changes in the fauna occur over
time in response to large-scale
environmental alterations. Changes
in the fauna due to climate shifts or
atmospheric perturbations are not
expected in the near future. However, analyses of changes in amphibian abundance and distribution
must take into account possible
changes in the abiotic environment.
For the biotic and abiotic results to
be comparable in space and time,
standardized protocols must be
established for both data sets.
Information on changes in
amphibian communities and the
abiotic environment provides
insights into how management and
restoration efforts may be able to
restore amphibian species and
communities in the impacted areas.
The baseline inventory of amphibians and reptiles at San Martin-3
and Cashiriari-2, along with an
understanding of the differences in

habitat structure and layout of
trails, roads, and streams, allows
identification of which species
groups will respond to standardized
monitoring techniques. Based on
preliminary inventories of amphibians and reptiles at the two sites
(Reynolds et al., Icochea and

Mithchell, this volume), the known
amphibian fauna of the study
region consists of salamanders (one
species), caecilians (one species),
and frogs (48 species). The reptile
fauna is comprised of 15 lizards and
24 snakes. Because these groups
Figure 1. Example of nocturnal microhabitats of selected frog species in

the Lower Urubamba region.
237
Frogs are
diverse,
numerous,
and relatively well
known
taxonomically and
are sufficiently
active at
predictable
times.
238
have not been inventoried completely in this region, additional

species will be discovered as more
time is spent at the well sites. A
comparison of the known amphibian faunas in rain-forest habitats of
Lower Urubamba, Manu, and
Iquitos, Per (Table 1) demonstrates that most amphibian species
in Peruvian rain forests belong to
two families of frogs (treefrogs and
leptodactylids). The former is
primarily arboreal, whereas the
latter is primarily terrestrial and
arboreal on low vegetation.
Leptodactylids are especially wellrepresented at San Martin-3 and
Cashiriari-2. Figure 1 shows the
structure of the frog community in
primary forest based on observations in the Lower Urubamba.
because many (especially caecilians
and snakes) are secretive and are
encountered in too few numbers to
provide information other than
presence or absence. Lizards must
be monitored with specialized
trapping techniques. Frogs are
diverse, numerous, and relatively
well known taxonomically and are
sufficiently active at predictable
times. For these and other reasons
(see below), frogs are the primary
target group for this long-term
monitoring plan. Baseline information on the entire herpetofauna will
be accumulated as the monitoring
project continues.
The primary goals of the amphibian monitoring plan are to: 1)
evaluate the effects of the construction and operation of each
well site and the proposed pipeline
on frog communities, 2) evaluate
the long-term trend in amphibian
community structure and species
distributions relative to habitat
changes in the Lower Urubamba,

and 3) add to the faunal inventories
and knowledge of the natural
history of each species at each site
using data from the monitoring
project. These goals will be met by
using monitoring protocols proposed in Heyer et al. (1994) and
adapted to the topography and
logistical support system that will
be in place at the well sites. This is
the first long-term (10 to 20 years)
standardized monitoring study
proposed for amphibians in
neotropical rain forests. The results
will be invaluable to the continued
evaluation of the world-wide
decline in these animals because we
will be able to ascertain whether
amphibians are declining in numbers of species here for reasons
other than the habitat modifications
generated by the well sites.
Methodology
Study Species
Two characteristics make frogs
especially amenable to monitoring
techniques: most frogs are active at
night when it is wet and males
produce species-specific vocalizations (Duellman and Trueb 1986).
Frogs are readily visible at night on
the ground and while sitting on
leaves and branches with the aid of
a strong headlamp. Once learned,
their calls are easily recognizable
and can be taped with a tape recorder for permanent record (see
Fig. 2 for an example of a frog
sonogram) and for verification in
those cases where the field researcher is uncertain of the identification. These characteristics also
make frogs especially appropriate
for monitoring projects that incorporate students and relatively
inexperienced biologists. In addition, methods for standardized
Table 1. Comparison of the known amphibian fauna for tropical sites
in Per (the number of amphibian species found in the Lower

Urubamba region at San Martin-3 and Cashiriari-2 [data from Reynolds et al. and Icochea and Mitchell this full report] are compared
to Manu [data from Morales and McDiarmid 1996] and Iquitos [data
from Rodriguez and Duellman 1994] )
Family
Bufonidae
Dendrobatidae
Centrolenidae
Hylidae
Leptodactylidae
Microhylidae
Pipidae
Ranidae
Urubamba
Manu
3
4
3
6
1
27
26
4
11
30
1
monitoring techniques based on

frog vocalizations are currently
being worked out in the United
States and Canada (Heyer et al.
1994, Green, 1997).
Monitoring Techniques
Two commonly used techniques are being tested in the field
during the first phases of the am-
Iquitos
7
8
50
37
7
2
1
phibian monitoring program. They

are linear transects and plot sampling.
Linear transects operated at
night have been shown by a single
comparative study in Ecuador to be
an effective technique for monitoring terrestrial and arboreal frogs in
primary rain forest and other associated habitats in the tropics
Two
commonly
used techniques are
being
tested in
the field
during the
first phases
of the
amphibian
monitoring
program.
Figure 2. Sonogram of the mating call of a male Ischnocnema

quixensis from Per (from Schluter and Icochea 1995).
239
Figure 3. Diagram of San Martin-3 well site at the Lower Urubamba

region. Transects for the study of amphibians are indicated along the
existing trails.
(Pearman et al. 1995). This technique allows one to encounter frogs
on the ground and on foliage and
branches. It has proven to be more
effective than searches in specified
plots and assessments using specialized trapping techniques. Frogs that
are calling 50 or more meters from
the trail can be identified by their
vocalizations, although rainfall will
attenuate detection distances
especially for high frequency calls
(Zimmerman 1994), and the numbers of individual males calling can
240
be quantified. Teams of trained

field biologists walk along the trail
and search carefully for frogs sitting
on the ground and perched on
leaves or branches. The effective
distance for visually encountering
frogs is about 1 to 3 m on each side
of the trail (about a 4 to 6 m wide
search area). This approach is been
termed the Visual Encounter
Survey technique (Crump and Scott
1994). Team members also record
the identification and number of
calling males in the transect (or the
audio strip transect; Zimmerman

1994). Calling males of many
tropical frogs are widely dispersed
in the forest or occur in small
enough groups to be counted accurately by sound.
To facilitate statistical comparisons, the number and length of
transects must be standardized
among sites. Two or more trails of
varying lengths will be associated
with each drilling site. In this
project, the trails are divided into
100-m segments on a map and
numbered individually. A total of
four segments, separated from each
other by one or more additional
segments, will be selected for
monitoring using a random numbers
table. The assumption is made that
frog vocalizations and activities in
one transect are independent of
activities in another when separated
by 100 m or more. An example of
the sampling design is illustrated in
Figure 3.
Monitoring frogs in permanent
1-ha plots has been shown to be an
effective method in tropical rain
forests (Salas et al. 1988).
Operationally, the methods are
similar to transect sampling, with
individual frogs identified and
counted as they are encountered
visually and aurally. The primary
differences are that a specified area
(1 ha) is searched intensively to
obtain density estimates and most
of the plots are not located randomly.
A total of four plots per drilling
site, two of which are permanent
SI/MAB plots (see Alonso et al.
this volume), are being searched by
teams of trained field biologists
over a six-night period. One plot
encompasses all or part of the drill
site. A fourth will be established in
a random location. Each plot must
be sampled a minimum number of

times to account for daily variation
in rainfall and frog activity.
Field Teams
Teams of at least two people,
all of whom have training in identification of the frogs (and their
vocalizations) in the Lower
Urubamba, monitor each plot or
transect for six nights. A field
operations coordinator oversees
sampling at transects, plots, and
gathering abiotic data and assists in
field monitoring. Each team is led
by a person with experience in
identifying frogs and their vocalizations. Given the number of
transects and plots (four each) at
each drilling site and because of
hourly variation in rainfall patterns
and frog activity, project staff are
considering the possibility of
gathering three teams of two people
to conduct future monitoring
surveys (one team for transects and
two teams for the plots).
This plan is based on monitoring four sites annually over a onemonth period during the rainy
season (see below). If additional
sites are added, the field team will
extend the time spent in the Lower
Urubamba
Seasonal Timing
Because of the strong seasonal
nature of frog activity in the tropics
(species-specific male vocalizations
are tied to reproduction, which
occurs during the rainy season), all
transects at all sites must be run
when the probability of encountering frogs is highest. That period is
the first month of the rainy season
(Duellman 1978, Morales and
McDiarmid 1996). Few frogs vocalize in the dry season, and conducting monitoring efforts at that time
241
The
amphibian
monitoring
project in
the Lower
Urubamba
region
should be
operational
for at least
10 years
with a
target
length of
20 years.
242
will produce insufficient numbers

for statistical analysis and skewed
results. To ensure comparability
among sites, all plots and transects
will be monitored during the same
month each year of the long-term
project. Because of logistics and
helicopter schedules, it is unrealistic
to alternate among sites on a daily
or short-term basis, the preferred
protocol used in other studies to
minimize differences in results
based on weather patterns. Weekly
rotation is preferred, if possible,
with the monitoring period covering
a total of four weeks (assuming four
sites are available for monitoring).
Thus, in the Lower Urubamba the
target month for the long-term
monitoring study is November or
December.
Field Methods for
Individual Animals
Each individual frog is being
identified through visual inspection
by experienced herpetologists
(catching and holding for identification, if necessary, so that keys and
identification guides can be consulted) or by its species-specific
vocalization. The position of each
frog relative to the nearest transect
marker and its microhabitat is noted
on the data sheet. Frog calls are
recorded on cassette tape recorders,
and estimates of male population
density ranked as 1 for single
males, 2 for a chorus of two to
five males, 3 for a chorus of six
to10, and 4 for choruses of more
than 10 individuals, following
guidelines in Bishop et al. (1994).
Presence of eggs, tadpoles, or
amplectant (mating) pairs of frogs
serve as indicators of reproductive
activity. All metamorphosed frogs
observed are recorded as subadult
or adult as appropriate for each
species or species group. Environmental data (air temperature,

rainfall, relative humidity) are taken
at the start of each search.
Data Management and
Length of Study
All data are entered into a
standardized database for storage
and statistical analysis. Copies of
this database will reside with the
appropriate Peruvian museum
(original data in, for example, the
Marcos), the Smithsonian Institution, and with the principal researchers (both Peruvian and North
American). Example data fields are
date, site, transect, names of field
biologists, air temperature, soil
temperature, rainfall, transect
section, species, visual identification or call recorded and identified,
location of frog above ground,
perch type (leaf, branch, ground),
distance to nearest transect marker,
sex, relative age (juvenile or adult),
and number of individuals in the
chorus.
Detection of trends over time
(stable, declining, increasing) in
amphibian populations has been a
challenging task. This is because
populations sizes of many species,
especially frogs, fluctuate dramatically from year to year (Pechmann
et al. 1991). The wide variation
induced by the fluctuating numbers
of breeding individuals makes
detection of statistical significance
difficult. Attempts to deal with this
problem have focused on the use of
power analysis to determine if
existing data are sufficient to
provide statistical confidence in the
hypothesis that there was no change
in population size over time (Reed
and Blaustein 1995). Recent inves-
tigations suggest that 10 to 20 years

of annual data are needed to determine if any real change in the status
of a population has occurred
(Hayes and Steidl 1997; S. Droge,
personal communication). Although
no information is available on the
long-term dynamics of amphibian
populations in the neotropics, this
suggests that the amphibian monitoring project in the Lower
Urubamba region should be operational for at least 10 years with a
target length of 20 years.
Data Acquisition and Abiotic
Measurements
The accumulation of abiotic
measurements is essential to all of
the monitoring projects in the
Lower Urubamba. The preferred
method for measuring maximum
and minimum temperature, rainfall,
relative humidity, barometric pressure, wind speed, and wind direction is to set up a weather station at
each site that automatically records
these data on a daily basis and
records them in electronic format.
The following abiotic measurements
should be taken during each day of
the monitoring period and preferably for several weeks prior to
monitoring. If automatic recording
of abiotic data is not possible or if
these instruments are inoperative,
hand-held instruments to obtain
these data should always be brought
to the site by the field crew.
General Data Acquisition
1. Data sheet for data records.
2. Map of each plot and
transect with segments and habitats
delineated.
3. Descriptions (written with
sketches and photographs) of the
micro-habitats and vertical struc-
ture of the habitat for each plot and

segment studied.
Abiotic measurements
1. Temperature (soil, air):
minimum/maximum for each 24hour period at each site.
2. Precipitation (daily, with rain
gauge).
3. Acidity (pH) of aquatic sites
(including tree holes and bromeliad).
4. Relative humidity.
5. Wind speed and direction.
Voucher Specimens
Scientific documentation of the
occurrence of any amphibian or
reptile species at a particular site is
accomplished by the collection of a
voucher specimen or voucher series.
A small number of each species
previously unrecorded for each site
is being sampled and preserved
properly for permanent storage in a
museum devoted to maintaining
such collections. Appropriate
museums are the Museo de Historia
Mayor de San Marcos and the
Smithsonian Institution. These
specimens provide the core of the
baseline inventory database and
represent a permanent record of the
species that occur in the Lower
Urubamba.They also are like books
in a library from which scientists
can obtain much pertinent information on the biology of each species
that is not possible to learn in the
field (e.g., condition of reproductive organs at the time of monitoring).
Each voucher specimen is
euthanized humanely and preserved
with a small quantity of dilute
formaldehyde, tagged with an
individually numbered tag, and
stored in the museums herpetological collection in alcohol. The
243
project field operations coordinator

is fully trained in all techniques to
conduct this procedure.
Accurate
identification guides
to adult
frogs and
male vocalizations
(cassette
tapes) are
essential
for the
successful
execution
of this
project.
Training Sessions
and Materials
Sessions designed to train field
personnel in field monitoring techniques and data collection is critical
to a successful program. Two- to
four-day training sessions immediately prior to each annual monitoring period will assist new participants. The sessions are conducted
by leaders experienced in the Lower
Urubamba.
Accurate identification guides
to adult frogs and male vocalizations (cassette tapes) are essential
for the successful execution of this
project. Unfortunately, such field
guides and tapes do not exist for
most of the neotropical frog fauna,
and none are available for use in the
Lower Urubamba. Photographs of
frogs and recordings of frog calls
obtained during the inventory phase
at San Martin-3 and Cashiriari-2 are
providing some of the materials
needed to create these guides. In
the next phases of the project,
researchers will continue to develop
accurate field guides and tapes of
male vocalizations.
Logistical Support
Coordination of amphibian
monitoring project personnel with
access at the right time of year in
all sites is crucial to the successful
execution this project. All personnel, their support staff, and other
logistics must be in place at the
start of the rainy season. Project
staff will work with appropriate
representatives to ensure that the
logistical support is appropriate for
the amphibian monitoring project.
244
Integration with Other

Monitoring Projects
Because other monitoring
projects are taking place concurrently with amphibian monitoring,
project staff are responsible for
discussing plans and plotting maps
where their various teams will be
working on a daily basis. This level
of coordination is needed to ensure
interference is minimized and that
certain types of data such as such
as weather data are sampled in a
standardized manner across disciplines by all members of the team.
Products
Several products will result
directly from the amphibian monitoring project. These include management recommendations for the
recovery of the amphibian community, a field guide to the amphibians
of the Lower Urubamba region, a
cassette tape of the frog calls of
this area, several technical scientific
papers in the conservation biology
and scientific literature based on
the quantitative results of the
study, and, of course, annual reports. Popular articles for the lay
public will also be written for
magazines. All publications will
acknowledge the contributions and
financial support of Shell Prospecting and Development (Per) B.V.
The technical products will also
serve as the literature base for
future decisions on management
and restoration.
References
Anderson, I. 1995. Is a virus wiping

out frogs? New Scientist (January):
7.
Bishop, C. et al. 1994. A proposed
North American amphibian monitoring program. Unpublished
report, U.S Program, IUCN Declining Amphibian Population Task
Force.
Blaustein, A. R., P. D. Hoffman,
D. G. Hokit, J. M. Kiesecker, S. C.
Walls, and J. B. Hays. 1994a. UV
repair and resistance to solar UV-B
in amphibian eggs: A link to
population declines? Proceedings of
the National Academy of Science 91:
1791-1795.
Blaustein, A. R., D. G. Hokit, R. K.
OHara, and R. A. Holt. 1994b.
Pathogenic fungus contributes to
amphibian losses in the Pacific
Northwest. Biological Conservation
67: 251-254.
Blaustein, A. R., and D. B. Wake.
1990. Declining amphibian populations: A global phenomenon?
Trends in Ecology and Evolution 5:
203-204.
Blaustein, A. R., D. B. Wake, and
W. P. Sousa. 1994c. Amphibian
declines: Judging stability, persistence, and susceptibility of populations to local and global extinctions. Conservation Biology 8: 60-71.
Crump, M. L., and N. J. Scott, Jr.
1994. Visual encounter surveys.
Pages 84-92 in Measuring and
Monitoring Biological Diversity:
Standard Methods for Amphibians.
(W. R. Heyer, M. A. Donnelly,
R. W. McDiarmid, L. C. Hayek,
and M. S. Foster, eds.) Smithsonian
Institution Press, Washington, DC.
Delis, P. R., H. R. Mushinsky, and
E. D. McCoy. 1996. Decline in
some west-central Florida anuran
populations in response to habitat
degradation. Biodiversity and Conservation 5: 579-1595.

Duellman, W.E. 1978. The biology
of an equatorial herpetofauna in
Amazonian Ecuador. Miscellaneous Publication 65. University
of Kansas, Museum of Natural
History, Topeka.
Duellman, W. E., and L. Trueb.
1986. Biology of Amphibians.
McGraw-Hill, New York.
Fisher, R. N., and H. B. Shaffer.
1996. The decline of amphibians
in Californias Great Central
Valley. Conservation Biology 10:
1387-1397.
Green, D. M. 1997. Amphibians in
decline: Canadian studies of a
global problem. SSAR Herpetological Conservation 1: 1-340.
Hayes, M. P., and M. R. Jennings.
1986. Decline in ranid frog species
in western North America: Are
bullfrogs (Rana catesbeiana) responsible? Journal of Herpetology 20:
490-509.
Hayes, J. P., and R. J. Steidl. 1997.
Statistical power analysis and
amphibian population trends.
Conservation Biology 11: 273-275.
Heyer, W. R., M. A. Donnelly, R. W.
McDiarmid, L. C. Hayek, and
M. S. Foster (eds.). 1994. Measuring and Monitoring Biological
Diversity: Standard Methods for
Amphibians. Smithsonian Institution Press, Washington, DC.
Lannoo, M. J., K. Lang, T. Waltz,
and G. S. Phillips. 1994. An altered
amphibian assemblage: Dickenson
County, Iowa, 70 years after Frank
Blanchards survey. American
Midland Naturalist 131: 311-319.
Morales, V. R., and R. W.
McDiarmid. 1996. Annotated
checklist of the amphibians and
reptiles of Pakitza, Manu National
Park Reserve Zone, with comments on the herpetofauna of
245
Madre de Dios, Per. Pages 503522 in Manu. (D. E. Wilson and A.

Sandoval, eds.) Smithsonian Institution Press, Washington, DC.
Pearman, P. B., A. M. Valasco, and
A. Lopez. 1995. Tropical amphibian monitoring: a comparison of
methods for detecting inter-site
variation in species composition.
Herpetologica 51: 325-337.
Pechmann, J. H. K, D. E. Scott,
R. D. Semlitsch, J. P Caldwell, L. J.
Vitt, and J. W. Gibbons. 1991.
Declining amphibian populations:
The problem of separating human
impacts from natural fluctuations.
Science 253: 892-895.
Phillips, K. 1990. Where have all
the frogs and toads gone? Bioscience
40: 422-424.
Reed, J. M., and A. R. Blaustein.
1995. Assessment of
nondeclining amphibian populations using power analysis. Conservation Biology 9: 1299-1300.
Rodiguez, L. O., and W. E.
Duellman. 1994. Guide to the
frogs of the Iquitos region, Amazonian Per. Special Publication
22. University of Kansas, Natural
History Museum, Topeka, Kansas.
Sadinski, W. J., and W. A. Dunson.
1992. A multilevel study of effects
of low pH on amphibians in
temporary ponds. Journal of Herpetology 26: 413-422.
Salas, A. W., G. Tello, W. Arizabal,
P. J. Sehgelme Ible, and D. Neyra.
1998. Monitoring leaf-litter amphibians in Manu National Park.
Pages 615-632 in Forest
Biodiversity in North, Central and
South America, and the Caribbean:
Research and Monitoring (F.
Dallmeier and J. A. Comiskey, eds.)
Man and the Biosphere Series, v.
21 UNESCO, Pearl River, New
York.
246
Schluter, A. and J. Icochea. 1995.

Natural history: Ischnocnema
quixensis, vocalization. Herpetological Review 36: 144.
Schwalbe, C. R., and P. C. Rosen.
1988. Preliminary report on effect
of bullfrogs on wetland
herpetofaunas in southeastern
Arizona. Pages 166-173 In Management of Amphibians, Reptiles,
and Small Mammals in North
America. (C. S. Szaro, K. E.
Severson, and D. R. Patton, eds.)
General Technical Report RM-166,
U.S. Department of Agriculture,
Forest Service, Washington, DC.
Sinsch, U. 1990. Migration and
orientation in anuran amphibians.
Ethology, Ecology & Evolution 2:
65-79.
Wake, D. B. 1991. Declining amphibian populations. Science 253:
860.
Wyman, R. L. 1988. Soil acidity and
moisture and the distribution of
amphibians in five forests of
southcentral New York. Copeia
1988: 394-399.
Wyman, R. L. 1990. Whats happening to the amphibians? Conservation
Biology 4: 350-352.
Zimmerman, B. L. 1994. Audio
strip transects. Pages 92-97 in
Measuring and Monitoring Biological Diversity: Standard Methods
for Amphibians. (W. R.Heyer, M.
A. Donnelly, R. W. McDiarmid,
L. C. Hayek, and M. S. Foster,
eds.) Smithsonian Institution Press,
Washington, DC.
Birds:
Biodiversity
Assessment
in the Lower
Urubamba
Region
George Angehr
Smithsonian Tropical Research
Institute
Constantino Aucca
Abad del Cusco
Introduction
Birds are probably the most

widely studied vertebrate animal
group, primarily because of their
great diversity, their considerable
influence on the environment, and
their place in human and cultural
history. In all communities, even
those dominated by humans, birds
are important consumers, predators,
and prey of other organisms. Their
role in controlling insect populations, including agricultural pests, is
well known. In many environments,
birds are the key dispersal agents of
plants, usually by spreading seeds
after feeding on a variety of fruits,
cones, and nuts. A few important
groups of birds have evolved as
exclusive plant pollinators, feeding

on flower nectar and pollen. In their
roles as seed dispersers and pollinators, birds serve as mobile links
among various food webs. In this
capacity and as predators and
consumers, some birds are keystone species that largely determine the structure and function of
natural communities. Ultimately,
birds act to shape habitat for other
organisms, including other birds. In
addition, the nests of birds serve as
habitat for a wide range of organisms, and the birds themselves are
hosts to a suite of specialized
parasites and commensals.
The bird communities of the
Amazonian lowlands of Per are
among the most diverse in the
world. However, relatively few sites
within this vast area have been
surveyed by ornithologists. In
southeastern Per, good information exists for several sites in Madre
de Dios province, including Manu
National Park and its environs
(Terborgh et al. 1990, Servat 1996),
the Tambopata Reserve (Donahue
et al. MS.) and the Reserva Cusco
Amaznico (Davis et al. 1991). In
247
contrast, little work has been done

in the lowlands of Cusco Province
immediately to the west. The upper
course of the Urubamba River, the
areas major drainage, has been
extensively surveyed (Chapman
1921, Parker and ONeill 1980); the
Lower Urubamba region has not.
As part of the Smithsonian
Institution Monitoring and Assessment Program and Shell Prospecting
project, we surveyed birds at the
sites. Baseline surveys and subsequent monitoring efforts will help
the company anticipate and mitigate
any negative impacts of their
activities on biodiversity in the
region. Our objectives were: 1) to
obtain preliminary bird species lists
at the two well sites and 2) gather
data such as mist-net capture rates
and species accumulation curves
that are needed to devise a longterm monitoring plan for tracking
changes in the bird community that
might result from gas development
activities.
Methods
At the two well sites, we

surveyed birds from April 11 to
May 7 (San Martin-3) and May 7 to
May 16 (Cashiriari-2), 1997. Base
camps at the sites are located on
ridges at 474 m and 579 m above
sea level, respectively, so we
worked in area from about 400 to
500 m in elevation at San Martin-3
and 500 to 600 m in elevation at
Cashiriari-2. Rainy periods were
frequent during the survey at San
Martin-3, but were less frequent by
the second week of May during our
stay at Cashiriari-2. We originally
planned to spend equal amounts of
time each site, but logistical
248
challenges resulted in considerably

more time being spent at San
Martin-3 than at Cashiriari-2.
As noted throughout this
report, San Martin-3 and
Cashiriari-2 differ substantially in
their forest composition. San Martin
is heavily dominated by Guadua
sarcocarpa (bamboo), while
Cashiriari is located in a more
typical moist forest (see Alonso et
al. this volume). At each site, the
habitat is relatively homogeneous,
at least with respect to variables
that have been found to be of
significance to birds in southeastern
Per (Terborgh et al. 1990,
Robinson and Terborgh 1990,
Servat 1996, Kratter 1997). Neither
site is close to a large river, lake,
swamp, or seasonally flooded forest.
In Amazonia, a large number of
birds are confined to habitats near
water (Remsen and Parker 1983),
including a variety of forest species,
not just aquatic and semi-aquatic
species. In addition, large rivers
produce a series of successional
habitats by eroding their banks and
depositing new land in backwaters
that is soon colonized by vegetation. Each of these successional
habitats may have its own characteristic bird community. Both San
Martin-3 and Cashiriari-2 have a
number of small streams, but they
are generally so small that the forest
canopy is usually closed above
them. A somewhat larger stream,
approximately 5 m wide and generally open to the sky, runs within
one km of San Martin-3. However,
neither this stream nor the others at
the sites are sufficiently large to
attract many of the birds typical of
Amazonian water-side habitats, nor
are they large enough to produce
significant habitat heterogeneity
through erosion and deposition.
Although quite homogenous

with regard to major habitat type,
the forest composition at each site
does show some local variation. At
San Martin-3, the main habitat
variable is the relative proportion
of bamboo in different sites, which
appears to be influenced primarily
by drainage. The area is a complex
mosaic of forest and bamboo, with
the moister ravine bottoms containing a higher percentage of trees
than the drier ridge tops that are
more strongly dominated by bamboo. At Cashiriari-2, the most
noticeable habitat variable is forest
age. Most of the forest in the
vicinity of the site appears to be
primary. However, when the site
was originally developed approximately 10 years ago, roads were
constructed to the villages of
Shivankoreni and Segakiato. The
area adjacent to these roads was
apparently cleared 10 to 15 m in
many places, and in some cases
trees were cut to construct bridges.
These roads are now abandoned,
and a dense second-growth forest,
dominated by Cecropia and other
sucessional species, has developed.
Sampling Protocols
We used a combination of mistnetting and visual and auditory

observations to document the bird
communities at the two sites.
However, we concentrated primarily on mist-netting because of the
lack of previous work in the area
and the difficulty of visual and
auditory identification of many of
the species in this extremely diverse
community.
We used ATX type mist-nets,
12 m long by 2.6 m high with a 36mm mesh. We operated nets at
three locations in San Martin-3 and
two at Cashiriari-2. Descriptions of
the specific netting locations follow:

* San Martin-3: Location 1in
forest along the banks of a small
stream just below biodiversity plot
#1, approximately 500 m west of
the edge of the well-site clearing.
This area had relatively less bamboo
than the other two locations at San
Martin.
* San Martin-3: Location 2a
drier bamboo-dominated site on the
ridge just above biodiversity plot
#2, approximately 500 m east of
the San Martin camp.
* San Martin-3: Location 3on
a slope just below and to the south
of Location 2. The upper most nets
were in drier bamboo-dominated
habitat; the lower most were placed
along the banks of a small stream
where trees were more common.
* Cashiriari-2: Location 1in
primary forest along the waterdischarge pipeline from the well
site, beginning approximately 300 m
from the well pad.
* Cashiriari-2: Location 2in
secondary forest along the former
road to Cashiriari.
At each netting location, we
initially set up 11 to 13 nets. On
subsequent days, we added additional nets for a total of 15 to 20 at
each well sites. Individuals nets
were usually set at least 10 m apart,
but we sometimes set two or three
nets adjacent to one another. We
operated nets for three to six days
at each well site. In mist-netting
studies, the capture rate at an
individual net typically falls off
after the first day because birds
learn the nets location and avoid it.
During this study, overall capture
rates remained high enough over the
sampling period at each location so
that we did not have to move the
nets to maintain a desirable capture
At San
Martin-3,
the main
habitat
variable is
the relative
proportion
of bamboo in
different
sites,
which
appears to
be influenced
primarily
by
drainage.
249
We
identified a
total of 198
species at
the two sites
combined
164 at San
Martin-3
and 98 at
Cashiriari-2.
250
rate. We only had to relocate a few

unproductive nets.
Weather permitting, we opened
nets shortly after dawn and closed
them just before dusk. However, at
San Martin rain often limited netting to only parts of a day; we
netted for a few hours in the morning and for several hours during a
second session in the afternoon.
Data taken on captured birds
included weight, wing chord, tail
length, exposed culmen, and tarsus.
Birds were checked for molt and for
presence of a brood patch or other
evidence of breeding. Captured
birds were temporarily marked by
clipping a tail feather to avoid
taking data on the same bird more
than once. (However, a few birds
escaped or were released without
being marked, and others may have
lost the clipped feather through
molting.) At least the first individual of each species captured was
photographed to provide a voucher
of the species occurrence.
It should be emphasized that
mist-nets only sample a portion of
the entire bird community present
at a site. They primarily sample
birds of the forest understory and
rarely capture species that spend
most of their time foraging above
the height of the nets. They effectively capture most species between
5 and 100 grams in body weight,
but larger birds fail to become
entangled and often escape, while
the smallest hummingbirds are
sometimes able to pass through the
mesh. Nets rarely capture birds that
forage primarily by walking on the
ground, possibly because such
species move along slowly enough
so that they see the net and avoid
it.
On days when we did not mistnet, we walked the trails and along
streams at each site, particularly in
the morning and late afternoon, to
observe species not susceptible to
capture with mist-nets. Whenever
possible, we recorded calls with a
Sony TCM-5000 portable tape
recorder with a Sennheiser short
shotgun microphone to provide a
voucher of the species occurrence
at the site.
Results and
Discussion
The species found at San

Martin-3 and Cashiriari-2 are shown
in Appendix 1. We identified a total
of 198 species at the two sites
combined164 at San Martin-3
and 98 at Cashiriari-2. The lower
total for Cashiriari most likely is
due to the fact that we spent much
less time there. In addition, nearly
all of our time at the latter site was
occupied in mist-netting so that we
did not have much opportunity to
observe those species not susceptible to capture with nets. Sixty-six
of the total species were observed
at both sites, 99 only at San Martin3, and 32 only at Cashiriari-2.
Again, the lower number of species
observed only at Cashiriari is likely
a result of the shorter amount of
time we spent there.
Several sites in Per, including
Cocha Cashu in Manu National
Park and the Tambopata Reserve,
have species lists of well over 500.
Much of that diversity is due to the
high diversity of habitats found at
those sites, which include river
edges, swamps, and oxbow lakes in
addition to upland and bamboo
forests. Also, those species lists
have been compiled over a period
of many years. The lower number

of species reported for San Martin-3
and Cashiriari-2 stems from the
relatively homogeneous habitat at
each site and to the fact that the
surveys were carried out over a
brief period of time.
An examination of the number
of species recorded just within a
single habitat shows that the Lower
Urubamba sites are indeed more
similar to others in southeastern
Per. For example, at Cocha Cashu,
Robinson and Terborgh (1990)
reported 212 resident species (plus
an additional 70 species recorded
irregularly) from transition forest,
239 resident and 29 irregular species from high-ground forest, and
245 resident and 28 irregular species from upland forest. These
figures were obtained from intensive studies over 17 years. In a nineyear study, Servat (1996) reported
415 species from Pakitza, of which
102 species were found in Guadua
bamboo thickets.
Species Richness and Species

Accumulation Curves
Mist-net samples provide a
standardized means to compare
species richness of the understory
bird communities at different sites.
Although mist-nets do not sample
the entire bird community, Karr et
al. (1990) found that overall differences in species richness at four
sites in Costa Rica, Panama, Brazil,
and Per were paralleled by differences in species richness in their
understory bird communities, as
shown by mist-net studies in those
tropical sites.
Data from mist-net sampling at
the Lower Urubamba sites in San
in Appendices 2 and 3. At San
Martin-3, we obtained 86 species
in a total of 440 captures (representing 360 individuals). At
Cashiriari, we obtained 62 species
in a total of 248 captures (representing 224 individuals). Species
90
80
70
Species
60
50
40
San Martin
Cas hiriari
30
Manu
20
Panam a
Brazil
10
Cos ta Rica
55
0
44
0
40
0
35
9
32
3
30
0
28
3
24
8
20
0
17
1
10
9
10
0
87
61
Captures
Figure 1. Species accumulation curves for San Martin-3 and Cashiriari-2 (diamonds
= San Martin-3, squares = Cashiriari-2). The line with triangles indicates the species
accumulation curve from Manu National Park, derived from Karr et al. 1990, the
line with crosses represent Panama, the one with asterisks represents Brazil, and the
one with circles Costa Rica (see text).
251
30
San Martin-3
25
Individuals
20
15
10
0
1
13
17
21
25
29
33
37
41
45
Rank
49
53
57
61
65
69
73
77
81
85
18
Cashiriari-2
16
14
Individuals
12
10
8
6
4
2
0
1
10
13
16
19
22
25
28
31
Rank
34
37
40
43
46
49
52
55
58
61
Figure 2. Ranked species abundances from mist-net samples at San

Martin-3 and Cashiriari-2.
accumulation curves are shown in
Figure 1 and graphs of ranked
species abundances in Figure 2.
Karr et al. (1990) compared
species accumulation curves from
mist-netting studies in high-ground
forests at Cocha Cashu in Manu
National Park and from other
tropical forest sites in Costa Rica
(La Selva), Panama (Soberania
National Park), and Brazil (Biological Dynamics of Forest Fragments
Project area 70 km north of
Manaus). In general, Manu showed
the highest species richness and the
highest species accumulation curves
of any of these sites
Cashiriari-2 shows a species
accumulation curve almost identical
252
to that at Manu, while San Martin is

substantially higher (Fig. 1). Karr et
al. provided data on 1000-capture
samples in their figure 14.1. From
that figure, we estimate that a 248capture sample from the Manu
would yield 62 species, equal to our
actual result from Cashiriari-2. In
contrast, a 440-capture sample from
Manu would yield only 80 species,
substantially lower than the 86
species we recorded at San
Martin-3.
As with many other tropical
sites, there were only a few common species and many rare ones in
our samples. At San Martin-3, 83%
of the species captured each made
up 2% or less of the total captures,
while at Cashiriari-2 the corresponding figure was 77%. Corresponding figures from the other
sites discussed by Karr et al. were
between 78% and 88%. At San
Martin-3, 30% of all species and at
Cashiriari-2, 34% of all species
were represented by only a single
capture.
Species accumulation curves
for individual habitats in Manu
National Park are provided by
Robinson and Terborgh 1990.
According to the habitat classification of those authors, upland
forests grow on terraces well above
the floodplain of the Manu River.
They are dissected by steep ravines
and contain extensive patches of
bamboo, they appear to be the
closest parallel to the habitat type
of San Martin-3. As shown in
Robinson and Terborghs Figure
12.1, species accumulation curves
from upland forest closely parallel
those from highland forest, discussed by Karr et al. Therefore, it
appears that San Martin-3 exceeds
any single forest habitat at Manu in
species diversity.
We are not certain why there is
such high species richness at San
Martin-3. One reason could be that
mist-nets sample a high proportion
of the total community in this
habitat. Because the canopy is not
continuous, birds that normally
forage in higher strata of the forest
may descend and be more susceptible to capture. However, canopy
species were generally scarce at San
Martin-3, as discussed below, and
this did not appear to be a significant factor in producing high species richness in the mist-net
samples. Another possibility is that
the discontinuous canopy allows
sunlight to penetrate into lower
levels, producing a dense and
presumably highly productive

understory layer. Another factor
could be the highly complex mosaic
of forest and bamboo present at the
site. A large number of bamboo
specialists are present at San
Martin-3 (see below), as well as a
variety of birds typical of moist
forest. San Martin-3 may represent
an intersection between the avifaunas of bamboo and moist forest.
Breeding and Molt
There was little evidence of
breeding activity at either site. In
the mist-net samples, no birds were
observed with brood patches One
female rufous-breasted piculet
(Piculus rufiventris) appeared to be
gravid and ready to lay. Only two
recently fledged birds were captured, one black-spotted bare-eye
(Phlegopsis nigromaculata) and one
slate-colored grosbeak (Pitylus
grossus), both at San Martin-3.
Singing and other vocal activity
generally was very low.
In contrast, many individuals
were in molt at both sites. At San
Martin-3, 64% of all mist-netted
birds examined (316) showed some
sign of molt. The percentage at
Cashiriari, 43%, was somewhat
lower.
In the seasonal tropics, breeding activity usually occurs from the
late dry season to the early part of
the rainy season when insect abundance is highest as insects respond
to the flush of leaves at the start of
the rains. In addition, fruit and
nectar availability may peak at this
time. Because molt is an energetically costly process, most birds wait
until after the breeding season is
finished to carry it out. The fact
that a large number of birds at each
site were in molt implies that they
were post-breeding. Since our
It appears
that San
Martin-3
exceeds
any single
forest
habitat at
Manu in
species
diversity.
253
Table 1. Mist-net capture rates and numbers of species captured at each location sampled
(net-hours calculated by multiplying the number of nets by the number of hours they are
in operation; first-day capture rates are those on the rist day of operation at the location
[see text])
Site
San Martin-3
San Martin-3
San Martin-3
Total
Location
1
2
3
1
Cashiriari-2
2
Cashiriari-2
Total
Project Totals
Habitat
Net hours Captures
First-day Species
Captures/
100 net-hours capture rate
729.25
stream side bamboo
392.25
ridgetop bamboo
ridgetop+stream bamboo 674.5
1802
180
122
138
440
25
31
20
24
45
33*
36*
64
42
54
87
401.75
263.25
665
2467
109
139
248
27
53
39
35
83
37
45
62
primary forest
secondary forest
* To calculate first day capture rates for San Martin Locations 2 and 3, we combined data from the
first two days of net operation, because nets were open for less than half a day each of the first
two days due to rain.
sample took place in the late rainy

season, approximately six months
after the presumed peak of breeding
in the early rainy season, this
conforms with the general pattern
of breeding in tropical birds.
Capture Rates
Mist-net capture rates for San
in Table 1. Capture rates can be
compared on the basis of the
number of captures (including
recaptures) per 100 net-hours, with
net-hours calculated by multiplying
the number of nets in operation by
the number of hours they were
open. In mist-net studies, capture
rates typically are highest for the
first day of operation at a location
and gradually decline as birds learn
where the nets are. Because we
operated nets at each location for
different numbers of days and
sometimes added nets over the
course of the sample, first-day
capture rates provide the best index
of the relative capture rate at each
location. (However, at both San
Martin-3 locations 1 and 2, rain
254
limited us to less than half a day of

operation during the first two days
at each site. Therefore, for these
two locations, we combined data
from the first two days of operation
to obtain a first-day capture rate).
At San Martin-3, first-day capture
rates were highest at location 1
(streamside) with 45 captures/100
net-hours and somewhat lower at
location 2 (ridgetop bamboo) with
33 captures/100 net-hours and at
location 3 (combination of bamboo
and streamside) with 36 captures/
100 net-hours in each. At
Cashiriari-2, location 2 (secondary
vegetation along a former road) had
the highest capture rate of any
sample site83 captures/100 nethours. Cashiriari location 1 (primary
forest) was much lower with 35
captures/100 net-hours.
After three days of operation at
the same location, capture rates
typically droppedto 22-27 captures/100 net-hours (San Martin-3
locations 1-3 and Cashiriari-2
location 1). After five days, the
rates dropped to 14-17 captures/
100 net-hours (San Martin-3,
locations 1-3). Overall capture rates

for the entire sample period for
individual locations ranged from a
high of 53 captures/100 net-hours
at Cashiriari-2 location 2 to 20
captures/100 net hours at San
Martin-3 location 3. Overall capture
rates for the sites themselves were
24 captures/100 net-hours at San
Martin-3 and 36 captures/100 nethours at Cashiriari-2. The somewhat
higher overall rate at Cashiriari was
due to the very high capture rate at
location 2 and to the fact that nets
were operated for only two and
three days, respectively, at location
1 and location 2 versus five and six
days at San Martin-3.
Robinson and Terborgh (1990)
provided mist-net capture rates for
a variety of habitats near Cocha
Cashu in Manu National Park,
where they operated net-lines at
each study site for four days. The
net-lines consisted of 12-m nets
strung end to end in straight lines
of 50 to 500 m through a habitat.
Their data are displayed in Table 2.
Capture rates for forested habitats
in general were somewhat lower
than those found in this study,

ranging from a high of 36 captures/
100 net hours in transition forest to
a low of 15 captures/100 net-hours
in high-ground and ridge-top forest.
This difference could possibly be
seasonal, since all of their sampling
was carried out between August
and October. Another possibility is
that by setting nets in straight lines,
the authors may have limited their
ability to set individual nets in
optimal locations for maximizing
capture rates. Interestingly, their
highest capture rates of all, 50 to
70 captures/100 net-hours, were in
low successional vegetation along
river banks (Tessaria/cane). Our
highest capture rates, 83 captures/
100 net hours, were in more mature
successional vegetation along roads.
Avifaunal Patterns
Several differences between the
bird communities at the two well
sites are apparent (although comparisons are somewhat complicated
by the different amounts of time
spent at the two sites). The most
notable differences are the relative
Our
highest
capture
rates, 83
captures/
100 net
hours,
were in
more mature successional
vegetation
along
roads.
Table 2. Comparative data on mist-net capture rates and numbers of

species captured in different habitats in Manu National Park
(Robinson and Terborgh 1990; number after babitat indicates the
month and year of the sampling)
Habitat
Net-hours Captures Captures/100 net-hours Species
Tessaria/cane 8/83
Tessaria/cane 11/85
Tessaria/cane 8/86
Transition forest 9/80
Transition forest 10/86
High ground forest 8/81
Upland bamboo10/85
Upland ravine 10/86
Upland ridgetop 8/74
325
230
144
1000
722
1064
593
1467
1223
1165
1672
1672
1782
194
154
73
273
260
189
152
219
252
306
250
251
294
60
70
50
27
36
18
26
15
21
26
15
15
16
58
46
39
51
51
55
44
55
66
68
49
55
65
255
absence of canopy flocks at San

Martin-3 and the presence of a large
number of bamboo specialists at
the same site.
Canopy Flocks and Canopy
Frugivores
Mixed-species canopy flocks
composed of tanagers, flycatchers,
and vireos are a prominent feature
of many neotropical forests. Their
composition and ecology have been
studied in detail by Munn (1985) in
Manu National Park. However, we
rarely observed such flocks at San
Martin-3, and their diversity was
very low compared to those studied
by Munn. Of the 35 species that
Munn observed in more than half
of the canopy flocks he studied, we
found only seven (Tangara chilensis,
Tangara mexicana, Lanio versicolor,
Tachyphonus rufiventer, Dacnis cayana,
Dacnis lineata, and Cacicus cela) at
San Martin-3. The species not
found were Monasa nigrifrons, Capito
niger, Sittasomus griseicapillus,
Lepidocolaptes albolineatus, Philydor
erythropus, Philydor ruficaudatus,
Xenops rutilans, Cymbilaimus lineatus,
Pygiptila stellaris, Myrmotherula
sclateri, Myrmotherula menetriesii,
Terenura humeralis, Tyrannulus elatus,
Myiopagis gaimardii, Tolmomyias
assimilis, Tyranneutes stolzmanni,
Pachyramphus marginatus,
Pachyramphus minor, Euphonia
chrysopasta, Euphonia minuta,
Euphonia rufiventris, Tangara
schrankii, Tangara velia, Tangara
callophrys, Tachyphonus luctuosus,
Chlorophanes spiza, Vireolanius
leucotis, and Hylophilus hypoxanthus.
We spent a relatively small
proportion of our time at San
Martin-3 looking for or observing
canopy flocks as these species are
in general not very susceptible to
capture in mist-nets. We may have
256
missed a number of species that

were actually present, but they
likely were either absent or at very
low density.
Canopy and sub-canopy
frugivores are also an important
component of the avifauna in many
tropical forests. This group includes
parrots, trogons, parrots, cotingas,
and many tanagers. (Most of these
species eat insects as well, but fruit
is an important component of their
diets. Many of the tanagers listed as
members of canopy flocks above
eat both fruit and insects.) Although we recorded a number of
these species at San Martin-3, in
general their densities and diversity
appeared to be low. For example,
four species of macaw were recorded but rarely observed. We
recorded scarlet macaw (Ara macao)
on six of the 25 full days we were
at the site, chestnut-fronted macaw
(Ara severa) for six days, the redand-green macaw (Ara chloroptera)
for four days, and the blue-andyellow macaw (Ara ararauna ) once.
We never observed more than one
or two pairs of any of these species
at any one time. The mealy parrot
(Amazona farinosa) was recorded on
nine days, although few individuals
were present, and the yellowcrowned parrot Amazona ochrocephala
was recorded once. Trogons were
particularly notable by their absence, with only a single blacktailed trogon (Trogon melanurus)
observed on one occasion. Cuviers
toucan (Ramphastos cuvieri) was
recorded on 16 days, but again there
appeared to be no more than one or
two pairs present. The emerald
toucanet (Aulacorhynchus prasinus)
was observed twice, and a chestnuteared aracari (Pteroglossus castanotis)
once. Of frugivorous cotingas, the
plum-throated cotinga (Cotinga
maynana) was recorded on 3 days,

and the purple-throated fruitcrow
(Querula purpurata) once.
The most obvious explanation
for the rarity and low diversity of
mixed-species canopy flocks and
canopy frugivores at San Martin-3 is
that over most of this area, the
canopy of broad-leafed trees is not
continuous. Except in the ravines,
isolated trees rise out of the bamboo understory at some distance
from one another. It may be energetically unprofitable for canopy
species to fly the relatively long
distances to forage in these emergent trees.
We did not spend enough time
at Cashiriari-2 to record many
canopy species. However, our
impression was that canopy flocks
were much more prominent at this
site. In particular, we observed
several species of canopy tanagers
that we never saw at San Martin-3,
including Tangara schrankii, T.
xanthogastra, T. nigrocincta, and T.
cyanocollis.
Bamboo specialists
A number of bird species of
Amazonia are found exclusively or
predominantly in thickets of
Guadua sarcocarpa (bamboo). Such
species are particularly characteristic of southeastern Per, where
extensive areas of bamboo are
common. In addition, several bird
species that are found in other
habitats elsewhere in their ranges
show a preference for bamboo in
southeastern Per. These bamboo
specialists have been the focus of
studies by Kratter (1997) on the
Tambopata River (mainly at Collpa
de Guacamayos) and by Servat
(1996) at Pakitza on the Manu
River.
Kratter found that 19 species

were exclusively restricted to
bamboo thickets at his study site.
He identified four species
(Drymophila devillei, Cercomacra manu,
Hemitriccus flammulatus, and
Poecilotriccus flammulatus) as obligate
bamboo specialists, which were
restricted to bamboo thickets
throughout their geographic range.
He found that nine species were
near-obligate specialists, which
may use other habitats sparingly
away from southeastern Per, and
that six species were facultative
bamboo specialists, which frequently use non-bamboo habitats
away from southeastern Per. In
addition, he found seven other
species to be common in bamboo,
although they also used other
habitats at his study site, and six
species that were apparent bamboo
specialists, but which were too
uncommon to characterize definitely. Combining these 13 species
with the 19 confirmed bamboo
specialists yielded a total of 32
species associated with bamboo at
Tambopata.
Servat (1997) reported that 20
species were found exclusively in
bamboo at Pakitza, although three
of them (Chlorostilbon mellisuga,
Malacoptila semicincta, and Casiornis
rufa) were found in other habitats
elsewhere in the region. Two other
species, Lophotriccus eulophotes and
Capsiempis flaveola, were reported in
this habitat in previous studies at
Pakitza (Servat and Pearson 1991,
Servat 1993). Servat did not distinguish between obligate and facultative bamboo specialists.
Stotz et al. (1996) provided
lists of indicator species for key
habitats in various sub-regions of
the neotropics. As defined by those
authors, Indicator species are
The most
obvious
explanation
for the
rarity and
low diversity of
mixedspecies
canopy
flocks and
canopy
frugivores
at San
Martin-3 is
that over
most of this
area, the
canopy of
broadleafed trees
is not
continuous.
257
those that, taken as a group, exclusively define a specific ecological

and geographic province. Lists of
indicator species provide a standard
for assessing the type and condition
of the biological community being
studied. Good indicators for identifying sites for the conservation of
habitat share four characteristics:
(1) they typically occur in only one
or a very few habitats; (2) they are
relatively common; (3) they can be
detected easily; and (4) they show
high sensitivity to habitat disturbance (i.e., they become rare, or
disappear, in habitats that are
altered, overhunted, or fragmented). Stotz et al. listed 29
indicator species for bamboo habitats in their southern Amazonia
region, of which 24 have ranges
that extend to southeastern Per.
Of the 41 species identified as
associated in some way with bamboo by either Kratter, Servat, or
Stotz et al., 29 were found at San
Martin-3 (Appendix 2).These
species represent 18% of the total
species list for the site and include
17 of the 19 species identified as
bamboo specialists by Kratter and
18 of the 24 indicator species for
bamboo habitats listed by Stoltz et
al. that extend to southeastern
Per. Of the 13 species indicated as
bamboo specialists by all three
studies, 12 were found at San
Martin-3. It is likely that more
extended studies at San Martin-3
will confirm that additional species
of bamboo-associated birds are
present. Clearly, the dominance of
Guadua bamboo this well site plays
a major part in determining the
composition of the avifauna at this
site.
In contrast, only four of the
bamboo-associated species
(Malacoptila semicincta, Automolus
258
rufipileatus, Neopelma sulphureiventer,

and Leptopogon amaurocephalus) were
found at Cashiriari-2. None of the
19 bamboo specialists identified by
Kratter were found at this site.
Kratter found that the non-bamboo
habitats used by near-obligate and
facultative specialists included early
successional vegetation, among
others. Interestingly, Neopelma
sulphureiventer and Automolus
rufipileatus were only netted in
secondary vegetation at
Cashiriari-2.
Micro-habitats
Although San Martin-3 was
relatively homogeneous in terms of
habitat, certain species were only
observed or captured or appeared to
be much more common in certain
micro-habitats. Differences in
micro-habitat preferences are likely
to occur at Cashiriari-2 as well, but
we were not there long enough to
document them.
Certain fish-eating species
were, naturally enough, observed
only along streams. The agami
heron (Agamia agami), fasciated
tiger-heron (Tigrisoma fasciatum), and
green-and-rufous kingfisher
(Chloroceryle inda) were found along
small streams within the forest,
while the green kingfisher
(Chloroceryle americana) was observed
on the larger Camisea tributary
stream. Other arboreal species were
only observed along the edges of
this larger stream, including whitethroated jacamar (Brachygalba
albogularis), gray-capped flycatcher
(Myiozetetes granadensis), and silverbeaked tanager (Ramphocelus carbo).
Buff-rumped warbler (Basileuterus
fulvicauda) foraged primarily along
the banks of small streams within
the forest.
Certain bamboo-associated
species were much more common
on the drier ridge tops where bamboo dominated and were absent or
rare in patches of broad-leafed trees
in ravines. These include Cabanis
spinetail (Synallaxis cabanisi), Manu
antbird (Cercomacra manu), whitelined antbird (Percnostola lophotes),
Goeldis antbird (Myrmeciza goeldi,)
and sulphur-bellied tyrant-manakin
(Neopelma sulphureiventer) (Appendix
4). Conversely, several other species
were more common in patches of
forest in ravines, including bluishslate antshrike (Thamnomanes
schistogynus), green manakin
(Chloropipo holochlora), and whitewinged shrike-tanager (Lanio versicolor). Band-tailed manakin (Pipra
fasciicauda) showed an interesting
pattern. Adult males were virtually
restricted to ravines, while femaleplumaged birds, possibly mostly
immatures, were much more common on the drier ridge tops. It is
possible that adult males exclude
younger birds from more favorable
habitat in ravines.
Patterns for other species were
suggestive, but our observations
were too few to make any definite
statement about their micro-habitat
preferences. Additional work in the
area is likely to document further
species differences.
Game Species
Large game birds such as
tinamous, guans, curassows, and
trumpeters are typically among the
first species to disappear in response to human activity in an area,
chiefly as a result of over-hunting.
At San Martin-3, small groups of
Spixs guan (Penelope jacquacu) were
noted regularly, and the bluethroated piping-guan (Pipile
cumanensis) was seen on three occa-
sions. The razor-billed curassow

(Mitu tuberosa) and pale-winged
trumpeter (Psophia leucoptera) were
reported in the area by other observers. We also recorded 3 species
of tinamougreat (Tinamus major), cinereous (Crypturellus cinereous), and black-capped (C.
atrocapillus). Tinamous are secretive
and generally recorded primarily by
call. Calling activity by many species is low during the late rainy
season, and it is possible that
surveys at another season would
record additional species of
tinamou.
The frequency of encounter of
th game species plus their relative
lack of wariness when encountered
indicate that hunting pressure in the
immediate area of San Martin-3 is
fairly low. A small indigenous
settlement of three houses is located approximately three hours
(walking) from the site. Evidently,
this density of human population is
too low to have had any severe
effect on local game species.
Our stay at Cashiriari-2 was too
brief to obtain any definite impression of the relative abundance of
game species. However, Spixs guan,
razor-billed curassow, and palewinged trumpeter were reported to
us by other observers. Given the
remoteness of the site, it would be
expected that game species should
be relatively common.
Large parrots, macaws, and
toucans also are liable to disappear
rapidly because of human activity,
being hunted not only for food but
also for the pet trade. As mentioned
above, these species were rare or
uncommon at San Martin-3. However, their low density at this site is
more likely to be the result of forest
structure than of hunting pressure.
The
frequency
of encounter of the
game
species
plus their
relative
lack of
wariness
when
encountered indicate that
hunting
pressure in
the immediate area
of San
Martin-3 is
fairly low.
259
Every
effort
should be
made to
see that
this exceptional avifauna does
not suffer
degradation as a
result of
future gas
development in
the area.
260
Species of Disturbed Areas

Certain bird species are characteristically associated with open
and/or disturbed sites, either
natural or those caused by human
activity. These species are rarely or
never found within extensive areas
of undisturbed forest, but sometimes may be able to colonize even
isolated disturbed sites. These
include certain species of doves,
raptors, hummingbirds, flycatchers,
wrens, seedeaters, cowbirds, and
others.
Few such species were observed at either well site. At San
Martin-3, on one occasion we
observed a little ground-tyrant
(Muscisaxicola fluviatalis) foraging at
the edge of the well site. This
terrestrial species typically is found
on river banks and islands. We also
observed a female vermilion flycatcher (Pyrocephalus rubinus), which
was present at the site over the
course of four days. This species, a
migrant from southern America, is
usually found in open areas. These
were the only two species whose
presence at San Martin-3 appeared
to be due primarily to the open area
provided by the well site. Two other
species, silver-beaked tanager
(Ramphocelus carbo) and troupial
(Icterus icterus) may be found both
along river courses and other edge
habitats and in areas disturbed by
human activity. At San Martin-3, we
observed them only along the main
stream in the area and never at the
well site.
Two cattle egrets (Bubulcus ibis)
were present at Cashiriari-2 during
the first three days of our stay, but
they subsequently left. This highly
invasive species is typically found
in pastures and open areas, feeding
on insects and other animals disturbed by grazing cattle. A roadside
hawk (Buteo magnirostris) was also

seen at the site during our stay. As
the name implies, the species is
usually found in open and disturbed
areas. These were the only species
at the site that appeared to have
been attracted to the open area, a
particularly interesting occurrence
because the former road to the area
provides a corridor of somewhat
disturbed vegetation that could
potentially facilitate colonization by
birds of open areas.
At Cashiriari-2, we also observed a pair of bat falcons (Falco
rufigularis) at the edge of the well
site every morning during our stay.
On several mornings, they engaged
in courtship displays, apparently
oblivious to the activities and noise
at the well site.
Conclusion
The avifauna of San Martin-3

and Cashiriari-2 is highly diverse,
and the species richness at these
sites may equal or exceed that of
equivalent habitats in Manu National Park, one of the richest
known sites for birds in the world.
At present, the bird community
appears to be in nearly pristine
condition, with little evidence that
human activity to date has had
significant impacts. Every effort
should be made to see that this
exceptional avifauna does not suffer
degradation as a result of future gas
development in the area.
References
Chapman, F. M. 1921. The distribution of birdlife in the Urubamba

valley of Per. Bulletin of the
Natural History Museum 7: 1-138.
Davis, T. J., C. Fox, L. Salinas,
G. Ballon, and C. Arana. 1991.
Annotated checklist of the birds
of Cusco Amazonico, Per. Occ.
Pap. Mus. Nat. Hist., Univ. Kansas
144: 1-19.
Donahue, P., T. A. Parker III, and
B. Sorrie. MS. The birds of the
Tambopata Reserve.
Karr, J. R., S. K. Robinson, J. G.
Blake, and R. O. Bierregaard, Jr.
1990. Birds of four neotropical
forests. Pages 237-269 in Four
Neotropical Rainforests. (A. H.
Gentry, ed.) Yale University Press,
New Haven.
Kratter, Andrew W. 1997. Bamboo
specialization by Amazonian birds.
Biotropica 29(1): 100-110.
Munn, Charles A. 1985. Permanent
canopy and understory flocks in
Amazonia: species composition
and population density. Pages 683712 in Neotropical Ornithology.
Ornithological Monographs 36. (P.
A. Buckley, M. S. Foster, E. S.
Morton, R. S. Ridgely, and F. G.
Buckley, eds.) American Ornithologists Union, Washington, DC.
Parker, T. A., III, and J. P. ONeill.
1980. Notes on little known birds
of the Urubamba valley, southern
Per. Auk 97: 167-76.
Parker, T. A., III, S. A. Parker, and
M. A. Plenge. 1982. An Annotated
Checklist of Peruvian Birds. Buteo
Books, Vermillion, South Dakota.
Remsen, J.V. Jr., and T. A. Parker,
III. 1983. Contribution of rivercreated habitats to bird species
diversity richness in Amazonia.
Biotropica 15: 223-31.
Robinson, S. K., and J. J. Terborgh.

1990. Bird communities of the
Cocha Cashu Biological Station in
Amazonian Per. 199-216 in Four
Neotropical Rainforests. (A. H.
Gentry, ed.) Yale University Press,
New Haven.
Servat, G. P. 1993. First records of
the Yellow Tyrannulet (Capsiempis
flaveola) in Per. Wilson Bulletin
105: 534.
Servat, G. P. 1996. An annotated
list of birds of the BIOLAT
Biological Station at Pakitza,
Per. Pages 547-568 in Manu: The
Biodiversity of Southeastern Per
( D. E. Wilson and A. Sandoval,
eds.) Editorial Horizonte, Lima.
Servat, G. P., and D. L. Pearson.
1991. Natural history notes and
records for seven poorly known
bird species from Amazonian Per.
Bulletin of the British Ornithological
Club 111: 92-95.
Stotz, D. F., J. W. Fitzpatrick, T. A.
Parker, III, and D. K. Moskovits.
1996. Neotropical Birds: Ecology
and Conservation. University of
Chicago Press, Chicago.
Terborgh, J., J. W. Fitzpatrick, and
L. Emmons. 1984. Annotated
checklist of birds and mammal
species of Cocha Cashu biological
station, Manu National Park, Per.
Fieldiana (Zoology) 21: 1-29.
Terborgh, J., S. K. Robinson, T. A.
Parker, III, C. A. Munn, and N.
Pierpont. 1990. Structure and
organization of an Amazonian
forest bird community. Ecological
Monographs 60: 213-38.
261
Appendix 1. Bird species found at San Martin-3 and Cashiriari-2. Evidence

of occurrence: C = captured; P = photographed; V = vocalization recorded;
s = sight record; v = vocalization, not recorded; + = species reported to us
by other observers at the site; * = species identification needs to be confirmed; final identification for captured species pending on comparison of
photos and other data with museum specimens and reference material; sight
records not listed for species whose occurrence was confirmed by capture
or photographs).
Species
Common Name
Tinamus major
Crypturellus cinereus
Crypturellus atrocapillus
Crypturellus variegatus
Bubulcus ibis
Agamia agami
Tigrisoma fasciatum
Cathartes melambrotus
Leucopternis albicollis
Buteo magnirostris
Spizaetus tyrannus
Daptrius americanus
Herpetotheres cachinnans
Micrastur ruficollis
Falco rufigularis
Penelope jacquacu
Aburria pipile
Mitu tuberosa
Odontophorus stellatus*
Psophia leucoptera
Columba subvinacea
Leptotila rufaxilla
Ara ararauna
Ara macao
Ara chloropterus
Ara severa
Brotogeris cyanoptera
Amazona ochrocephala
Amazona farinosa
Piaya cayana
Piaya melanogaster
Nyctiphrynus ocellatus
Hydropsalis brasiliana
262
Great Tinamou
Cinereous Tinamou
Black-capped Tinamou
Variegated Tinamou
Cattle Egret
Agami Heron
Fasciated Tiger-Heron
Greater Yellow-headed Vulture
White Hawk
Roadside Hawk
Black Hawk-Eagle
Red-throated Caracara
Laughing Falcon
Barred Forest-Falcon
Bat Falcon
Spixs Guan
Common Piping-Guan
Razor-billed Curassow
Starred Wood-Quail*
Pale-winged Trumpeter
Ruddy Pigeon
Gray-fronted Dove
Blue-and-yellow Macaw
Scarlet Macaw
Red-and-green Macaw
Chestnut-fronted Macaw
Cobalt-winged Parakeet
Yellow-crowned Parrot
Mealy Parrot
Squirrel Cuckoo
Black-bellied Cuckoo
Ocellated Poorwill
Scissor-tailed Nightjar
San Martin-3 Cashiriari-2

V
s
s
s
s
s
s
Vs
V
P
CPV
Vs
s
+
sv
+
Vs
s
v
sv
sv
v
s
v
Vs
Vs
v
CP
v
s
s
s
v
s
+
+
+
v
s
sv
s
s
s
Appendix 1. Bird species found at San Martin-3 and Cashiriari-2 (Cont.).

Species
Common Name
Streptoprocne zonaris
Threnetes leucurus
Phaethornis superciliosus
Phaethornis hispidus
Phaethornis koepckeae
Phaethornis ruber*
Eutoxeres condamini
Campylopterus largipennis
Florisuga mellivora
Klais guimeti
Thalurania furcata
Heliodoxa aurescens
Trogon melanurus
Chloroceryle americana
Chloroceryle inda
Baryphthengus martii
Brachygalba albogularis
Galbula cyanescens
Nystalus striolatus
Malacoptila semicincta
Nonnula ruficapilla
Monasa flavirostris
Capito niger
Eubucco richardsoni
Aulacorhynchus prasinus
Pteroglossus castanotis
Selenidera reinwardtii
Ramphastos cuvieri
Picumnus aurifrons
Picumnus rufiventris
Melanerpes cruentatus
Veniliornis passerinus
Celeus spectabilis
Dryocopus lineatus
Campephilus melanoleucos
Dendrocincla fuliginosa
Deconychura longicauda
Glyphorynchus spirurus
Xiphocolaptes promeropirhynchus
Dendrocolaptes picumnus
Xiphorhynchus ocellatus
Xiphorhynchus spixii
Campylorhamphus trochilirostris*
Synallaxis cabanisi
White-collared Swift
Pale-tailed Barbthroat
Long-tailed Hermit
White-bearded Hermit
Koepckes Hermit
Reddish Hermit*
Buff-tailed Sicklebill
Gray-breasted Sabrewing
White-necked Jacobin
Violet-headed Hummingbird
Fork-tailed Woodnymph
Goulds Jewelfront
Black-tailed Trogon
Green Kingfisher
Green-and-rufous Kingfisher
Rufous Motmot
White-throated Jacamar
Bluish-fronted Jacamar
Striolated Puffbird
Semicollared Puffbird
Rufous-capped Nunlet
Yellow-billed Nunbird
Black-spotted Barbet
Lemon-throated Barbet
Emerald Toucanet
Chestnut-eared Aracari
Golden-collared Toucanet
Cuviers Toucan
Bar-breasted Piculet
Rufous-breasted Piculet
Yellow-tufted Woodpecker
Little Woodpecker
Rufous-headed Woodpecker
Lineated Woodpecker
Crimson-crested Woodpecker
Plain-brown Woodcreeper
Long-tailed Woodcreeper
Wedge-billed Woodcreeper
Strong-billed Woodcreeper
Black-banded Woodcreeper
Ocellated Woodcreeper
Spixs Woodcreeper
Red-billed Scythebill*
Cabanis Spinetail
s
CP
CP
CP
CP
CP
CP
CP
CP
s
s
CP
CPV
s
CP
Vs
CP
CP
Vs
sv
s
v
Vs
CP
s
CP
s
s
s
CP
CP
CP
CP
CP
CP
CP
s
C
CP
C
CP
C
C
CP
C
C
C
v
C
C
C
C
sv
s
s
C
CP
C
C
CP
263

Species
Common Name
Synallaxis cherriei
Cranioleuca gutturata
Hyloctistes subulatus
Simoxenops ucayalae
Automolus ochrolaemus
Automolus dorsalis
Automolus infuscatus
Automolus rubiginosus
Automolus rufipileatus
Sclerurus mexicanus
Sclerurus albigularis
Sclerurus caudacutus
Xenops tenuirostris*
Xenops minutus
Cymbilaimus lineatus
Cymbilaimus sanctaemariae
Frederickena unduligera
Taraba major
Thamnophilus aethiops
Thamnophilus schistaceus
Thamnomanes ardesiacus
Thamnomanes schistogynus
Myrmotherula brachyura*
Myrmotherula hauxwelli
Myrmotherula leucophthalma
Myrmotherula ornata
Myrmotherula axillaris
Myrmotherula longipennis
Myrmotherula menetriesii
Microrhopias quixensis
Drymophila devillei
Cercomacra cinerascens
Cercomacra serva
Cercomacra manu
Myrmoborus leucophrys
Myrmoborus myotherinus
Hypocnemis cantator
Sclateria naevia
Percnostola lophotes
Schistocicla leucostigma
Myrmeciza hemimelaena
Myrmeciza goeldii
Myrmeciza fortis
Rhegmatorhina melanosticta
Hylophylax naevia
264
Chestnut-throated Spinetail
Speckled Spinetail
Striped Woodhaunter
Peruvian Recurvebill
Buff-throated Foliage-gleaner
Crested Foliage-gleaner
Olive-backed Foliage-gleaner
Ruddy Foliage-gleaner
Chestnut-crowned Foliage-gleaner
Tawny-throated Leaftosser
Gray-throated Leaftosser
Black-tailed Leaftosser
Slender-billed Xenops*
Plain Xenops
Fasciated Antshrike
Bamboo Antshrike
Undulated Antshrike
Great Antshrike
White-shouldered Antshrike
Plain-winged Antshrike
Dusky-throated Antshrike
Bluish-slate Antshrike
Pygmy Antwren*
Plain-throated Antwren
White-eyed Antwren
Ornate Antwren
White-flanked Antwren
Long-winged Antwren
Gray Antwren
Dot-winged Antwren
Striated Antbird
Gray Antbird
Black Antbird
Manu Antbird
White-browed Antbird
Black-faced Antbird
Warbling Antbird
Silvered Antbird
White-lined Antbird
Spot-winged Antbird
Chestnut-tailed Antbird
Goeldis Antbird
Sooty Antbird
Hairy-crested Antbird
Spot-backed Antbird
CP
CP
CPV
CPV
CPV
CPV
CP
CP
CP
CP
CP
Vs
CP
CPV
CP
CPV
CP
CPV
s
s
C
CP
C
CP
C
C
CV
C
C
s
C
CP
CP
CP
CP
C
CP
C
C
Vs
CPV
CPV
CPV
C
CP
CPV
s
CPV
s
CPV
CPV
CP
CP
sv
CP
C
CP

Species
Common Name
Hylophylax poecilonota
Scale-backed Antbird
Phlegopsis nigromaculata
Black-spotted Bare-eye
Formicarius analis
Black-faced Antthrush
Hylopezus berlepschi
Amazonian Antpitta
Myrmothera campanisona
Thrush-like Antpitta
Conopophaga Peruviana
Ash-throated Gnateater
Liosceles thoracicus
Rusty-belted Tapaculo
Iodopleura isabellae
White-browed Purpletuft
Lipaugus vociferans
Screaming Piha
Cotinga maynana
Plum-throated Cotinga
Querula purpurata
Purple-throated Fruitcrow
Chloropipo holochlora
Green Manakin
Pipra fasciicauda
Band-tailed Manakin
Pipra chloromeros
Round-tailed Manakin
Pipra coronata
Blue-crowned Manakin
Neopelma sulphureiventer
Sulphur-bellied Tyrant-Manakin
Mionectes olivaceus
Olive-striped Flycatcher
Mionectes oleagineus
Ochre-bellied Flycatcher
Mionectes macconnelli
McConnells Flycatcher
Leptopogon amaurocephalus
Sepia-capped Flycatcher
Poecilotriccus albifacies
White-cheeked Tody-Tyrant
Hemitriccus flammulatus
Flammulated Bamboo-Tyrant
Corythopis torquata
Ringed Antpipit
Zimmerius gracilipes
Slender-footed Tyrannulet
Ramphotrigon megacephala
Large-headed Flatbill
Platyrinchus coronatus
Golden-crowned Spadebill
Platyrinchus platyrhynchos
White-crested Spadebill
Myiophobus fasciatus
Bran-colored Flycatcher
Myiobius erythrurus
Ruddy-tailed Flycatcher
Myiobius atrocaudatus
Black-tailed Flycatcher
Myiobius barbatus
Sulphur-rumped Flycatcher
Lathrotriccus euleri
Eulers Flycatcher
Contopus borealis
Olive-sided Flycatcher
Vermilion Flycatcher
Pyrocephalus rubinus
Little Ground-Tyrant
Muscisaxicola fluviatilis
Long-tailed Tyrant
Colonia colonus
Bright-rumped Attila
Attila spadiceus
Empidonomus aurantioatrocristatu Crowned Slaty-Flycatcher
Gray-capped Flycatcher
Myiozetetes granadensis
Masked Tityra
Tityra semifasciata
Tawny-crowned Greenlet
Hylophilus ochraceiceps
White-necked Thrush
Turdus albicollis
Thrush-like Wren
Campylorhynchus turdinus
Moustached Wren
Thryothorus genibarbis
Southern Nightingale-Wren
Microcerculus marginatus
CPV
V
CP
CPV
CP
CPV
s
s
V
CP
CP
CP
CP
CP
C
CP
s
CP
CP
s
Vs
CP
CP
CP
CP
CP
s
s
s
s
v
s
s
s
CP
CP
Vs
CPV
CPV
C
C
C
v
V
CP
C
C
C
C
C
C
C
C
C
C
C
C
C
C
C
265

Species
Common Name
San Martin - 3 Cashiriari - 2
Ramphocaenus melanurus
Long-billed Gnatwren
Basileuterus chrysogestee
Golden-bellied Warbler
Basileuterus fulvicauda
Buff-rumped Warbler
Arremon taciturnus
Pectoral Sparrow
Cissopis leveriana
Magpie Tanager
Lanio versicolor
White-winged Shrike-Tanager
Tachyphonus rufiventer
Yellow-crested Tanager
Chlorothraupis carmioli
Olive Tanager
Habia rubica
Red-crowned Ant-Tanager
Ramphocelus carbo
Silver-beaked Tanager
Thraupis palmarum
Palm Tanager
Euphonia xanthogaster
Orange-bellied Euphonia
Tangara mexicana
Turquoise Tanager
Tangara chilensis
Paradise Tanager
Tangara schrankii
Green-and-gold Tanager
Tangara xanthogastra
Yellow-bellied Tanager
Tangara nigrocincta
Masked Tanager
Tangara cyanicollis
Blue-necked Tanager
Dacnis lineata
Black-faced Dacnis
Dacnis flaviventer
Yellow-bellied Dacnis
Dacnis cayana
Blue Dacnis
Pitylus grossus
Slate-colored Grosbeak
Saltator maximus
Buff-throated Saltator
Cyanocompsa cyanoides
Blue-black Grosbeak
Psarocolius bifasciatus yuracares Olive Oropendola
Psarocolius decumanus
Crested Oropendola
Psarocolius angustifrons
Russet-backed Oropendola
Cacicus cela
Yellow-rumped Cacique
Icterus cayanensis
Epaulet Oriole
Icterus icterus
Troupial
Total = 198
266
CP
s
CP
CP
s
CP
s
CP
s
s
CP
s
s
s
s
s
CP
CP
CP
s
s
Vs
s
sv
164
C
CP
C
s
s
s
s
s
s
s
C
C
s
s
s
98
Appendix 2. Mist-netting data from San Martin-3 (species ranked by the

number of individuals captured).
Species
Pipra fasciicauda
Phlegopsis nigromaculata
Hylophylax naevia
Myrmeciza hemimelaena
Campylorhamphus sp.
Myrmeciza goeldii
Basileuterus fulvicauda
Myrmotherula ornata
Dendrocincla fuliginosa
Xenops minutus
Thalurania furcata
Myiobius atricaudus
Automolus dorsalis
Thamnomanes schistogynus
Lanio versicolor
Myrmotherula leucophthalma
Pitylus grossus
Terenotriccus erythrurus
Habia rubica
Simoxenops ucayalae
Eutoxeres condamini
Saltator maximus
Threnetes leucurus
Glyphorhynchus spirurus
Galbula cyanescens
Corythopis torquata
Captures
26
19
14
14
11
10
10
9
9
9
8
7
7
7
7
6
6
6
6
6
5
5
5
5
5
5
5
5
5
5
4
4
4
4
4
4
4
4
4
3
3
3
3
3
Recaptures Total Percent of total

12
7
8
4
4
1
0
1
1
2
1
3
4
5
0
1
4
1
0
0
0
0
0
1
0
0
0
0
1
0
1
1
1
0
6
0
0
0
0
0
3
0
0
0
38
26
22
18
15
11
10
10
10
11
9
10
11
12
7
7
10
7
6
6
5
5
5
6
5
5
5
5
6
5
5
5
5
4
10
4
4
4
4
3
6
3
3
3
8.6
5.9
5
4.1
3.4
2.5
2.3
2.3
2.3
2.5
2
2.3
2.5
2.7
1.6
1.6
2.3
1.6
1.4
1.3
1.1
1.1
1.1
1.4
1.1
1.1
1.1
1.1
1.4
1.1
1.1
1.1
1.1
0.9
2.3
0.9
0.9
0.9
0.9
0.7
1.4
0.7
0.7
0.7
267
Appendix 2. Mist-netting data from San Martin-3 (Cont.).

Species
Captures
Turdus albicollis
Pipra chloromeros
Hypocnemis cantator
Arremon taciturnus
Sclerurus mexicanus
Polyplancta aurescens
Hyloctistes subulatus
Myiobius barbatus
Synallaxis cabanisi
Myrmeciza fortis
Mionectes maconnellii
Chloroceryle inda
Myrmothera campanisona
Frederickena unduligera
Phaethornis cf. ruber
Liosceles thoracicus
Baryphthengus ruficapillus
Dendrocolaptes picumnis
Rhamphocaenus melanurus
Cercromacra manu
Hylopezus berlepschi
Mionectes olivaceus
Nonnula ruficapilla
Micrastur ruficollis
Taraba major
Synallaxis cherriei
Sclerurus albigularis
Venilovis passerinus
Thamnophilus aethiops
Cercromacra serva
Total species = 86
268
3
3
3
3
3
2
2
2
2
2
2
2
2
2
2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
360

0
1
0
0
0
0
2
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
3
4
3
3
3
2
4
2
2
2
2
2
2
2
3
1
2
1
1
1
1
1
1
1
1
2
1
2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
440
0.7
0.9
0.7
0.7
0.7
0.5
0.9
0.5
0.5
0.5
0.5
0.5
0.5
0.5
0.7
0.2
0.5
0.2
0.2
0.2
0.2
0.2
0.2
0.2
0.2
0.5
0.2
0.5
0.2
0.2
0.2
0.2
0.2
0.2
0.2
0.2
0.2
0.2
0.2
0.2
0.2
0.2
100
Appendix 3. Mist-netting data from Cashiriari-2 (species ranked by the

number of individuals captured).
Species
Phaethornis koepckeae
Glyphorhynchus spirurus
Pipra coronata
Pipra chloromeros
Chlorothraupis carmioli
Mionectes olivaceus
Turdus albicollis
Hylophylax naevia
Myrmoborus myotherinus
Threnetes leucurus
Thalurania furcata
Platyrinchus coronata
Eutoxeres condamini
Conopophaga Peruviana
Deconychura longicauda
Myrmotherula menetriesii
Terenotriccus erythrurus
Saltator maximus
Hylophylax poecilonota
Mionectes oleagineus
Myrmotherula hauxwelli
Selenidera reinwardtii
Xiphorhynchus spixii
Myiobius barbatus
Formicarius analis
Lanio versicolor
Florisuga mellivora
Myrmeciza fortis
Automolus infuscatus
Captures
16
16
14
13
10
9
8
8
8
6
6
6
5
5
5
5
4
4
3
3
3
3
3
3
3
3
3
2
2
2
2
2
2
2
2
2
2
2
2
2
2
1
1
1

5
3
1
1
0
0
5
0
0
2
0
1
0
0
0
1
2
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
21
19
15
14
10
9
13
8
8
8
6
7
5
5
5
6
6
5
3
3
3
3
3
3
3
3
3
2
2
2
2
2
2
2
2
2
3
2
2
2
2
1
1
1
8.5
7.7
6
5.6
4
3.6
5.2
3.2
3.2
3.2
2.4
2.8
2
2
2
2.4
2.4
2
1.2
1.2
1.2
1.2
1.2
1.2
1.2
1.2
1.2
0.8
0.8
0.8
0.8
0.8
0.8
0.8
0.8
0.8
1.2
0.8
0.8
0.8
0.8
0.4
0.4
0.4
C
P
G
P
P
C
M
N
C
Tu
H
Th
M
Th
Th
C
P
Sc
M
P
E
C
D
M
Te
S
H
M
M
M
S
269
Appendix 3. Mist-netting data from Cashiriari-2 (Cont.).

Species
Captures
Platyrinchus platyrhynchos
Xenops minutus
Attila spadiceus
Klais guimetii
Capito niger
Cymbilaimus lineatus
Polyplancta aurescens
Corythopis torquata
Galbula cyanescens
Xenops tenuirostris
Total species = 62
270
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
224

0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
24
0.4
0.4
0.4
0.4
0.8
0.4
0.4
0.4
0.4
0.4
0.4
0.4
0.4
0.4
0.4
0.4
0.4
0.4
100
1
1
1
1
2
1
1
1
1
1
1
1
1
1
1
1
1
1
248
Appendix 4. Birds associated with Guadua bamboo thickets in southeastern Per (San Martin = this study, Kratter (1997) = data from the Ro
Tambopata, Servat (1996) = Pakitza on the Ro Manu; Column codes:
Kratter (1997)O = obligate bamboo specialist, N = near-obligate bamboo specialist, F = facultative bamboo specialist, c = common in bamboo
thickets, r = apparent bamboo specialist, but too rare to determine exact
status; Servat (1996)X = species found exclusively in bamboo at Pakitza,
x = species found exclusively in bamboo at Pakitza, but found in other
habitats at other sites in the region; Stotz et al. (1996)X = indicator
species for bamboo habitat in southern Amazonia region with ranges that
extend to southeastern Per).
Species
Crypturellus atrocapillus
Dromococcyx pavoninus
Chlorostilbon mellisuga
Bucco macrodactylus
Monasa flavirostris
Nonnula ruficapilla
Celeus spectabilis
Campylorhamphus trochilirostris
Synallaxis cabanisi
Synallaxis cherriei
Simoxenops ucayalae
Automolus dorsalis
Automolus melanopezus
Cymbalaimus sanctaemariae
Myrmotherula ornata
Myrmotherula iheringi
Myrmotherula obscura
Drymophila devillei
Cercomacra manu
Cercomacra nigrescens
Hypocnemis cantator
Myrmeciza goeldii
Lophotriccus eulophotes
Poecilotriccus albifacies
Ramphotrigon megacephala
San Martin-3
Kratter
c
r
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
r
r
r
F
N
F
F
N
N
N
c
r
N
F
F
F
O
O
c
c
N
N
O
O
N
Servat
X
X
X
X
Stotz
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
271
Appendix 4. Birds associated with Guadua bamboo thickets (Cont.).

San Martin-3
Species
Ramphotrigon fuscicauda
Casiornis rufa
Capsiempis flaveola
Machaeropterus pyrocephalus
272
X
X
Kratter
N
c
c
Servat
Stotz
X
X
X
X
X
Long-term
Monitoring
of Bird Fauna
in the Lower
Urubamba
Region
James Siegel
U.S. Fish and Wildlife Service,
National Education and
Training Center
George Angehr
Smithsonian Tropical
Research Institute
Introduction
Although tropical bird communities are typified by high species

richness, many species are rare,
making them vulnerable to local
extinction caused by human activity. Part of the high diversity of
these communities is due to the
high habitat heterogeneity suitable
for many species. Many species are
found in only one or a few habitats
or micro-habitats, and are unable to
survive if that habitat is altered or
degraded.
Some impacts of human activity are obvious. When a tropical
forest is cut and replaced by agriculture or other land uses, most of the
forest birds originally found there
will disappear. In the neighborhood

of human settlements, game species
and those popular in the pet trade
may be affected directly by overhunting or capture. Other effects
may be more subtle. For example,
Canaday (1997) found a decrease in
the number of insectivorous forest
birds within forest that was intact
but in the vicinity of large clearings.
Forest fragmentation (i.e., division
of previously contiguous forest into
smaller blocks) may result in drastic
losses in species diversity (Willis
1974, Karr 1982, Bierregaard and
Lovejoy 1989, Bierregaard et al.
1992). Cleared areas within forest,
particularly if connected to larger
clearings by continuous strips of
disturbed habitat such as those
along roads, could potentially
facilitate colonization of an area by
predators, parasites, and competitors.
Here, we discuss potential
impacts on local bird communities
stemming from current and future
activities by Shell Prospecting and
Development (Per) B.V. in the
Lower Urubamba region. We also
describe various methods that may
273
be used to monitor these impacts. In

the final section we present a
prioritized long-term monitoring
plan for the area.
Current Conditions
There is
considerable noise
by heavy
equipment
and aircraft
at both
sites, but
we saw no
evidence
that this
had
significant
impacts on
the behavior of birds
in the area.
274
We visited the Lower

Urubamba study area from late
April to early May, 1997. At that
time, approximately 3.5 ha at the 5month old San Martin-3 well site
had been cleared, and the site was
being prepared for well installation
with heavy equipment. A similar
area at Cashiriari-2 had been also
been cleared, and the well there had
been in operation for several
months.
Cashiriari-2 was initially developed approximately 10 years ago. At
that time, it connected by road to
the villages of Shivankoreni and
Segakiato. These roads are currently
overgrown, although the areas of
second-growth along their length
could potentially provide a route of
colonization for bird species associated with disturbed habitats. There
is no current or former road access
to San Martin-3.
At the time of our visit, there
were no obvious changes in the bird
fauna that could be attributed to
the relatively small clearings at San
Martin-3 and Cashiriari-2. A few
vagrant or migrant individuals of
open-country species were present,
but these did not appear to have
established resident populations
(Angehr and Aucca this volume).
Eventually, colonization by some
early successional species and forest
edge species may be anticipated. As
grass and brush fill in the edges and
bare ground of the site, the area
may become suitable habitat for
certain open-area seed eaters and
insect hawkers.
Game species such as guans,

curassows, and others were relatively common at San Martin-3.
Although we visited Cashiriari-2 for
only a brief time, game species
appeared to be reasonably common
at this site as well (Angehr and
Aucca this volume). Given the
apparently low population density
of indigenous people in the area
and the companys prohibition on
hunting by workers at the sites,
these species may be at or close to
their natural densities. Although
macaws and large parrots were
relatively uncommon, there is no
evidence that this was due to
human activity.
There is considerable noise by
heavy equipment and aircraft at
both sites, but we saw no evidence
that this had significant impacts on
the behavior of birds in the area. To
the contrary, a pair of bat falcons
was observed at Cashiriari carrying
out courtship activities within
about 30 m of the well itself.
Sedimentation as the result of
construction and other development activities or the release of
formation water or chemicals into
local streams could potentially
affect species such as kingfishers or
herons or other species that rely on
aquatic prey. Sedimentation or
pollution could also affect insects
dependant on streams that are fed
on by insectivorous birds. We did
not observe such impacts as the
result of current activities at San
Martin-3 or Cashiriari-2.
Sampling by Angehr and Aucca
(this volume) was not designed to
detect more subtle effects on the
bird community, effects such as a
decrease in the numbers of insectivores in the vicinity of clearings as
described by Canaday (1997).
Current activities at the sites are
not resulting in any forest fragmentation, so we do not anticipate any

impacts on the bird community
from this cause.
Future
Developments and
Land-Management
Implications
More significant impacts may

be expected if the Lower Urubamba
is developed for full-scale gas
production. Additional well sites
will be cleared, and a pipeline will
be installed. Installation of the
pipeline will create a corridor of
disturbed habitat that may allow
predators, nest parasites (e.g.,
cowbirds), or competitors to colonize the well sites, probably during
the revegetation process to primary
forest.
The most severe long-range
effects may be expected if a permanent road is constructed to the well
sites. We understand one alternative
under consideration by Shell Prospecting and Development (Per)
B.V. is to install a buried pipeline
and maintain no permanent road
access to the well sites. We agree
with their policy and discuss below
the potential impacts in case this
alternative is not pursued.
While the Lower Urubamba gas
project itself is not designed to
increase access to forest resources
and to potentially arable land, that
may well be the most significant
environmental impact of road
construction: the opening of formerly inaccessible tropical forest to
increasing levels of resource use
and clearing for agriculture and
other forms of land use. Pipeline
right-of-ways, trails, and roads
constructed for the project would
likely increase access to the area by

both local people and newcomers
seeking natural resources (fish and
game, timber, fruits, nuts, resins,
medicinal plants, and minerals) and
unoccupied land. Increased access
would result in greater exploitation
of these forest resources and increased clearing of the land for
subsistence and cash crops, with
significant changes in biodiversity
associated with forest fragmentation
and habitat loss. Thus, controlling
access to the region on the roads,
trails, and pipelines will be a crucial
management action in conserving
biodiversity in the area. Changes in
the natural environment because of
gas well platforms and pipelines
may be ultimately less important
than the environmental impacts that
increased forest access brings to the
region.
The project will raise expectations of local people and new
settlers, who will likely demand that
gas development bring tangible
benefits to the local communities.
These might include electricity,
arable land, timber-based industry,
ecotourism, and other activities
based on natural resources that will
have impacts on biodiversity. Such
impacts are difficult to quantify at
this initial stage of the project.
Objectives of the
Long-Term
Monitoring Program
It is not feasible to monitor all
conceivable impacts of development on bird populations in the
project area. For the purposes of
this manuscript, we limit our discussion to the impacts of the
minimal expected future development in the area if full-scale gas
production (i.e., installation of
275
permanent production wells, their

long-term operation, and the construction of an underground gas
pipeline with subsequent reforestation of the pipeline route) is initiated. We make the assumptions that
no permanent access road will be
built into the area, that the company or other authorities will continue to control access to the area,
and that spontaneous, uncontrolled
colonization will not take place. In
the event that these assumptions
prove false, additional monitoring
programs would be necessary to
examine the effects of forest fragmentation or of widespread deforestation.
Given these assumptions, the
major impacts of development will
be those caused by the presence of
the well sites and the pipeline. The
actual area of forest that would be
removed for each well site and the
pipeline is small in comparison to
the entire project area, so the direct
effect of this deforestation on bird
populations in the area should also
be small.
However, the possible indirect
effects of creating the clearings
must also be taken into account. It
is possible that the presence of
these clearings within the forest
could affect bird populations over a
much wider area. The most relevant
study in relation to this issue is that
of Canaday (1997), who investigated the impact of petroleum
development in the Cuyabeno
region of Ecuador. Canaday surveyed birds within four zones
defined by their distances from
large clearings, roads, and small
clearings: 1) coffee plantations
within 100 m of uncut forest, 2)
edge forest or uncut forest with
300 m of roads and 100 m of the
contiguous clearings bordering the
276
roads, 3) intermediate forest or

uncut forest 2 km from roads and
about 200 m from small clearings,
and 4) interior forest or uncut
forest at least 3.5 km from roads
and one km from any clearings.
Canaday found that the number of
insectivorous bird species varied
significantly among the four zones,
with forest near clearings having
fewer species than the interior
forest. In contrast, the number of
species of frugivores and omnivores
did not differ significantly. Canaday
was not able to identify the specific
causes of these differences, but
proposed that several factors could
be involved: 1) micro-climatic
changes affecting the arthropod
prey base, 2) greater habitat sensitivity by insectivores as a result of
the high degree of ecological specialization of many of these species, 3) changes in predation, and 4)
competition from opportunistic,
disturbance-adapted omnivores.
Canaday did not indicate how long
the roads and clearings he studied
had been in existence, but the
Cuyabeno region has been undergoing petroleum development for
more than 20 years. It is possible
that the decline in the number of
insectivorous species near clearings
described by Canaday has taken
place over a period of many years.
Given these results, a major
objective of the monitoring program in the Lower Urubamba
should be the detection of such
edge effects in the vicinity of the
well-sites and along the pipeline
route and at least an approximate
estimation of the distance they
extend from artificial clearings. If
edge effects extend for a considerable distance from clearings, they
could have an impact on bird
populations over a significant area

of forest.
At present, Shell Prospecting
and Development (Per) B.V. is
controlling hunting near the well
sites by banning the importation of
firearms into the project area by
company personnel. Once a pipeline is constructed, controlling
access to the project area by hunters will be more difficult, even if
the pipeline route is reforested.
Hunters from settled areas may be
able to obtain easier access to
remote areas of the forest by entering along the pipeline route. Therefore monitoring of game bird
species along the pipeline route
would be desirable. The company
may be able to control such hunting
by instituting patrols along the
pipeline route. An assessment of
the effectiveness of this control
could be obtained through a monitoring program.
Indigenous hunters and some
environmental groups have speculated that noise and human activities at the well sites might adversely
affect the forest fauna in the area,
possibly by frightening animals
away. Based on our observations in
the Lower Urubamba and other
tropical forest areas, we do not
believe that noise and human
activity in the absence of hunting,
other direct persecution, and habitat alteration are likely to have a
major long-term effect on the forest
fauna. However, this question could
be examined by instituting studies
of game species at the gas well sites
and along the pipeline.
Monitoring
Methodologies
Birds species differ widely in

their foraging zones, population
densities, ease of detection, daily

and seasonal activity patterns, and a
host of other factors. Because of
this variability, no single monitoring
methodology is capable of assessing
all species with equal accuracy and
efficiency. A wide variety of methodologies for monitoring bird
populations exists (see Verner
1985), but trade-offs must be
considered between the percentage
of bird community sampled, costs,
the degree of training needed, and
other considerations. Because
different methodologies may be
most efficient at sampling different
parts of the community, using a
combination of techniques is
generally the best approach to
assessing the community as a
whole.
We consider three widely-used
techniquesmist netting, point
counts, and transectsfor assessing
bird populations to be the most
feasible methods in remote areas
such as the Lower Urubamba.
Certain other techniques (e.g., spot
mapping) may yield more detailed
information, but they are highly
labor-intensive and require a very
high level of training.
Mist-netting surveys consist of
operating an array of mist-nets in
an area over a period of one to
several days. Advantages include: 1)
accuracy of the data collected is
relatively high; 2) relatively little
training is necessary for field personnel because identification
manuals can be consulted while the
bird is in hand; 3) data can be
collected on the physical condition
of the birds (very difficult to accomplish with other methods); 4) in
long-term programs, the recapture
frequency of marked birds provides
data on annual survivorship, productivity, and other demographic
We
consider
mist netting, point
counts, and
transects to
be the most
feasible
methods
for assessing bird
populations in the
Lower
Urubamba.
277
Figure 1. Example of mist net set-up.

parameters; and 5) secretive and
inconspicuous species may be
detected. Disadvantages include: 1)
mist netting is relatively labor
intensive; 2) it samples only a part
of the community, primarily understory species between 5 and 100 g
in body weight; and 3) because
capture rates depend strongly on
foraging patterns and behavior, mist
netting can not provide accurate
information on true population
densities or be used to make comparisons of the abundances of
different species (see Fig. 1).
The point-count technique
allows an observer to count all
individual birds seen or heard from
a fixed point within a standard
period of time, usually 8 to 10
minutes. Advantages include: 1) it
is much less labor intensive than
mist netting; 2) it samples a much
larger percentage of the bird community than mist-netting; and 3)
estimates of population density can
278
be obtained, which is not possible

with mist-netting. Disadvantages
include: 1) a much higher level of
training is necessary than with mist
netting, 2) surveys by different
observers may not be readily comparable, 3) no information is obtained on physical condition or on
demography, and 4) surveys must
be carried out during a certain part
of the yearthe breeding season,
when most species are vocalizing.
Recommendations
for Long-Term
Monitoring
In this section we outline the

elements of a monitoring program
for birds in the Lower Urubamba
region. Because funds to carry out
such a program may be limited, we
also assign priorities to the different
components within the program.
These priorities are based on the
costs versus expected benefits of
each component, as well as the

relative probability of the impacts
each component is designed to
monitor.
As discussed above, the primary objectives of a monitoring
program should be assessment of
impacts caused by edge effects
related to clearing at the well sites
and construction of the pipeline
and assessment of possible declines
in game birds and other hunted
species because of increased access
by hunters coming from settlements.
Impacts due to the first cause
might be expected at both the well
sites and along the pipeline route; a
monitoring program should be set
up in both areas. Impacts due to the
second cause would be expected
primarily along the pipeline route;
surveys of game birds would be a
high priority only in this area.
Establish Control Sites
Bird populations show natural
fluctuations depending on changing
environmental conditions such as
variations in rainfall, insect populations, and the fruiting of forest
vegetation. Monitoring bird populations at the well sites and along
pipelines over a multiple-year
period and comparing the results to
a preliminary survey may reveal
changes in the species complement
and in the relative abundance of
species over time. These changes
may be attributed to the effects of
gas developments. But the comparison may be strongly confounded by
other environmental factors unrelated to the development activities.
Monitoring at sites near the gas
developments as well as at control
sites established beyond the potential influence zone of the wells or
pipeline route is the best way to
separate changes in bird populations
caused by edge effects from those

caused by natural environmental
changes. The control plots should
be located several km away from
the wells and pipelines. A high
priority is to establish several small
base camps away from the well sites
or the pipeline route to facilitate
carrying out surveys at the control
sites.
Assessment of impacts at the
control sites may be carried out by
either mist-net surveys or point
counts, or preferably both. We
consider a mist-net survey to be of
higher priority than point counts
because it requires less intensive
training and because it will be easier
to replicate surveys over a longterm study even if there is a change
in personnel.
Assessment
of impacts
at the
control sites
may be
carried out
by either
mist-net
surveys or
point
counts.
Priorities for Monitoring

In summary, we recommend the
following priorities for the monitoring program:
A. Well Sites
Initially, monitoring should
include San Martin-3 and
Cashiriari-2. Additional sites could
be included if new wells are established in substantially different
habitat types.
Priorities:
* Establish a system or six
survey areas at each well site to
include two control sites and corresponding base camps.
these survey areas to assess edge
effects.
* Initiate a point-count survey,
also to assess edge effects.
species.
279
B. Pipeline Route
Monitoring should be carried
out at a minimum of two sites, one
within a few km of the point where
the pipeline starts and 1 within
intact forest.
Priorities:
* Establish a system of three
survey areas at each pipeline study
site that includes 1 control area and
corresponding base camp.
these areas to assess edge effects.
species.
* Initiate a point-count survey
to assess edge effects.
C. Training
A two- to three-week orientation and training course conducted
by expert ornithologists from Per
and elsewhere will begin the process
of creating a cadre of experience
field workers for the Lower
Urubamba project and related bird
monitoring programs. The course
should include instruction in basic
bird identification skills (both sight
and sound), mist netting, point
counts, and transects approaches.
We recommend a class of 10 to15
university students, post-graduate
ecologists, and local community
participants for the first year of
monitoring. Each year thereafter,
we recommend that a new class be
trained with assistance from the
most adept graduates and field
workers from earlier courses. In
addition to providing needed expertise and assistance for the Lower
Urubamba project, the course will
help the National Museum of Per
and other ecological and conservation organizations in advancing the
280
environmental knowledge, skills,

and capabilities of the Peruvian
scientific community and local
people.
References Cited
Bierregaard, R. O., Jr., and T. E.

Lovejoy. 1989. Effects of fragmentation on Amazonian understory bird communities. Acta
Amazonica 19: 215-241.
Bierregaard, R. O., Jr., T. E.
Lovejoy, V. Kapos, A. A. dos
Santos, and R. W. Hutchings.
1992. The biological dynamics of
tropical forest fragments. BioScience
42: 859-866.
Canaday, C. 1997. Loss of insectivorous birds along a gradient of
human impact in Amazonia.
Biological Conservation 77: 63-77.
Karr, J. R. 1982. Avian extinction
on Barro Colorado Island, Panama:
a reassessment. American Naturalist
119: 220-239.
Verner, J. 1985. Assessment of
counting techniques. Current
Ornithology 2: 247-302.
Willis, E. O. 1974. Populations and
local extinctions of birds on Barro
Colorado Island, Panama. Ecological
Monographs 44: 153-169.
Small,
Non-volant
Mammals:
Biodiversity
Assessment
in the Lower
Urubamba
Region
Sergio Solari and
Juan Jos Rodriguez
Mayor de San Marcos
Introduction
The Lower Urubamba region is

part of the Amazon rainforest. The
Amazon Basin is one of the most
highly diverse areas in the world in
relation to mammals (Voss and
Emmons 1996). Traditionally,
mammals are divided into groups
for study: small-non volant mammals (e.g., marsupials and rodents
this paper), volant mammals (e.g.,
bats, Wilson et al. this volume) and
medium and large mammals (e.g.,
agouti, deer, jaguars, Boddicker this
volume). We studied the community
of small, non-flying mammals,
which are estimated constitute
between 15% and 25% of the

mammal fauna in tropical rain forest
(Voss and Emmons 1996).
Studies of small, non-flying
mammals have been done in other
tropical areas of Per. This study
will be compared to those in
Pakitza, Cusco Amazonico, and
Balta, all of which are near our
study site (Fig. 1). The diversity
found at these locations confirmed
the previous estimates of the
mammal diversity in Amazonia
(Woodman et al. 1991, Patton et al.
1994, Pacheco and Vivar 1996,
Voss and Emmons 1996).
Because the Lower Urubamba
study area is very hilly with many
streams, its habitats have rarely
been sampled. Thus, their biological
diversity is not well known, and the
particular aspects of the species
composition have not been treated
properly. As part of the Lower
Urubamba biodiversity assessment,
sampling of small, non-flying
mammals was conducted at San
Martin-3 and Cashiriari-2, where
(Per) B.V. is exploring for natural
gas.
281
Figure 1. Map of southeastern Per, showing the Urubamba

location and three other sites used for comparison.
Methods
Sampling was done at San

Martin-3 from May 2 -14 and at
Cashiriari-2 from May 16 - 28,
1997, at the end of the rainy season. During this time, heavy rains
and abnormally low ambient temperatures prevailed.
At San Martin-3, we set a series
of traps parallel to the narrow,
muddy trial that goes east of the
camp to sample the mammal fauna.
We also sampled in a wide stream
that crosses biodiversity plot #1
west of the camp in a relatively flat
area (see Alonso et al. this volume).
The terrain at Cashiriari-2 is flatter
than that of San Martin-3. At
Cashiriari-2, we sampled south of
the drilling platform in a flat area
where the proposed biodiversity
plot #4 will be established. An old
road with secondary vegetation is
located north of the platform.
We set traps in lines, with
stations approximately every 10 m.
At each station, we set several
282
traps, including Victor, Sherman

and Tomahawk traps. The trap lines
were set parallel to the existing
trails and streams on the ground
and/or in vegetation (e.g., branches,
logs). Traps were baited every day
or every two days, depending on
how quickly the bait was removed.
Bait consisted of oats, peanut
butter, and vanilla. We occasionally
added fruits and vegetables for
traps set near San Martin-3. The
total trapping effort was 1651
nights-traps in San Martin-3 and
1065 night-traps in Cashiriari-2.
Even though we spent about the
same time at both sites, trapping
was higher at San Martin-3 because
we did not use Tomahawk traps at
Cashiriari-2. Tomahawks had very
low capture rates; trapping at San
Martin-3 without the Tomahawk
traps was 1464 night-traps. We also
set a series of pitfall traps at
Cashiriari-2.
Museum specimens were prepared and have been deposited at
the Museo de Historia Natural,
Unversidad Nacional Mayor de San

Marcos in Lima.
Results and
Discussion
We found a total of 13 species

of small rodents and six species of
marsupials. Table 1 lists the species
sampled at each location, how they
were recorded, and the number of
specimens sampled. Identifications
that require confirmation are indicated by a cf= prefix before the
specific name. The rodent
Oxymycterus inca and the marsupial
Monodelhis emiliae are noteworthy
records. This last species if the first
record for the Department of

Cusco, and just the third record for
the country (also found in Loreto
and Madre de Dios). The genus
Oxymycterus of the Akodontitni tribe
is a common group in the Andes
region at higher altitudes. Only a
few species of this genus live in the
lowland tropics (Reig 1987,
Hershkovitz 1994). Thus, the
taxonomic situation of this species
needs to be determined.
We found a high diversity of
small, non-volant mammals. At San
Martin-3, we found 18 species; at
Cashiriari-2, we found 8 (Table 1).
Our work at Cashiriari-2 added only
Table 1. Small, non-volant mammals found at San Martin-3 and

Cashiriari-2, Lower Urubamba region (number of specimens
listed; Ob = observed, and Vo = registered by vocalization, cf
means the identification requires confirmation)
San Martin - 3
Cashiriari - 2
DIDELPHIMORPHIA
Family Didelphidae
Chironectes minimus
Didelphis marsupialis
Marmosops noctivagus
Marmosops parvidens
Monodelphis emiliae
Philander opossum
RODENTIA
Family Muridae
Neacomys spinosus
Nectomys squamipes
Oecomys bicolor
Oligoryzomys microtis
Oryzomys capito
Oryzomys macconnelli
Oryzomys nitidus
Oryzomys cf. yunganus
Oxymycterus inca
Rhipidomys cf. couesi
Family Echimyidae
Dactylomys dactylinus
Proechimys cf. brevicauda
Proechimys cf. steerei
Total species
Total individuals
Ob
2
1
1
Ob
9
2
14
1
3
5
27
1
4
2
Vo
5
6
18
83
1
1
5
1
4
4
2
2
8
20
283
20
18
16
# Of Species
14
12
10
8
6
4
San Martin - 3
Cashiriari - 2
0
0
10
11
12
13
# of Days
Figure 2. Species accumulation curves for the small, non-volant

mammals at San Martin-3 and Cashiriari-2.
one additional species, Marmosops
parvidens, to those recorded at San
Martin-3. Despite the short time
period for sampling (15 days at each
location) and the difference in
trapping effort, we were able to
compare this study with others
conducted in nearby areas, including Pakitza (1157S, 7117W; ca.
360 m), Cusco Amazonico (1233S,
6903W, 200 m), and Balta (10
08S, 13W; ca. 300 m). Capture
rates were about 5% in San Martin3 (1464 night-traps and 83 captures) and 2% at Cashiriari-2 (1065
night-traps and 20 captures). In
Pakitza, the capture rate was about
1%; at Cusco Amazonico, it was
slightly higher than 5%. The relatively high capture rate at San
Martin-3 is possibly due to the
numerous rodents trapped near the
camp.
The most common species were
the rodents Oryzomys nitidus and
Oecomys bicolor. The marsupials were
all equally low in abundance at San
Martin-3 and Cashiriari-2. San
Martin-3 had a higher number of
species, which might be due to the
abundance of the bamboo Guadua
sarcocarpa at the site (see Alonso et
al. this volume). The forest struc284
ture at this site may also provide

additional places for shelter,
sources of food, and ways to reduce
predation. Both sites had a lower
number of species than the other
studies indicated as well as fewer
individuals (Appendix 1). The
species accumulation curve shows
the time and the sampling intensity
needed to record 18 species (Fig. 2).
Distribution and
Abundance of
Species
We compared the distribution

and abundance of the species
sampled in this study with other
locations in Per and found that
these factors do not vary much for
the majority of the groups (Appendix 2). The didelphimorfos
(Didelphidae) marsuphials are a
very diverse group, but they are
seldom represented in studies
because they are difficult to assess
unless specialized techniques are
used (Voss and Emmons 1996). We
registered two species from the
common genus Marmosops (Appendix 1). Other species are very rarely
observed and sampled because of
their particular habits. The potential
diversity in the area, based on data

from nearby locations, may well be
13 species in 10 groups (Appendix
1). Within the rodents, two families
(the Muridae represented by the
subfamily Sigmodontinae and the
Echimyidae) are the most common
in tropical forests. The tribe
Oryzomyni, and in particular the
genus Oryzomys, constitute the most
diverse and rich group among the
sigmodontinos. In our study,
Oryzomys nitidus was the most
abundant species (Table 1). In the
other studies, Oryzomys capito was
the most common species. Oecomys
bicolor was the second most abundant species (Table 1), mainly
because it was found feeding near
the camp.
Among the Echimyidos, the
group of spiny rats (Proechimys)
constitutes the most diverse group
with up to four or five sympatric
species at some areas (Emmons and
Feer 1990; Voss and Emmons
1996). In our study, at least two
species were recognized from the
relatively low number of individuals
sampled, which could be due to
forest structure and weather
conditions.
Biodiversity
Analysis
We found a relatively low

species diversity among the
Didelphidae (marsupilas), especially
among the small marmosas
(Marmosa, Marmosops, Gracilinanus,
and Micoureus). In our study, we
found only two of the four to seven
species registered in other locations
(Appendix 2; Voss and Emmons
1996). Most of the species in this
group are arboreal; thus, our ability
to capture them is very low.
Marmosops seems to use the ground
more frequently since we captured

it twice in ground traps. The other
registered groups, including the
Didelphis, Philander and Chironectes,
are commonly found in tropical
forests represented just by one
species. The first two are mainly
terrestrials, while Chironectes has
semi-aquatic habits and is found
near streams and rivers (Emmons
and Feer 1990), which is where we
found it in this study. The other
arboreal groups can only be registered by conducting night walks.
These animals feed mostly on fruits
and insects, although there are some
that have carnivorous diets, usually
depending on their size (Emmons
and Freer 1990). Monodelphis is a
group of small didelphids with
diurnal and terrestrial habits. They
are rarely sampled (Emmons and
Feer 1990, Voss and Emmons
1996). We predict the presence of
three species (M. adusta, M.
brevicaudata, and M. emiliae) in the
study area. One adult female (sensu
Pine et al. 1985) of M. brevicaudata
was was found in lactate state (nine
nipples).
The Sigmodontins (Muridae)
rodents are widely distributed in
tropical rain forest. They can have
arboreal, terrestrial, or semi-aquatic
habitats, depending on their different morphological adaptations
(Emmons and Feer 1990). The most
common group was the Oryzomyni,
considered as the most primivitive
of the rodents (Reig 1987).
Oryzomys usually groups up to four
sympatric species in the Amazonian
rain forest (Voss and Emmons
1996), and they are relatively
common on the forest floor. Even
though the characterization of
species has not been worked out
properly, we identified at least three
species, based on the characteristics
Among the
Echimyidos,
the group
of spiny
rats
(Proechimys)
constitutes
the most
diverse
group with
up to four
or five
sympatric
species.
285
Dactylomys
dactilinus,
the bamboo rat, is
easily
recognized for
its peculiar night
calls,
although
it is very
difficult
to see.
286
described by Musser and Williams

(1985). O. nitidus was the most
abundant; it is the smallest (45 to
75 g in weight) and was found only
at San Martin-3, while O. capito and
O. macconnelli were found in similar
quantities at both well sites (Table
1). Oryzomys, Oligoryzomys, and
Neacomys are omnivorous but not
commonly found on the ground
(Emmons and Feer 1990) and
usually are represented by one
species (Voss and Emmons 1996).
Oligoryzomys prefers open areas and
has been found at human settlements and camps. Neacomys prefers
forests and more dense vegetation
(Emmons and Feer 1990).
In comparison with the previous groups, Oecomys includes arboreal species that prefer feeding on
fruits and vegetation (Emmons and
Feer 1990). However, they can be
found living close to human settlements. O. bicolor could have displayed Oligoryzomys microtis, since we
found many individuals close to the
camp and very few in the forest.
Rhipidomys cf. couesi, was found only
at biodiversity plot #1 in San
Martin-3. Stomach analysis indicated that it had been feeding on
insect larvae living in the bamboo
stems.
The genus Nectomys, represented by N. squamipes, was found
near sources of water. It has semiaquatic life style and can also be
found in secondary forests and
cultivated areas (Emmons and Feer
1990). Our specimens were found
near the streams that crossed
biodiversity plots #1 and #2 at San
Martin-3.
The rodents of the Akodontini
tribe include a group that is not
very diverse in lowland rainforest
(Voss and Emmons 1996). Among
the few Amazonian genus,
Oxymycterus includes forms from the

high Andes (up to 2500 m) and
from the Amazon. The Amazon
forms prefer marshy areas with very
little arboreal vegetation (Emmons
and Feer 1990), similar to some
areas within the bamboo-dominated
forest at San Martin-3 where all of
our samples were found. These
animals are terrestrial, diurnal, and
strictly insectivorous (S. Solari
personal observation). We found O.
inca, a species whose biology,
taxonomic classification, and
geographic and altitude distribution
are poorly known (Hinojosa et al.
1987, Reig 1987, Hershkoitz 1994).
Reig (1987) suggested an Andean
origin for this genus, with migration
and speciation towards lower
altitudes in Amazonia, a proposition
that has been confronted by
Hershkovitz (1994).
The second family of rodents,
the Echimyidae, include mediumsized arboreal rodents,with some
terrestrial groups such as Proechimys
(Emmons and Feer 1990,Voss and
Emmons 1996). The genus
Proechimys (spiny rats) is highly
diverse; three to five species have
been found in one location (Voss
and Emmons 1996). It is a fairly
common genus that it is generally
found on trails, especially at night
(E. Vivar personal communication).
We found this group in different
types of forests, including bamboodominated areas, along streambeds,
and mature forest. One species in
rodents, Dactylomys dactilinus (the
bamboo rat), is easily recognized for
its peculiar night calls, although it is
very difficult to see (Emmons and
Feer 1990). We heard spiny rat calls
every night in our camp at San
Martin-3.
The species richness of small,
non-volant mammals at San
Martin-3 and Cashiriari-2 is similar

to that at other sites in southeastern
Per. We found at least six species
of marsupials didelphimorphos and
13 species of small rodents
(Muridae and Echimyidae). Even
though the species accumulation
curves of the two well sites are
relatively different (Fig. 2), the
averaged capture rate was close to
3.2%. This indicates that some
species have high populations.
Notable records include Monodelphis
brevicaudata and Oxymycterus inca,
which reinforce the concept of high
variability in the composition of
small, non-volant mammals among
locations (Voss and Emmons 1996).
References
Emmons, L. H., and F. Feer. 1990.

Neotropical Rainforest Mammals:
A Field Guide. University of
Chicago Press, Chicago.
Hershkovitz, P. 1994. The description of a new species of South
America Hocicudo, or long-nose
mouse, genus Oxymycterus
(Sigmodontinae, Muroidea), with a
critical review of the generic
context. Fieldiana (Zoology, n.s.)
79: 43.
Hinojosa, F., S. Anderson, and J. L.
Patton. 1987. Two new species of
Oxymycterus (Rodentia) from Per
and Bolivia. American Museum
Novitates 2898: 1-17.
Musser, G. G., and M. M. Williams.
1985. Systematic studies of
Oryzomyine rodents (Muridae):
definitions of Oryzomys villosus and
Oryzomys talamancae. American
Museum Novitates 2810: 1-22.
Pacheco, V., B. D. Patterson, J. L.
Patton, L. H. Emmons; S. Solari,
and C. F. Ascorra. 1993. List of
mammal species known to occur in
Manu Biosphere Reserve, Per.
Publications of the Museum de

Historia Natural, UNMSM, (A), 44:
1-12.
Pacheco, V., and E. Vivar. 1996.
Annotated checklist of the
non-flying mammals at Pakitza,
Manu Reserve Zone, Manu National Park, Per. Pages 577-592
in Manu: The Biodiversity of
and A. Sandoval, eds.)
Editorial Horizonte, Lima.
Patton, J. L. 1987. Species groups
of spiny rats, genus Proechimys
(Rodentia: Echimyidae). Fieldiana
(Zoology, n.s.) 39: 305-345.
Patton, J. L., M. N. F. DaSilva, and
J. R. Malcolm. 1994. Gene genealogy and differentiation among
arboreal spiny rats (Rodentia:
Echimyidae) of the Amazon basin:
a test of the riverine barrier hypothesis. Evolution 48: 1314-1323.
Pine, R.H., P.L. Dalby, and J. O.
Matson. 1985. Ecology, postnatal
development, morphometrics, and
taxonomic status of the
short-tailed opossum, Monodelphis
dimidiata, an apparently
semelparous annual marsupial.
Annals of the Carnegie Museum 54:
195-231.
Reig, O. A. 1987. An assessment of
the systematics and evolution of
the Akodontini, with the description of new fossil species of
Akodon (Cricetidae:
Sigmodontinae). Fieldiana (Zoology, n.s.) 39: 347-399.
Voss R. S., and L. H. Emmons.
1996. Mammalian diversity in
neotropical lowland rainforests: A
preliminary assessment. Bulletin
American Museum of Natural History
230: 1-115.
287
Woodman, N., R. M. Timm, R.

Arana, C. V. Pacheco, C. Schmidt,
E. D. Hooper, and C. Pachecho.
1991. Annotated checklist of the
mammals of Cusco Amazonico,
Per. Occasional Papers of the
Museum of Natural History, University of Kansas 145: 1-12.
288
Appendix 1. Comparative analysis of small, non-volant mammals found in

southeastern Per, in Balta, Pakitza, Cusco Amazonico, and Urubamba.
Balta Pakitza
DIDELPHIMORPHIA
Family Didelphidae
Caluromys lanatus
Chironectes minimus
Gracilinanus agilis
Marmosa murina
Marmosops parvidens
Metachirus nudicaudatus
Micoureus regina
Monodelphis adusta
Monodelphis brevicaudata
Monodelphis emiliae
Philander mcilhennyi
Philander opossum
RODENTIA
Family Muridae
Neacomys spinosus
Neacomys cf. tenuipes
Nectomys squamipes
Neusticomys Perviensis
Oecomys bicolor
Oecomys roberti
Oecomys superans
Oryzomys capito
Oryzomys nitidus
Oryzomys cf. yunganus
Oxymycterus inca
Rhipidomys couesi
Family Echimyidae
Dactylomys dactylinus
Isothrix bistriata
Mesomys hispidus
Proechimys brevicauda
Proechimys cf. simonsi
Proechimys cf. steerei
Proechimys sp. nov. 1
Proechimys sp. nov. 2
TOTAL
x
x
x
x
x
x
x
x
x
Cusco Amazonico Urubamba
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
24
25
27
x
x
x
x
x
x
x
x
x
x
19
289
290
Long-term
Monitoring
of Small,
Non-volant
Mammals
in the Lower
Urubamba
Region
Sergio Solari
Mayor de San Marcos
Introduction
The biodiversity assessment in

the Lower Urubamba region has
resulted in a great improvement in
knowledge about this area. We have
learned that this region of Per has
very rich biological diversity in all
groups studied to date. The recent
explorations of hydrocarbons
(principally gas) in the region by
(Per) B.V., calls for a long-term
monitoring plan to continue
biodiversity studies. This will be
one of the few studies in which
knowledge of the population dynamics of several species will
influence the decision-making
process as the natural resources are

exploited.
The goal of this proposal is to
identify the relation between natural gas exploration and the community of small, non-volant mammals.
Focus should also be directed at
methods for transporting hydrocarbons from the study area. This
proposed five-year plan to study
small, non-volant communities will
help establish:
* the distribution and abundance of the small mammal populations in different localities where
hydrocarbon exploration activities
continue,
* a base study to test possible
environmental impacts and determine the risk of disturbance that
the small, non-volant mammal
communities will experience during
construction and maintenance of
the gas line, and
* a method for detecting environmental variables and their
relation to the natural history of
small, non-volant mammal communities.
The proposal calls for the
application of standardized metho-
291
dologies for long-term monitoring at

the well sites and in those areas that
will be affected by the development
and extension of the gas line. The
principal objective is to determine
the distribution and abundance of
the small, non-volant mammal
populations. This will require
efficient trapping procedures to
optimize sampling while also facilitating a rapid and reliable comparison of the studied variables. The
proposal calls for selecting an
adequate number and type of
habitats directly related to the use
of a variety of traps and methods
of identifying the mammals (audio
and visual).
Description of
Monitoring
Activities
A. Monitoring at the well sites
Monitoring the communities of
small, non-volant mammals at the
well sites should include sampling
twice a year, during the wet and dry
seasons, for a total of three months
with an equal amount of time
devoted to each well site.
Period of evaluation: five years
(1998-2003).
Staff: two investigators
Field work: three months/year
(12 months during the dry season,
12 months during the wet season)
Laboratory work: two months/
year (one month each field season)
Activities: Sample small nonvolant mammals. Clean and prepare
the samples. Evaluate the diversity
and abundance. Determine the
environmental impacts.Prepare a
report and distribute it.
B. Monitoring of the Pipeline
Since the pipeline will cross
several types of habitats, it is
292
necessary to conduct a biological

assessment for the communities of
non-volant mammals in all distinct
habitats before monitoring proceeds. This initial assessment will
provide information on the geographic variation of the species
present in those areas. The number
and location of the localities to be
evaluated will depend on the
possible route of the pipeline. Each
site will be assessed in 10 days. An
outline for the subsequent monitoring program follows.
Period of evaluation: five years
(1998-2003).
Staff: two scientists and one
field assistant.
Field work: two months/year
(distributed among the different
localities to be evaluated).
Lab work: two months/year
(after the field work sessions).
Activities: Sample small mammals; Clean and prepare the
samples; Evaluate the diversity and
abundance; Determine the risks
and environmental impacts; Devise
recommendations with respect to
the planned pipeline route; Prepare
a report and distribute it.
C. Additional Considerations
Field should be accomplished
in coordination with other
biodiversity monitoring programs to
take advantage of logistical support
and on-site personnel.
The samples should be deposited in agreement with current
Peruvian legislation (INRENA),
with coordination of the Museo de
Historia Natural, Universidad
Nacional Mayor de San Marcos
(UNMSM, Per) and the National
Museum of Natural History
(Smithsonian Institution, USA).
Bats:
Biodiversity
Assessment
in the Lower
Urubamba
Region
Don Wilson
Robert Baker
Texas Tech University
Sergio Solari, and
Juan Jos Rodrguez
Mayor de San Marcos
Introduction
Bats are the only group of

mammals that can fly. They comprise the second largest order of
mammals with 177 genera and 925
species (Koopman 1993). They
occur throughout the tropical and
temperate regions of the world,
where they occupy a wide array of
ecological niches. Bats are mostly
active at night, finding shelter
during the day in caves, tree cavities, and crevices. In cities, they can
be found roosting on buildings,
under bridges, and in tunnels. Bats
are known to echolocate; i.e.,

navigate and find food through a
system of high frequency sounds
and their echoes. The bats emit
pulses of sound, then analyze the
returning echoes from the environment around them to avoid obstacles and locate food (Wilson
1997). They can catch insects while
flying using echolocation, but they
also feed on mammal blood, fishes,
small vertebrates and plant material
such as pollen, nectar, and fruits
(Wilson 1973).
Bats play important roles in
ecosystem functioning. They consume large quantities of insects,
they aid in the pollination of hundreds of flowering plant species
that have evolved to open their
flowers at night, and they serve as
seed dispersers by feeding on fruits
(Wilson 1997). Farmers use the
nutrient-rich guano (feces) produced by bats to fertilize fields.
The lowland tropical rain forest
in southeastern Per is one of the
most diverse ecosystems in the
world. The bat fauna there is likely
no exception; Ascorra et al. (1996)
studied the bat fauna at Manu
293
Figure 1. Map of southeastern Per, showing the Urubamba

location and three other sites used for comparison.
National Park in the region and
considered it one of the most
diverse tropical sites in relation to
bat fauna. In our study, we conducted a biodiversity assessment of
the bat fauna of the Lower
Urubamba region.
Methods
Bats were captured with mistnets during May 1997. We sampled

the bat community during the first
two weeks of the month at the San
Martin-3 well site and for the last
two weeks of the month at the
Cashiriari-2 well site (Fig. 1). Each
night, one to 15 mist nets were
placed across trails or streams and
left open between four to 12 hours.
Capture effort varied as nets were
placed to maximize capture of both
individuals and diversity of species.
The sampling effort at San Martin-3
was 58 nets/night; at Cashiriari-2, it
was 95 nets/night (see Fig. 2).
A representative sample of
each species was prepared as a
294
study specimen, either as a standard

museum study skin and skull or
preserved whole in 10% formalin.
Each specimen was also sampled
for both endo- and ecto-parasites by
participating parasitologists (see
Guerrero this volume and Chavez
this volume). Parasites encountered
were preserved separately. In
addition, we saved a small sample
of tissue in lysis buffer for DNA
analysis. Standard karyotypes were
prepared from specimens of particular interest or where taxonomic
problems were known to occur.
All specimens are now stored in
the mammal collection at the
Marcos, Lima, Per.
Results and
Discussion
A total of 51 species of 31
genera and five families of bats
(379 individuals) were captured in
this study (Table 1). The identifica-
tions may change slightly as the

skulls are cleaned and the species
identifications are confirmed. Bats
of the Family Phyllostomidae were
the most abundant with 351 individuals of 40 species; Carollia
perspicillata was the most abundant
species with 64 individuals, followed by Platyrrhinus brachycephalus
with 40, Artibeus obscurus with 25,
Artibeus planirostris with 18, Carollia
brevicauda with 21, and Carollia
castanea with 20. The other four
families were less abundant. The
Vespertilionidae was represented by
five species and 10 individuals, the
Emballonuridae by three species
and nine individuals, the
Molossidae by three species and
eight individuals, and the
Thyropteridae by one species and
one individual. Twenty species were
captured only once, for a total of
38.5% of the sample. Thus, as in
other tropical sites (Wilson et al.
1996), the bat fauna is represented
by a few species with a large number of individuals and many species
with only a few captures. Notewor-
thy records include the finding of

two rare nectivore species, Anoura
latidens and Choeroniscus intermedius,
three Molossidae species at San
Martin-3, and the sympatry of three
species of Dermanura at
Cashiriari-2.
Species richness was different
at San Martin-3 and Cashiriari-2.
We found 30 species at San
Martin-3 and 43 species at
Cashiriari-2. Of these, 22 were
found in both localities; eight were
found only at San Martin-3 and 21
only at Cashiriari-2 (Appendix 1).
The most common species at each
site were the canopy frugivorous
Artibeus planirostris and Platyrrhinus
brachycephalus at San Martin-3 and
the subcanopy frugivore Rhinophylla
pumilio and canopy frugivore
Uroderma bilobatum at Cashiriari-2
(Appendix 1). A species accumulation curve for data gathered at each
site suggests different patterns of
diversity for each locality (Fig. 2).
A large number of species were
captured in Camisea in a short
period of time. For example,
We found
30 species
at San
Martin-3
and 43
species at
Cashiriari2. Of
these, 22
were
found in
both
localities.
50
45
40
# of Species
35
30
25
20
15
10
San Martin - 3
Cashiriari - 2
0
0
10
11
12
13
14
# of Days
Figure 2. Species accumulation curve for bat species sampled at San Martin-3 and
Cashiriari-2 in the Lower Urubamba region.
295
Nectarivores
10%
Hematophagous
4%
Insectivores
23%
Frugivore-insectivore
10%
Carnivores
4%
Frugivores
49%
Figure 3. Bat diets and their related percentages.
Ascorra et al. (1996) reported 55

species (32 genera) for the Pakitza
Biological Station, Reserved Zone,
Manu National Park, Per (Appendix 2). They considered it one of
the most diverse tropical sites in bat
fauna. Their sample included field
work for 79 days, during the months
of September-November and
February-March over five years
(1987-1992). In Loreto, Per,
Ascorra et al. (1993) reported 62
species (33 genera) after three years
of intensive monthly sampling, and
Gardner (1988) found 62 species at
the Territorio Federal Amazonas,
Venezuela, in a 2.5-year study.
Moreover, Brosset and CharlesDominique (1990) reported 57
species (43 genera) in French
Guyana after 10 years of study, and
Wilson (1990) has found 61 species
(37 genera) after 20 years of studies
at La Selva, Costa Rica. In this
study, we report a similar number of
bat species and genera from a
sampling effort of just one month.
Bats in the Lower Urubamba
region feed on a great variety of
296
food items. Frugivores comprised

50.0%, insectivores 23.1%,
nectarivorous 9.6%, frugivoreinsectivores 9.6%, carnivores 3.8%,
and hematophagos (i.e. blood
feeders) 3.8% of the total number
of species of the sample (Appendix
1, Table 1; Fig. 3). Frugivores were
also the most abundant group in the
bat community of Loreto, Per, also
followed by the insectivores
(Ascorra and Wilson 1992). Most
bats are active at night, from sunset
to sunrise. The activity of insectivorous bats has been correlated
with the activity of the insects that
they feed on. Ascorra et al. (1996)
reported that insect-feeding bats
have a peak of activity just after
sunset, when many insects become
active; the bats decrease their
activity for most of the night and
increase it again right before sunrise
when the insect activity peaks
again. In contrast, frugivores have a
peak of activity during the first part
of the night, but they remain active
for most of the night.
References
Ascorra, C. F., and D. E. Wilson.

1992. Bat frugivory and seed
dispersal in the Amazon, Loreto,
Per. Publicaciones del Museo de
Historia Natural UNMSM 43: 1-6.
Ascorra, C. F., D. L. Gorchov, and
F. Cornejo. 1993. he bats from
Jenaro Herrera, Loreto, Per.
Mammalia 57:533-552.
Ascorra, C. F., S. Solari, and D. E.
Wilson. 1996. Diversidad y
Ecologia de los Quiropteros de
Pakitza. Pages 593-612 in Manu:
the Biodiversity of Southeastern
Perd. (E. Wilson and A. Sandoval,
eds.) Ed. Horizonte, Lima.
Brosset, A., and P. Charles Dominique. 1990. The bats form French
Guyana: a taxonomic, faunistic
and ecological approach. Mammalia
54: 509-560.
Gardner, A. 1988. The mammals of
Parque Nacional Serrania de la
Neblina, Territorio Federal
Amazonas, Venezuela. Pages 695765 in Cerro de la Neblina.
Resultados de la Expedicion 19831987. (C. Brewer-Carias, ed.)
Editorial Sucre, Caracas.
Koopman, K. F. 1993. Order
Chiroptera. Pages 137-241 in
Mammal Species of the World: A
Taxonomic and Geographic Reference. 2d Ed. (D. E. Wilson and

D. M. Reeder, eds.) Smithsonian
Institution Press, Washington, DC.
Patterson, B. D., V. Pacheco, and S.
Solari. 1996. Distribution of bats
along an elevational gradient on
Southeastern Per. Journal of
Zoology (London) 240.
Wilson, D. E. 1973. Bat faunas: a
trophic comparison. Systematic
Zoology 22: 14-29.
Wilson, D. E. 1990. Mammals of La
Selva, Costa Rica.Pages 273-286 in
Four Neotropical Rain Forests.
(A..H. Gentry, ed.) Yale University
Press, New Haven.
Wilson, D. E. 1997. Bats in Question: The Smithsonian Answer
Book. Smithsonian Institution
Wilson, D. E., C. F. Ascorra, and
S. Solari. 1996. Bats as indicators
of habitat disturbance. Pages 613625 in Manu: The Biodiversity of
Southeastern Perd. (E. Wilson and
A. Sandoval, eds.) Ed. Horizonte,
Lima.
Table 1. Relative abundance and diversity for trophic guilds (sensu

Patterson et al. 1996) at the San Martin-3 and Cashiriari-2 well sites
(numbers indicate the percentages of the values for each guild from
the total abundance or diversity)
Feeding guild
San Martin - 3
Abundance Diversity
10.0
Insect-carnivore gleaners
4.9
3.3
Hematophagous
1.0
16.7
Low flight frugivores
23.5
30.0
Hight flight frugivores
50.0
16.7
Low flight insectivores
7.8
10.0
High flight iinsectivores
5.9
10.0
Omnivorous glossophagines
5.9
3.3
Omnivorous phyllostomines
1.0
Cashiriari - 2
Abundance Diversity
9.3
2.4
2.3
0.0
18.6
38.8
37.2
46.9
14.0
4.8
4.7
1.2
11.6
5.7
2.3
0.3
297
Appendix 1. Bats recorded in the Lower Urubamba region in May, 1997, at

the San Martin-3 and Cashiriari-2 well sites; number of specimens sampled
and the diet for each are given.
San Martin-3
Family Emballonuridae
Peropteryx macrotis
Saccopteryx bilineata
Saccopteryx leptura
Family Phyllostomidae
Anoura caudifer
Anoura latidens
Artibeus lituratus
Artibeus obscurus
Artibeus planirostris
Carollia brevicauda
Carollia castanea
Carollia perspicillata
Chiroderma trinitatum
Chiroderma villosum
Choeroniscus intermedius
Chrotopterus auritus
Dermanura anderseni
Dermanura cinerea
Dermanura glauca
Desmodus rotundus
Diphylla ecaudata
Enchisthenes hartii
Glossophaga soricina
Lonchophylla thomasi
Mesophylla macconnelli
Micronycteris minuta
Mimon crenulatum
Phylloderma stenops
Phyllostomus hastatus
Platyrrhinus brachycephalus
Platyrrhinus helleri
Platyrrhinus infuscus
Rhinophylla fischerae
Rhinophylla pumilio
Sturnira lilium
Sturnira magna
Sturnira tildae
Tonatia brasiliense
Tonatia saurophila
Tonatia sylvicola
Uroderma bilobatum
298
1
3
1
3
7
14
7
8
2
8
4
1
1
1
1
6
3
1
13
1
6
1
1
2
Cashiriari-2
Primary Diet
1
2
3
insects
insects
insects
1
1
8
18
5
20
19
22
5
11
5
1
18
2
5
1
nectar, pollen
nectar, pollen
fruit
fruit
fruit
fruit
fruit
fruit
fruit
fruit
nectar, pollen
carnivore
fruit
fruit
fruit
blood
blood
fruit
nectar, pollen
nectar, pollen
fruit
insects, fruit
insects, fruit
fruit
carnivore, insects
fruit
fruit
fruit
fruit
fruit
fruit
fruit
fruit
insects, fruit
insects, fruit
insects, fruit
fruit
10
2
1
1
22
20
3
2
27
21
4
14
1
5
23
Appendix 1. Bats in the Lower Urubamba region (Cont.).

San Martin-3
Vampyressa bidens
Vampyressa pusilla
Vampyrodes caraccioli
Family Thyropteridae
Thyroptera tricolor
Family Vespertilionidae
Eptesicus brasiliensis
Myotis nigricans
Myotis riparius
Myotis simus
Family Molossidae
Molossus ater
Molossus molossus
Promops centralis
Total species
Total individuals
Cashiriari-2
6
1
2
1
1
2
2
1
3
30
102
Primary Diet
fruit
fruit
friut
insects
2
6
2
1
3
43
333
insects
insects
insects
insects
insects
insects
insects
299
Appendix 2. Comparative list of bat diversity among three localities

Balta, Pakitza, and Cusco Amazonicoin southesastern Per and the San
Martin-3 and Cashiriari-2 well sites in the Lower Urubamba region (see Fig.
1 in text; number of species present at each locality given for each genera).
Balta Pakitza
Family Emballonuridae
Peropteryx
Rhynchonycterys
Saccopteryx
Family Noctilionidae
Noctilio
Family Phyllostomidae
Chrotopterus
Macrophyllum
Micronycteris
Mimon
Phylloderma
Phyllostomus
Tonatia
Trachops
Vampyrum
Anoura
Choeroniscus
Glossophaga
Lonchophylla
Carollia
Rhinophylla
Artibeus big
Artibeus small
Chiroderma
Platyrrhinus
Sturnira
Uroderma
Vampyressa
Vampyrodes
Desmodus
Diphylla
Furipteridae
Furipterus
Famaily Thyropteridae
Thyroptera
Family Vespertilionidae
Eptesicus
Lasiurus
Myotis
300
1
2
1
1
1
1
2
1
2
1
1
3
1
1
2
3
1
2
4
1
2
1
1
1
3
1
4
2
2
3
2
2
3
1
1
1
1
2
1
3
1
3
3
2
3
2
2
3
1
1
1
1
2
2
1
1
1
1
1
3
2
1
1
1
1
1
1
3
1
3
2
2
2
2
2
2
1
2
1
1
1
3
2
3
4
2
3
3
1
3
1
1
1
1
1
2
2
4
1
4
1
1
3
1
3
Appendix 2. Comparative list of bat diversity among three localities (Cont.).

Balta Pakitza
Family Molossidae
Molossops
Molossus
Nyctinomops
Promops
TOTAL
2
1
56
1
1
55
44
1
51
301
302
Long-term
Monitoring
of Bats in
the Lower
Urubamba
Region
Don Wilson
Introduction
Bats play a number of essential

roles in an ecosystem. Although
some can be minor pests in orchard
crops in certain areas, they are
extremely important pollinators of
many plants and dispersal agents for
many seeds. Bats occur around the
world. They can form some of the
largest colonies of vertebrates on
earth, up to 20 million individuals
in a single cave for some North
American species. However, even
small colonies play an important
role in insect control over a large
area of countryside. Bats are also
among the fastest flying species
known.
The primary objective of
surveying and censusing bats is to
determine community composition,
species richness, and abundance.
Before beginning a survey, an

investigator should have some
knowledge of roosting habits,
nightly foraging activities, seasonal
movements, and effects of environmental factors such as local topography, temperature, humidity, light
intensity, and habitat structure on
abundance patterns. Because roost
sites of some bat species are relatively easy to locate and often
house large aggregations of individuals, they offer considerable
potential for assessing population
characteristics.
The following is an outline for
a long-term monitoring program for
bats in the Lower Urubamba region.
Proposed Long-term
Monitoring of Bats
In the Lower Urubamba, as

with elsewhere in the world, there is
no single method suitable for
monitoring all bats. A combination
of observation techniques and
sampling methods should be employed in the study region for
various bat species.
303
Observation techniques
tend to
be more
useful and
easier to
apply to
roosting
bats than
to flying
bats.
304
Observation Techniques for

Bats
Many types of studies require
the capture of bats to identify them
properly. If identification is not a
problem, observation techniques
provide a useful alternative. In
general, these techniques result in
far less disturbance to individuals
and colonies than do the more
disruptive capture techniques.
Additional material documenting
the techniques discussed below can
be found in Wilson et al. (1996). In
that volume, T. H. Kunz and colleagues outline a wealth of information useful to the study of bat
diversity.
A. Roosting Bats
Observation techniques tend to
be more useful and easier to apply
to roosting bats than to flying bats.
The tendency of most bats to seek
refuge in colonies allows the possibility of studying them while they
are at rest and relatively undisturbed.
* Direct roost counts
The most straightforward way
to determine the population size of
a colony of roosting bats is through
direct counts in the roost. Many
roosts allow adequate visibility for
counting each individual in the
colony, and with the aid of redfiltered lights or night-vision equipment, the possibilities are extended
to caves and mines that are otherwise in total darkness.
In very large colonies, it may be
necessary to count the number of
individuals occupying a given area
such as 1 square m and then multiply that number by the number of
square meters occupied throughout
the roost for an estimate of colony
size. Research design is particularly
important in this type of study. To

count all animals, care must be
taken to ensure that all areas of the
roost are accessible. Also, some
species are especially sensitive to
disturbance in roosting areas. In
such cases, it may be necessary to
make the estimates from a sufficient
distance that the bats are not
unduly alarmed.
Data analysis must explicitly
distinguish between complete
counts and estimates. Complete
counts are only obtained when
every individual in the roost is
visible and can be counted directly.
Estimates are far more common and
result from a number of variations
of quadrat counts and estimates of
area covered.
* Disturbance counts at roosts
In roosts that are impossible to
census directly, it is occasionally
possible to count the bats after
disturbing them. Bats can be stimulated to fly through the use of loud
noise or lights and then can be
censused by direct counts or
through photography or videography, as described above.
* Nightly dispersal counts
Rather than disturbing the bats,
it may be possible to census them as
they leave the roost to feed at night.
Occasionally, this can be done
through visual counts, if light levels
permit. In some situations, radar
imagery has been used to census
dispersing bats. Large flocks of
dispersing bats appear on radar
screens, and if the groups are large
enough, quantitative estimates of
the flock size can be made.
* Nightly emergence counts
A more direct census at the
roosting site is obtained by measuring the nightly emergence of the
colony. If light levels and other
conditions permit, visual counts are
the most direct and straightforward

method for estimating populations.
In some cases, remote devices can
be employed to count the bats when
the investigator is absent. Photography is the most common way to
record emergences (Altenbach et al.
1979) from roost sites. For small
colonies, the photographs can be
taken by hand; for larger, more
extended emergences, an automated
system can record a photograph
every few seconds. Cinematography
or videography may prove equally
useful in such situations. Another
common method is to use mechanical or electronic counting devices to
aid in the estimate. Sophisticated
systems may employ a combination
of photographs and electronic
counting devices to record large
numbers of emerging bats.
* Counts of foliage and cavityroosting bats
Bats that roost in more cryptic
situations require considerably more
effort to census. Frequently the only
way to encounter such roosts is
through systematic searching of all
available habitats. Another possibility is to capture an individual bat
and equip it with radio telemetry
equipment that will allow the
investigator to follow the bat back
to the roost.
* Counts of hibernating bats
Where winters are cold enough
to force bats to hibernate, counts
made at the hibernacula may be
relatively simple, as the bats will be
torpid. Investigators must be careful
not to disturb the bats to the point
where they begin to arouse; this will
result in considerable loss of body
fat and may limit the bats ability to
survive the winter. Such counts
should be done in mid-winter when
the populations are at peak levels.
Many species of bats hibernate in
such dense clusters that direct

counts may not be possible. In such
cases, it may be necessary first to
count a cluster of bats occupying a
smaller area and then multiply that
number by the total area covered by
hibernating bats. Sometimes it helps
to use a frame of known dimensions to determine cluster size. In
this situation, photographic techniques are sometimes useful. If the
areas containing hibernating bats
are relatively flat, photographs work
well for documenting cluster size
and counting the number of individuals per cluster. Stereo photography can sometimes be used for
irregular surfaces.
B. Flying Bats
* Direct visual counts
Direct visual counts of flying
bats are not usually an option,
except in special circumstances. If
the bats are readily observable and
identifiable, then it may be possible
to count them on the wing using
night vision devices or even
low-light-level videography. Such
counts may yield useful estimates
of bat usage over a given area, but
are less likely to result in complete
censuses.
* Counts with motion detectors
The use of motion detectors
can sometimes extend the range for
counting bats as they pass over an
area. Such devices are free from
constraints of low-light levels and
have the added advantage of minimizing disturbance. They may be
used on a nightly or seasonal basis
to obtain estimates of different
usage by flying bats. Infrared devices offer an additional possibility
for remote detection of flying bats.
All such methods have a considerable amount of uncertainty that
requires the investigator to be
305
In many
cases it will
be necessary to
capture
bats in
roosts and
flying bats
for proper
inventory
or monitoring studies.
306
intimately familiar with the habitat

and the habits of the bats being
censused.
* Ultrasonic bat detection
In recent years, the use of
ultrasonic detectors has grown
steadily. Such sophisticated instruments call for specialized knowledge of instrumentation and research design. Ultrasonic bat detectors come in three basic systems,
with varying advantages and disadvantages: 1) heterodyne systems
offer high signal-to-noise ratios,
have the greatest sensitivity, and are
the least expensive systems; 2)
frequency division systems have
lower sensitivity than heterodyne
systems and offer broad-band
detection; and 3) time expansion
systems with digital memory yield
signals that are both audible and
recordable, have lower sensitivity,
and provide a discontinuous signal.
The use of electronic bat
detectors carries additional special
considerations. The sounds must be
validated and a reference library
generated from captured individuals. This is no small task, and the
generation of such reference libraries in various parts of the world is
currently occupying a considerable
number of investigators. They have
usually found that station counts
are more effective than transects.
Another constraint is imposed by
habitat differences. Adequate
monitoring of the air space occupied by flying bats, for example,
requires detectors at different
heights.
Limitations on the use of
electronic bat detectors include the
very high learning curve associated
with the study of echolocation.
Bioacoustics is a field just now
coming into its own, and the pitfalls
of sound analysis and necessity for
maintaining an array of precise

electronic equipment may cause all
but the most intrepid of investigators to opt for more conventional
techniques. At the current state of
knowledge, many call characteristics still overlap between species.
For instance, most species of
Myotis, the most common genus in
most parts of the world, sound alike
on ultrasonic detectors. Species
discrimination can vary tremendously depending on the skill of the
observer.
Finally, many groups of bats
are not particularly detectable using
current technology; e.g., all members of the family Phyllostomidae
are known as whispering bats
because they produce very quiet
signals. There is also a considerable
amount of atmospheric attenuation
caused by factors such as dense
vegetation and high humidity.
Sampling Techniques for
Bats
In many cases it will be necessary to capture bats in roosts and
flying bats for proper inventory or
monitoring studies. Just as with
observation techniques, such
studies require considerable preparation for field studies. In addition,
it is important to use proper procedures for handling bats and to
ensure the proper design of holding
devices for bats. Finally, an essential part of any such study is examining bats and keeping records.
A. Capturing Bats in Roosts
By first locating large colonies
of roosting bats, the capture process can be simplified. In roosts,
bats can be captured by hand, or by
use of various kinds of trapping
devices, including hoop nets,
bucket traps, and funnel and bag

traps.
* Hand capture
The simplest method of capture is to take the bats by hand
from the ceiling or walls of a roost.
For human safety, wear gloves while
handling bats; for bat safety, make
sure the gloves are light weight. If
bats are wedged into tight places, it
may be possible to use forceps or
other tools to extract them without
harm.
* Hoop nets
Hoop nets of the kind used for
insect collecting can be useful for
capturing bats in roosts. Mesh and
whole-cloth bags of the type used
for sweep sampling are also helpful.
A plastic lined bag will aid in
keeping the bats from escaping after
capture. The best nets have long
handles, deep bags, and a relatively
small hoop for maximum maneuverability.
* Bucket traps
Bucket traps can be used to
collect bats from high ceilings. The
bucket should have a wire bottom
to improve visibility and should be
attached to a long pole so it can be
raised to the ceiling and pressed
around a group of roosting bats.
Lining the rim of the bucket with
soft foam will improve the seal and
make it more difficult for bats to
escape. In addition, some sort of
wire spatula may be necessary to
dislodge the bats.
* Funnel and bag traps
These traps consist of two
parts: a funnel of variable length is
inserted over the exit from a roost,
leading down to a catchment bag
that accumulates the captured bats.
Each component can be made from
various kinds of fabric or from
plastic sheeting that may make it

more difficult for the bats to escape.
B. Capturing Flying Bats
Bats can be captured in flight
with hoop nets, mist nets, or harp
traps. Hoop nets of the type described above for roosts can sometimes capture flying bats that use
regular flyways or are somehow
constrained to fly in a particular
area.
* Mist nets
The most common way of
capturing flying bats is through the
use of mist netsfine mesh nets
strung between poles. Net types
include monofilament nylon,
braided nylon, or dacron polyester.
They commonly come in lengths of
6, 9, 12, and 18 m long by 2 m high.
Newly purchased nets should
be unfolded and the loops should be
arranged in order. The loops can
then be tied together for future use.
Net poles can be cut on site, or presized poles can be constructed from
electrical conduit or aluminum tent
poles. In general, two 3-meter
poles/net are needed. An alternative is to poles are nails driven into
trees at properly spaced distances.
Nets can be used efficiently
near roosts to catch emerging bats.
Most bats fly to nearby areas of
open water to drink soon after
emergence, and nets placed over
such water sources are often quite
effective. Another useful site is at
feeding sites such as areas of
localized insect abundance or near
fruiting or flowering trees, depending on the kinds of bats being
sought. Nets can also be placed
along flyways, including trails,
forest gaps, mountain ridges, or in
forest canopies through the use of
pulley systems that can easily
The most
common
way of
capturing
flying
bats is
through
the use
of mist
nets
fine
mesh
nets
strung
between
poles.
307
accommodate stacked nets to

increase the amount of vertical
coverage. Many species of bats
become accustomed to using familiar flyways, and this makes them
vulnerable to mist nets placed
unexpectedly across their routes.
Another technique is net flicking,
where one or both net poles are
held by hand and moved quickly to
entangle passing bats.
Care must be taken in removing
bats from nets to minimize damage
to the bats and avoid injury to the
investigator. One commonly used
technique is to wear a glove on one
hand, which is then used to hold the
bat; that leaves the other hand free
to deftly separate the net from the
bat. It is easiest to remove the bat
from that side of the net first
encountered by the bat. Free the
posterior first and work carefully
towards the head, freeing the wings
along the way.
Furling and dismantling mist
nets. Before closing the nets, all
debris should be removed, including
small leaves, sticks, or insects.
Gather the loops to the top of each
pole and spin or drape the net
around the shelf strings. Tie the
loops together, fold the net together, or carefully stuff it into a
cloth bag. Nets must be dry before
storing to prevent mildew or rot.
* Harp traps
Double and multiple frame harp
traps can be constructed by using
two or more frames of aluminum
tubing, two or more sets of vertically strung wires, and a canvas or
plastic-lined catch bag below the
wires. Monofilament line can be
substituted for wires in some designs. The general idea is to have
the bats encounter the first set of
vertically strung wires, fold its
wings allowing it to pass through,
308
and then as it re-opens its wings,

encounter the second set of wires,
which will cause it to fall into the
bag below.
Trap placement is along the
same principles as outlined above
for mist nets. In addition, harp traps
can be hoisted relatively easily into
the canopy of forests or into high
cave mouths. Trap effectiveness
often hinges on proper spacing and
wire tension.
C. Mist Nets versus
Harp Traps
Mist nets and harp traps are the
most common devices for sampling
bats because they are easily deployed and can be used in a variety
of situations. Disadvantages of mist
nets are that they must be tended
constantly, and when captured bats
become entangled, they must be
removed individually. Advantages
of mist nets are their ease in deployment, relatively low cost, and
portability. Harp traps have made it
possible to capture large numbers
of bats without the need for tedious
extraction of each bat separately.
Mist nets tend to work best for
small bats flying in the sub-canopy
of forests. Harp traps work better
for some small insectivores, and
they seem to be superior in light
rain, as mist nets soak up water or
collect drops that greatly increase
their detection by echolocating bats.
Mist nets cover a larger area, but
harp traps tend to be easier to use
in tight places.
Miscellaneous Methods
Sometimes local conditions will
dictate the development of specialized capture techniques such as
mobile traps in caves. Another
possibility is to use nylon netting
draped over roost entrances. Before
mist nets came into common usage,

trip lines were placed over bodies
of water to knock drinking bats into
the water, where they could be
retrieved. Baiting with fruit or
ultraviolet lights to attract insects
has been used successfully in some
venues. Using or mimicking distress
calls is another possibility that has
occasionally been successful.
Preparation for Field Study
All equipment should be assembled beforehand, including nets,
poles, ropes, extra parts and tools
for harp traps, extra shelf cord for
repairs, string for guy lines, duct
tape, wire, leather gloves, and cloth
bags for holding bats.
Handling Bats
Small bats can be held in the
palm of the hand. Larger bats can
be held by the forearms over the
back. Gloves should be worn to
avoid bites and scratches. Holding
devices for bats include cloth bags,
which should be made from a soft,
open-weave fabric such as monks
cloth. Drawstrings are easier to use
than ties, which means bags have to
be prepared in advance. Large
species should be bagged individually, but small species may do better
if allowed to cluster to reduce
stress and water loss.
Examining bats. Each captured
individual should be carefully
examined and identified by species
and sex. Age can sometimes be
determined through examination of
the degree of ossification of the
long bones or the amount of tooth
wear. Reproductive condition can
be determined for females by
palpating the mammary glands to
detect lactation or palpating the
abdomen to detect pregnancy.
Males should be checked for the

presence and development of chest
and skin glands, and the length and
width of testes should be measured
with calipers. Weight to the nearest
gram can be determined by weighing the bats in the cloth bags with
Pesola scales and then subtracting
the bag weight. Forearm lengths
should be measured to the nearest
mm with dial calipers or a ruler.
Pelage condition should be noted,
and pollen or hair samples can be
taken using scotch tape. Sometimes
valuable evidence of feeding can be
determined from food items carried
into the nets or by examination of
the feces and urine.
Keeping Records
It is a good idea to sketch maps
of capture localities to include
latitude, longitude, trails, streams,
roads, and an indication of the
placement of nets and traps. Also,
photograph roost trees, roosting
groups, individual bats, and sexually
dimorphic characteristics such as
epaulets and skin glands. Everything should be recorded in indelible ink or pencil on water-resistant
paper for safekeeping.
309
References
Altenbach, J. S., K. N. Geluso, and

D. E. Wilson. 1979. Population
size of Tadarida brasiliensis at
Carlsbad Caverns in 1973. Pages
341-48 in Biological Investigations
in the Guadalupe Mountains
National Park, Texas. National
Park Service Proceedings and
Transactions Series No. 4. (H. H.
Genoways and R. J. Baker, eds.)
Washington, DC.
Fenton, M. B. 1992. Bats. Facts on
File, New York.
Hill, J. E., and J. D. Smith. 1984.
Bats: A Natural History. University
of Texas Press, Austin.
Kingdon, J. 1974. East African
Mammals: An Atlas of Evolution
in Africa. II(A). Insectivores and
Bats. Academic Press, London.
Kunz, T. H. 1982. Ecology of Bats.
Plenum Press, New York.
Nowak, R. M. 1991. Walkers
Mammals of the World. Vol. 1.
Fifth ed. Johns Hopkins University
Press, Baltimore.
310
Van Der Merwe, M. 1978. Postnatal

development and mother-infant
relationships in the natal clinging
bat Miniopterus schreibersi natalensis
(A. Smith 1834). Proceedings of the
Fourth International Bat Research
Conference, Nairobi, Kenya 1:
309-322.
Wilson, D. E. 1997. Bats in Question. Smithsonian Institution Press,
Washington, DC.
Wilson, D. E., and D. M. Reeder,
eds. 1993. Mammal Species of the
World, Smithsonian Institution
Wilson, D. E., F. R. Cole, J. D.
Nichols, R. Rudran, and M. S.
Foster. 1996. Measuring and
Monitoring Biodiversity: Standard
Methods for Mammals.
Washington, DC.
Medium
and Large
Mammals:
Biodiversity
Assessment
in the Lower
Urubamba
Region
Major Boddicker
Rocky Mountain Wildlife
Enterprises
Introduction
Because of growing concerns

about the management of biological
systems impacted by mineral extraction, biodiversity surveys have been
suggested as a method to develop
baseline data about those systems.
Monitoring processes to follow up
on the baseline surveys are in the
process of being developed and
tested. At the well sites in the
Lower Urubamba, the goal is to
employ a long-term monitoring
program based on sound biological
science to measure changes brought
about by natural gas exploration and
production and use the results to
minimize or eliminate long-term
negative impacts on the ecosystem.
Through monitoring, human caused

changes may be detected early on so
that mitigation can be implemented.
Monitoring might also lead to
preventive actions prior to natural
gas extraction, thus lowering the
potential for negative impacts. This
strategy could be called an ecosystem management processa synthesis of modern biology and
management applied to large-scale
ecological manipulation for the
extraction of natural gas products
from a previously undisturbed area.
Methodologies for measuring
the diversity of large mammals in
temperate and arctic climes, open
grasslands, and deserts are often of
little use in tropical rain forest.
Appropriate methods of monitoring
in rain forest are often very expensive efforts prolonged over a very
long time. They are often not
practical given the limited financial
and personnel commitments and
time windows that are usually
associated with natural resource
development projects. At
Urubamba, the objectives were to
observe the first test drilling sites,
establish monitoring test areas,
311
Biodiversity
monitoring
of medium
and large
mammals
has no
generally
accepted
wide-range
testing
procedure.
312
conduct preliminary medium and

large mammal surveys, develop
recommendations for needed modifications of survey methods, and
prepare for monitoring. Despite the
challenges inherent in achieving
these objectives during introductory
visits to the sites, significant
progress was made, as detailed in
this report.
Biodiversity monitoring of
medium and large mammals has no
generally accepted wide-range
testing procedure. Hundreds of
excellent specific tests are available
for single species and, in some
instances, families of mammals; e.g.
aerial surveys of Cervidae on
African plains. The challenges
presented by the rain forest and
mammal adaption to it in the Lower
Urubamba preclude the use of any
one method for monitoring. Vegetative cover makes aerial surveys
difficult. Radio telemetry is greatly
restricted because of costs and
receiver access to mammal-use
areas. There are no hunter/trapper
harvest records, no animal damagecontrol reports, and no wildlife
agencies to track population trends
at large or small scales. Vehicle
access to the well sites is limited to
aircraft or canoe.
In effect, biodiversity monitoring in tropical rainforest areas such
as the Lower Urubamba takes
science back to its basicshiking, a
pencil, and a notebook (batterypowered computers can also be
used). There are no shortcuts.
Sophisticated technology meets its
limits because of climate, topography, logistics, sociological requirements, and health considerations.
Statistically, reliability tests require
data gathering beyond the scope of
some basic biodiversity surveys. If
those surveys indicate a need for
more sophisticated methods, then

that focus must be directed at the
species requiring it.
Methods
Upon arrival at the well sites, a

general walk-over was completed to
assess the size and impact of the
operations. The selection of survey
routes and lengths was made based
on an estimate of the sound, sight,
air currents, and water impacts of
the drilling operations and the
estimated home ranges of the
largest mammals that would be
surveyed. It was determined that
the test plots for monitoring plants,
arthropods, and other life forms
were of insufficient size for monitoring larger mammals.
At San Martin-3, where the
drilling pad was being established,
two trails were developed. Trail A
extended west from the camp
facilities across the pad to the top
of the highest point, about .5 km,
then approximately 0.5 km to the
bottom of biodiversity plot #2 (see
Alonso et al. this volume). The trail
then wound for 0.25 km to the
south along the creek bottom and
terminated at the southeastern
corner of the plot. A second leg of
this trail continued from the trail
intersection to the north approximately 1 km along the creek bottom
away from biodiversity plot #2.
After five days of surveying, it
became obvious that a longer
survey route was needed. An additional leg, approximately 1 km in
length, was cut due west to the top
of the ridge. The total length of
trail A is 3.25 km in length.
The eastern trail began at the
edge of camp and extended .75 km
to the stream bottom at the water
pumping station. This trail was
extended during the last week to the
south along the stream bottom for

2 km, for a total of 2.75 km (see
Fig. 1). The two trails included
portions of biodiversity plots #1
and #2 as well as the drilling site
and the two highest and lowest
physical points in the area of San
Martin-3. Both trails take researchers of hearing and sight range of
the drilling site, although they do
not take researchers out of the
range of helicopter traffic noise.
At Cashiriari-2, similar survey
routes were established. The northern route begins at camp, extends
across the drilling site and north on
a developed road approximately 2.5
km to the water pumping station,

then continues downstream for 1.5
km to the northwest in the stream
bed. A 0.5 km dogleg trail was cut
along the old Camisea road to
extend the survey into more traditional habitat at a higher elevation.
The Camisea road branches away
from the pump road about 2 km
from the camp. The eastern route
begins at camp and extends 2.5 km
east and northeast along the old
road to Cashiriari. About 1 km has
been reopened by botany and bird
researchers. Another 1.5 km was
opened to provide sufficient length
for getting out of sight and hearing
Figure 1. Diagram of San Martin-3, Lower Urubamba region,

Cusco, Per.
313
Figure 2. Diagram of Cashiriari-2, Lower Urubamba region,

Cusco, Per.
range of the drilling area (Fig. 2).
The trails contain the highest
geographical feature near the drilling site, lowest point, ridge tops,
stream bottoms, and transversed
altitudes in between. Both trails
passed along edges of the
biodiversity test plots. All survey
lines extended beyond sight and
sound of the drilling rigs, but not
beyond aircraft noise.
Five methods were used and
tested for practicality and effectiveness at the two well sites: visual
surveys; vocalization/calling surveys; modified scent-post surveys;
track, scat, and sign surveys; and
snare and conibear trapping surveys.
(Detailed descriptions of these
314
methods are given in Appendices 37.) All of the methods were used in
an integrated approach; therefore,
the efficiencies of each survey
method enhanced the others. Repetition, volume of observations,
and the variety of approaches used
to find the large mammals were
assumed to reduce negative biases,
mistakes, and problems with language communications. The visual,
track/scat/sign, and snare and
conibear trapping surveys were
"passive". The scent-post and
calling surveys were "active".
The objectives of the surveys
were to find all species of the
mammals in the area of influence
of the drilling site that weigh more
than 2 kilograms (kg), identify these

mammals, develop an estimate of
density for each species, and develop a method of monitoring the
factors of the preceding three
objectives over five consecutive
years to determine the effects on
these animals, positive or negative,
that might result from gas exploration and production.
The five methods were implemented immediately after the initial
scouting of survey lines was completed. The approximate order of
implementation was visual survey;
track, scat, and sign survey; modified scent-post survey; snare and
conibear trapping survey; and the
vocalization survey. Results of all
survey methods were pooled. All
duplications of results were noted,
and the results were adjusted to
eliminate duplication. Observations
were recorded in a notebook, in
daily log notes, and through photographs. Tangible evidence (scats,
tracks, hair, bone, etc.) was collected, identified, photographed,
and stored.
Approximately seven hours per
day were spent walking the survey
trails and obtaining data. Usually,
the trails were walked twice a day
to remove captured animals and to
renew baits.
Two simple indices were constructed to assist researchers in
synthesizing and analyzing data.
Both indexes are recommended for
use in long-term monitoring. The
purpose of the first index, the
occurrence index, is to provide a
confirmed species list based on
accumulated evidence from all five
survey methods (see Appendix 1).
In summary, it proposes a point
system that gives values to various
kinds of evidence according to the
reliability of the evidence. When
the accumulated points of evidence

reach a threshold, the researcher
concludes the species is present on
the surveyed area.
Only two types of evidencea
specimen of an individual whole
animal and observation of an
animal verified by a corroborator or
photographic evidencequalify for
a complete record. At least two
types of other tangible evidence are
required for confirmation of a
species presence. The second index,
the abundance index, is a function
of the first; it puts the occurrence
index into numerical form for
comparative purposes (see Appendix 2). It consists of multiplying
evidence scores from the occurrence index by the number of nonduplicated records of evidence.
Results from use of the five
survey methods can be synthesized
in the occurrence index and extended in the abundance index to
provide the following: 1) a highly
reliable species list; 2) a replicable
process for repeating the collection
of a species list for monitoring
purposes; 3) a numerical score for
ranking the value and strength of
the record; 4) a numerical extension
of the evidence into a comparable
index so that occurrence and abundance records can be compared
between survey areas, years, and
other parameters; 5) and a comparative format for identifying changes
in medium and large mammal
abundance that might call for
further study and determination of
the causes of the changes.
Results
from use
of the five
survey
methods
can be
synthesized in the
occurrence
index and
extended
in the
abundance
index.
Results and
Discussion
A list of species that could be

expected to occur at the two well
sites was compiled from range maps
315
Table 1. List of medium and large mammals at San Martin-3 (SM3) and Cashiriari-2
(C2) well sites in the Lower Urubamba region, Cusco, Per (in the Occurrence Index,
numbers of 8 or higher indicate confirmation of the presence of the species at the time
of the survey; numbers of 7 or below indicate the potential presence of the species but
its presence was not confirmed; in the Abundance Index, numbers are determined by
multiplying the type of evidence found for a species times the number of independent
observations of that evidence; the index expresses whether a species is more abundant
at one location or time than another, but it is not comparable between species)
Species
Occurence Index
SM3
C2
Family Didelphidae (Marsupials)

Caluromys lanatus (western woolly opossum)
Didelphis marsupialis (common opossum)
Philander opossum (common grey four-eyed opossum)
Chironectes minimus (water opossum)
Metachirus nudicaudatus (brown four-eyed opossum)
Family Myrmecophagidae (anteaters)
Myrmecophaga tridactyla (giant anteater)
Tamandua tetradactyla (collared anteater)
Cyclopes didactylus (pygmy anteater)
Family Bradypodidae and Megalonychidae (sloths)
Bradypus variegatus (brown-throated three-toed sloth)
Choloepus hoffmanni (Hoffmans two-toed sloth)
Choloepus didactylus (southern two-toed sloth)
Family Dasypodidae (armadillos)
Cabassous unicinctus (southern naked-tailed armadillo)
Priodontes maximus (giant armadillo)
Dasypus novemcinctus (nine-banded long-nosed armadillo)
Dasypus kappleri (great long-nosed armadillo)
Family Callitrichidae (marmosets and tamarins)
Cebuella pygmaea (pygmy marmoset)
Saguinus fusciollis (saddleback tamarin)
Saguinas mystax (black-chested mustached tamarin)
Saguinus imperator (emporor tamarin)
Aotus species (night or owl monkeys)
Callicebus moloch brunneus (dusky titi monkey)
Saimiri sciureus boliviensis (common squirrel monkey)
Cebus apella (brown capuchin monkey)
Cebus albifrons (white-fronted caphucin monkey)
Cebus pithecia (Guianan saki monkey)
Alouatta seniculus (red-howler monkey)
Lagothrix lagothricha (common wooly monkey)
Ateles paniscus chemek (black-spider monkey)
Family Canidae (dog family)
Atelocynus microtis (short-eared dog)
Speothos venaticus (bush dog)
316
Abundance
SM3
C2
5
32
5
22
5
5
10
5
5
5
0
32
0
17
0
0
5
0
0
0
7
7
5
5
5
5
2
2
0
0
0
0
15
5
5
5
5
15
10
0
0
0
0
10
5
5
10
5
5
5
15
5
0
0
7
0
0
0
23
0
5
5
5
15
5
5
20
20
20
5
20
5
20
5
20
20
5
5
5
5
20
20
15
5
5
5
0
0
0
10
0
0
45
205
125
0
35
0
25
0
25
25
0
0
0
0
45
210
0
0
0
0
Table 1. List of medium and large mammals (Cont.).

Species
Occurence Index
SM3
C2
Family Procyonidae (raccoon family)

Procyon cancrivorus (crab-eating raccoon)
Nasua nasua (South American coati)
Bassaricyon gabbi (olingo)
Potos flavus (kinkajou)
Family Mustelidae (weasel family)
Mustela africana (Amazon weasel)
Galictus vittata (grison or huron)
Eira barbara (tayra)
Lutra longicaudis (southern river otter)
Pteronura brasiliensis (giant otter)
Damily Felidae (cat family)
Felis pardalis (ocelot)
Felis wiedii (margay)
Felis yagouaroundi (jaguarundi)
Felis concolor (puma)
Panthera onca (jaguar)
Family Tapiridae (tapirs)
Tapirus terrestris (Brazilian tapir)
Family Tayassuidae (peccaries)
Tayassu tajacu (collared peccary)
Tayassu pecari (white-lipped peccary)
Family Cervidae (deer family)
Mazama americana (red brocket deer)
Mazama gouazoubira (grey brocket deer)
Family Sciuridae (squirrels)
Sciurus spadiceus (southern Amazon red squirrel)
Microsciurus flaviventer (Amazon dwarf squirrel)
Family Erethizontidae (porcupines)
Coendou prehensilis (Brazilian porcupine)
Family Agoutidae, Dasyproctidae, Hydrochaeridae
Hydrochaeris hydrochaeris (capybara)
Agouti paca (paca)
Dinomys branickii (pacarana)
Dasyprocta punctata (Central American common agouti)
Myoprocta pratti (green acouchy)
Family Leporidae (rabbits)
Silvilagus brasiliensis (Brazilian rabbit)
Abundance
SM3
C2
10
12
5
5
12
10
5
10
5
9
0
0
21
10
0
5
5
0
15
5
5
5
0
18
5
5
0
0
8
0
0
0
0
13
0
0
15
10
5
5
5
25
10
5
21
21
40
5
0
0
8
55
5
0
40
35
28
13
217
34
30
20
27
5
257
50
58
0
28
5
28
5
58
0
64
0
5
15
20
15
5
15
15
25
5
13
5
23
13
5
13
5
30
13
0
145
0
174
53
0
65
0
80
31
22
22
62
52
317
We confirmed the
presence
of 25
species at
San
Martin-3
and 26
species at
Cashiriari-2
for a total
of 33 nonduplicated
species.
318
and distribution descriptions found

in Emmons (1990), a checklist
developed by Guerrero and Patron
(1996), and Pacheco et al. (1993).
The plates Emmons were shown to
three local guides, who were asked
whether or not the species occurred
in the area. The species list in Table
1 reflects the sum of these four
sources, which were then confirmed
as present or not by the score
derived from accumulated evidence
at each site. (Table 1 is organized in
the same order as Emmons field
guide to make it convenient for the
reader to refer to that text for
details about the species.) While
there are differences of opinion
about the range maps developed in
Emmons, this author is not qualified to discuss taxonomic or historical range issues.
Fifty-eight species of mammals
were considered potentially present
at the two well sites. We confirmed
the presence of 25 species at San
Martin-3 and 26 species at
Cashiriari-2 for a total of 33 nonduplicated species. The total number of confirmed species was 57%
of the potential list. Twenty-four
or 41% of the possible species
were not confirmed as present, but
could be present, even common.
Some monkeys, canopy-inhabiting
species, species with large territories and/or low mobility, and
nocturnal animals can easily avoid
survey methods in the rain forest.
All species listed have a reasonable
probability of living on, or occasionally moving through, the study
area. Furthermore, the agouti, small
cats, and armadillo species have
very similar tracks and sizes and are
difficult to distinguish without
specimens. These species have a
tendency to be lumped.
All species listed with a numerical score of 8 or higher in the

first two columns are considered
confirmed records at the site listed.
Species with scores of 7 or less are
considered unconfirmed but could
exist at the site. The two columns
on the right side of the table compare abundance at the two well
sites. The higher the numbers, the
more individual animals were likely
present. Dividing the smaller number into the larger one will provide
a rough comparative ratio of abundance of a species at one site over
the other. The higher the occurrence
index number, the more reliable the
record.
Difficulties were encountered
in distinguishing the sign of the
small cats, particularly margays,
ocelots, and jaguarundis. According
to Emmons (1990), ocelot tracks
measure about 2.75 cm in females
and 3 cm in males. During this
study, measurements were taken of
four sets of tracks, all identified as
ocelot tracks by one hunter and the
author. They were found within two
hours over 3 km and were likely
different individual cats. The width
measurements were 4 cm, 4.5 cm,
and two at 5 cm. All tracks were
fresh on wet sand or clay and of
excellent quality. Cat tracks with
different heel pad configurations
and ranging in size from 2.5 cm to 5
cm across were also observed. It
was assumed that at least two
species of small cat were present,
probably margay and ocelot; however, further studies are needed to
verify these assumptions.
The following mammal groups
require further study and specimen
collection to clear up doubts and to
provide measurements and samples:
all of the Didelphidae (opossum)
species, two-toed sloths, armadil-
los, canines, the procyonids (raccoon) family, ocelots, oncillas,

margay and jaguarundi cats, grisons
(or hurons), the squirrel family, and
the agouti family.
General observations indicated
that the species expected at the
sites were present and that numbers
or densities of those populations
were in expected ranges. Large
mammal activity around the sites
was close and continuous; the noise
and operations did not appear to
have any severe impacts other than
directly on the drilling pad. Red
brocket deer and Brazilian rabbit
tracks were present at both well
sites. Ocelot, tapir, and peccary
tracks, beds, and other signs were
found within 100 m of the drilling
pads. Cat species that are often
considered the first to flee human
development (ocelots, pumas, and
jaguars) were also active to within
100 m of the pad at Cashiriari-2. A
major loafing and toilet area for
jaguars and pumas was found
approximately 2.5 km from the well
site where the engines could be
clearly heard. Sightings and signs of
large mammals did not appear to be
any less numerous near the drilling
activity than 2.5 km away.
The observations and signs
present at the well sites were sufficient to develop a density estimate
for 28 species, as reported in
Table 2.
San Martin-3 clearly had a
higher abundance index (1643) for
most animals than did Cashiriari-2
(1028). San Martin-3 is at a lower
elevation, is wetter, and has a
shorter overstory and more
bamboo.Thus, its carrying capacity
for large mammals may be greater
than Cashiriari-2, which is at a
slightly higher elevation with a
higher canopy, significantly less
water, and a much longer distance

to travel to water. In addition, the
predator populationjaguars,
pumas, ocelots, and short-eared
dogswas higher at Cashiriari-2
than at San Martin-3.
Five survey methods were used
to develop baseline biodiversity
data on large mammals at the two
well sites. Thirty-three species of
large mammals were confirmed
present at the sites25 at San
Martin-3, 26 at Cashiriari.
The most effective survey
method for confirming species
presence was the track/scat/sign
survey, followed by the scent-post
survey using trapping lures. Vocalization/calling, visual, and trapping
surveys were less effective, but they
nevertheless contributed significantly to the successful development of the base list.
Occurrence and abundance
indices were tested. Both show
promise for the monitoring program.
Ocelot,
tapir, and
peccary
tracks,
beds, and
other signs
were found
within 100
m of the
drilling
pads.
References
Anonymous. 1981. Indices of

Predator Abundance in The Western United States. U.S. Fish and
Wildlife Service, Washington, DC.
Emmons, L.H. 1990. Neotropical
Rainforest Mammals: A Field
Guide. The University of Chicago
Press, Chicago.
Guerrero, R., and H.Z. Patron. Los
Mamiferos del Rio Camisea, Cusco,
Per: Estudio Preliminar. Pages
91-94 in Proceedings from the
Workshop on Biological and
Cultural Diversity of the Lower
Urubamba, Per. (S. Udvardy and
A. Sandoval, eds.) Smithsonian
Institution, Washington, DC.
Pacheco, V., et al. List of mammal
species known to occur in Manu
Biosphere Reserve, Per. Mus.
Hist. Nat. UNMSM 44 (A): 1-12.
319
Table 2. Density estimate (per km2) of the medium and large mammals at
San Martin-3 and Cashiriari-2 in the Lower Urubamba region, Cusco, Per
(** not confirmed as present or too little information to support an
estimate)
Species
Mammals per km2

Cashiriari-2
San Martin-3
Common Opossum
Nine-banded long-nosed armadillo
Saddleback tamarin
Black-chested mustached tamarin
Common squirrel monkey
Brown capuchin monkey
White-fronted capuchin monkey
Red-howler monkey
Black-spider monkey
Short-eared dog
Bush dog
Crab-eating raccoon
South American coati
Tayra
Ocelot
Margay
Puma
Jaguar
Tapir
Collared peccary
White-lipped peccary
Red brocket deer
Southern Amazon red squirrel
Amazon dwarf squirrel
Paca
Common agouti - D. punctata
Green acouchy
Brazilian rabbit
320
4
**
**
**
4
8
4
2
2
**
0.25
2
1
0.25
1.5
0.5
**
**
3.5
10
2
5
**
2
10
20
6
6
1
2
2
2
**
2
8
**
**
0.25
**
2
1
0.25
1.5
0.5
0.25
0.1
0.5
2.5
**
6
2
4
6
10
3
6
Appendix 1. Biodiversity Occurence Index.

A method of standardizing
occurrence records for large mammal species, without physically
collecting them, is needed for
monitoring purposes. The author is
unaware that such a method exists.
Consequently, a biodiversity occurrence index is being proposed as a
preliminary effort to start the
process of developing such a
standardized method. The need for
such a system is obvious when one
accepts the job of monitoring large
mammalsa large, very diverse
group of animals, most of which
are difficult to find and count. For
short-term, general location, and
identification studies, it becomes
obvious that physical collection and
observation cannot provide the
necessary information.
Various survey methods provide a wide range of observations
and evidence, that, when pooled,
can provide reliable and verifiable
data. How can that data be used?
Assign numerical values to the
pieces of evidence and determine
by adding the values, how much
evidence is sufficient to confirm the
presence of a species at a given
time and place. This threshold value
at present is somewhat arbitrary. Its
value will depend on the experience
and knowledge of the decisionmakers and field testing to determine its best level.
The following evidence list was
used, along with the points this
researcher considered reasonable,
for reliability towards a confirmation. Discussions on each element
of evidence and scoring follows the
list. This is an invitation for discussion about such a proposed index.
Discussion of Elements of
Evidence and Points
Species Collected - 10 points.
Obviously, a specimen of each
species collected from the study
area, during the study period, would
be a definitive record and rates 10
points. Any further collection of
specimens becomes useful in determining how many of the species
occurs at the study site. The total
number of a given species collected
at the site should be kept for use in
the abundance index calculations.
Any evidence associated with the
collected specimens should not be
included in any further evidence
records, for it factors in duplication.
Species Observed - 10 points.
There are qualifiers for this
evidence. The observer must have
demonstrable experience at observing and identifying wildlife, and that
observation should be verified with
a witness, photograph, or other
tangible evidence to back it up. If
the observation fulfills the verification standards then it is worth 10
points.
Various
survey
methods
provide a
wide range
of observations
and evidence,
that, when
pooled,
can
provide
reliable
and verifiable data.
Sign.
Sign is a collective term for the
evidence animals leave which
indicates it has been, or is now,
present at a location. Sign includes
tracks (spoor), scats (feces), urine,
vomitus, feeding leftovers, trails,
hair, bone, beds, nests, burrows,
food caches, waste piles, toilets,
territorial markings, etc. An individual piece of sign may be strong
or weak evidence of presence
depending on many factors. Many
pieces of sign of the same type or
different types can result in confirmation of a species presence and
321
Appendix 1. Biodiversity Occurence Index (Cont.).
322
be equal to a collection or verified

observation. (This statement is an
expert observation subject to
discussion.)
erased so new tracks can be made

to confirm presence. A confirmed
track should be recorded only once
in the occurrence indextotal.
Tracks - 5 points.
Clear unambiguous tracks are
like finger prints of a species and
quite reliable for identifying the
presence of many species; e.g., tapir
and jaguar. Many species, however,
have tracks which are very similar
and difficult to differentiate under
the best of conditions: e.g., ocelot,
margay, and jaguarundi.
Identification by tracks requires
a reasonable skill level and experience not common among even
experienced naturalists. During this
monitoring effort, local guides
occasionally made errors in track
identification, as did the researcher.
It took discussion, further tracking,
and sometimes a third opinion to
come to an acceptable conclusion
on what the track represented. Soil
type, wind, rain, and artifacts are
factors in track determination.
Consequently, the value of tracks in
determining species occurrence
was assigned 5 points. One, or an
accumulation of tracks, by itself
was not deemed sufficient evidence
of occurrence. Another corroborating piece of evidence was required.
The number of times identifiable tracks were found which were
in a different place, or were of
different size or configuration,
should be noted since they should
signify a different animal, and
therefore, be useful in calculating
the abundance index.
Tracks generally are short lived
entities in the environment. After a
track is found and identified, photographed, and measured, it should be
Feces (scats) - 3 points.

The feces or scats of many
mammals are unique and can be
identified to family, genus, and even
species by the diameter, length,
form, segmentation, contents,
placement, and odor. The reliability
of a scat as an indicator of species
presence can vary from absolute to
no value, depending on the quality
of the scat and ability of the researcher. Many species produce
ambiguous scats which become very
difficult to identify. Species which
are closely related, like red and gray
brocket deer, may deposit scats
which are difficult to differentiate.
Ocelot, margay, jaguaroundi,
housecats, oncilla and juvenile
puma, and jaguar conceivably could
deposit scats of similar appearance
and size. Other corroborating
evidence would be necessary to
confirm an occurrence.
Scats are sources of much
additional information including
size of the individual, food consumed by the animal, social habits,
and territorial marking habits. They
are also good sources of bone, hair,
hooves, and claws of other species
which provide additional evidence
of occurrence not found by other
survey methods.
In the proposed index, scats
were assigned a point value of 3. It
is anticipated that scats could
introduce a significant error factor
in the identification and use for
confirming a species presence. At
least one other strong piece of

evidence would be necessary with it
to confirm a presence.
Hair - 5 points.
Hair is excellent evidence of
species presence and may deserve
more than 5 points in this index.
However, to identify hair requires
equipment, technology, and knowledge usually unavailable in field
research. Some species of large
mammals have many types of hair.
To be completely useful, a detailed
key to hair identification is needed,
but is unavailable on Neotropical
mammals. Hair as a definitive proof
is therefore possible, but not probable, so in this scheme gets a 5point rank. At least one other type
of evidence is needed to rate a
confirmation.
Bone - 5 points.
The skulls of most animals are
definitive proof of a species
presence at some time. Depending
on the including tooth patterns,
characteristic bite locations, entry
characters on carcasses, dirt movement patterns, refuse piles, seed or
fruit caches, etc. When feeding sign
is found, often scats, tracks, and
other evidence can be found with it
to corroborate the species identification. By itself, feeding characteristics may or may not provide strong
evidence. In this scheme, it is
assigned a low point score and
requires at least two more pieces of
evidence with it to confirm a
species presence. A photograph or
sample of the feeding sign should
be sampled to verify the observation.
Beds, Dens, Nests, Trails - 3

points.
For many species of mammals,
beds (resting places), den holes,
nests, wallows, and trails can be
characteristic to the species. The
location of these features, sizes,
shapes, and dirt placement are often
useful as evidence to verify a
mammal is present; e.g., tapir trails
and beds. Often bone, hair, scats,
and tracks will be found at these
features and will corroborate presence and refined identifications. By
themselves, these features do not
have the strength to support a
confirmation and require at least
one other corroborative piece of
evidence.
Photos and/or written descriptions should be obtained to verify
this evidence on significant species
or where it is central to the identification.
Vocalizations and Odors - 5
points.
Many large mammals communicate with distinctive vocalizations
and odors (mustelid musks and
peccary musks). These smells and
sounds often are excellent evidence
of a species presence and are quite
impressive; e.g., jaguar territorial
and mating calls. Monkey vocalizations are characteristic to the
species and very good evidence of
presence without other evidence.
However, monkey sounds can be
mimicked by birds, and people
which could render an incorrect
determination.
Vocalizations can be aggressively invited which makes them
very useful for a monitor, and can
be used to bring the mammal into
323

sight or photographic range for
corroborative identification.
When possible, record vocalizations for verification and play back
for calling.
Odors are occasionally useful
for recognizing species; e.g.,
mustelids musks, civit musks and
opossum musks can be used to
identify presence. In this scheme,
vocalizations and/or odors rate 5
points and would require at least
one other piece of quality evidence
to confirm a species presence.
Identification by Local Residents - 5 points.
The validity of an identification by a local resident can vary
from certain to false, depending on
the person chosen to do the identification. Choose a qualified hunter,
trapper, naturalist, or similar person
with interest and experience with
large mammals. Often, local people
will report what they think is
wanted. It is particularly true when
there is a language barrier and they
want to appear to be knowledgeable
beyond their capabilities. Do not be

afraid to ask them what their limitations are in identifying these animals, and factor that into the
formula. When possible, test them
in the field to ascertain how reliable
they are.
Our field guides were very
talented and credible, but misidentified some tracks. They had a
tendency to lump tracks into raton
(rats), anouje (agouti), and tigrillo
(small cats), which were obviously
several different species of about
the same size.
After consultation with two
mammalogists and one ecologist, it
was decided that at least one other
confirming piece of evidence was
needed to verify a confirmation of
presence for a species identified by
local residents.
This system assumes that only
conclusive sign is used in constructing the occurrence index. The
conservative point assignment
requiring one or two kinds of
corroborating evidence protects
Table 3. Example of Biodiversity Occurrence Index

Evidence
Specimen collected
Specimen observed and verified by a witness or evidence
Tracks
Feces (scats)
Hair
Bone
Feeding characteristics
Beds, dens, trails, and nests
Vocalizations and odors
Identification as occurring at the site by local people
Total possible points
Point total necessary to confirm the presence of a species
at the time of monitoring
324
Points
10
10
5
3
5
5
2
3
5
5
53
8

against errors. An example of the
Biodiversity Occurrence Index is
shown in Table 3.
The occurrence index is calculated by adding up the points of
evidence found for each species.
When the total reaches 8, the
species is declared present at that
monitoring place and time period.
The higher the score, the more
verification of its presence was
collected and more certain the
record. Any type of evidence is
counted only once for the occurrence index, even though that
evidence may have been collected
many times (Table 4). The balance
of the species and occurrence index
number calculations are on file,
available upon request. All index
values are listed in Table 1.
Discussion
Strengths
It is direct, simple, and uses an
accumulated evidence score to
verify a species record. The system
allows for some error and still
develops a verified record and
indicates its strength. The numerical score provides a basis for comparison of the strength of the
records between locations and time
within the same species.
Weaknesses
The system requires a lot of
woodsmanship and knowledge of
the species in order to produce the
evidence required. The score indicates only occurrence, not abundance. Index numbers are comparable only within the same species
between places and time, not
between species. For example, the
red brocket deer was much more
abundant than tapir at San Martin,
but the occurrence scores were the
same.
This index is offered as a
suggested tool that can and should
be modified to make it useful to
future researchers of large mammals.
The identification of species
present and the determination of
changes in the population status of
the species over time are both
essential to monitoring of medium
and large mammals. Based on those
two determinations, further judgements may be made on what caused
the changes.
325

Table 4. Biodiversity Occurrence Index value for each of the
medium and large mammal species found at the Lower
Urubamba region, Cusco, Per
San Martin-3
Species
326
Cashiriari-2
Brazilian Tapir
Collection
Observation
Tracks
Scats
Hair
Bone
Feeding
Beds and dens
I.D. by locals
Totals = Occurrence Index
0
10
5
3
0
0
2
3
0
5
28
0
0
5
0
0
0
0
3
0
5
13
Tayra
Collection
Observation
Tracks
Scats
Hair
Bone
Feeding
Beds and dens
I.D. by locals
0
0
5
0
0
0
0
3
0
5
13
0
0
0
3
0
5(skull)
2
3
0
5
18
Common Opossum
Collection
Observation
Tracks
Scats
Hair
Bone
Feeding
Beds and dens
Vocalization and odors
I.D. by locals
10
10
5
0
0
0
2
0
0
5
32
0
0
0
0
0
0
0
0
5
5
10

Table 4. Biodiversity Occurrence Index value (Cont.)
Species
San Martin-3
Cashiriari-2
Jaguar
Collected
Observed
Tracks
Scats
Hair
Bone
Feeding
Beds and dens
I.D. by locals
0
0
0
0
0
0
0
3
0
5
8
0
0
5
3
0
0
0
3
5
5
21
Puma
Collected
Observed
Tracks
Scats
Hair
Bone
Feeding
Beds and dens
I.D. by locals
0
0
0
0
0
0
0
0
0
5
5
0
0
5
3
0
0
0
3
5
5
21
Ocelot
Collected
Observed
Tracks
Scats
Hair
Bone
Feeding
Beds and dens
I.D. by locals
0
0
5
3
0
0
0
3
0
5
16
0
10
5
0
0
0
0
3
0
5
23
327

Table 4. Biodiversity Occurrence Index value (Cont.)
San Martin-3
Species
328
Cashiriari-2
Collared Peccary
Collected
Observed
Tracks
Scats
Hair
Bone
Feeding
Beds and dens
I.D. by locals
0
10
5
0
0
0
2
3
5
5
30
0
0
5
0
5
5
2
0
5
5
27
Common agouti
Collected
Observed
Tracks
Scats
Hair
Bone
Feeding
Beds and dens
I.D. by locals
10
0
5
0
0
0
2
3
0
5
25
0
10
5
0
0
5
2
3
0
5
30
Red brocket deer

Collected
Observed
Tracks
Scats
Hair
Bone
Feeding
Beds and dens
I.D. by locals
0
10
5
0
0
0
0
3
5
5
28
0
10
5
0
5
0
0
3
0
5
28
Appendix 2. Biodiversity Abundance Index.

Finding and identifying the
large mammals is relatively easy.
Counting them and tracking number
changes is difficult for individual
species. Trying to do so on a wider
scale is at best a guess, the quality
of the guess is based on the experience and knowledge of the researcher.
Generally speaking, the best
evidence animals leave, which
indicates numbers, is an accumulation and interpretation of the sign
of individual animals. In the absence of a count, which is rarely
possible, it is the only alternative. It
logically follows that the higher the
number of animals present at a
location, the more sign they will
leave. It also logically follows the
amount of sign can be counted and
will be proportional to the number
of animals at a location. It is probable that that number of sign events
is characteristic of each species and
therefore valid only between members of that species. There are
limits, of course, to comparing
apples to oranges.
If this logic is sound, then an
abundance index can be constructed
from the occurrence index evidence. The key to relative abundance would be in the care and
accuracy that is used to discriminate out the duplicative evidence,
or counting several times the sign
left by a few individuals. That can
be done in a number of ways,
including measuring differences in
the size, characteristics, how far
apart the collection occurs in time
and distance, removing individuals,
the sign features, etc.
For the purposes of monitoring
at the well sites, an abundance
index was constructed using the
same point system as for the occurrence index, minus the points for
the local identification. Each
evidence type was multiplied by the
number of times the evidence was
found where it was likely an independent, non-duplicating record.
These index numbers are found in
Table 1 as the two columns on the
right.
An example of how the index is
constructed follows (Table 5).
This proposed abundance index
has many features about it that may
or may not be useful. Basically, a
high index number means there was
a lot of activity of that mammal on
the monitored area; a low index
number means there was little
activity. With some knowledge of
mammal home-range size based on
size, food habits, radio telemetry,
ear-tagging, and other tracking
methods for similar mammals in
other parts of the world, an educated guess can be made on the
number of individuals of the
species using the monitored area.
329
Appendix 2. Biodiversity Abundance Index (Cont.).

Table 5. Biodiversity Abundance Index of medium and large mammals in
the Lower Urubamba region, Cusco, Per
San Martin-3
Species
330
Cashiriari-2
Brazilian tapir
Collection
Observed
Tracks
Scats
Hair
Bone
Feeding
Beds, dens, trails, nests
Vocalization & odors
Totals
0
10 x 1 = 10
5 x 7 = 35
3x3=9
0
0
2x1=2
3x3=9
0
65
0
0
5 x 3 = 15
0
0
0
0
3x2=6
0
21
Tayra
Collection
Observed
Tracks
Scats
Hair
Bone
Feeding
Beds and dens
Vocalization & odors
Totals
0
0
5 x 2 = 10
0
0
0
0
3x1=3
0
13
0
0
0
3x1=3
0
5x1=5
2x1=2
3x1=3
0
13
Common opossum
Collection
Observed
Tracks
Scats
Hair
Bone
Feeding
Beds and dens
Vocalizations & odors
Totals
10 x 2 = 20
10 x 1 = 10
5x1=5
0
0
0
2x1=2
0
0
37
0
0
0
0
0
0
0
0
5x1=5
5

Table 5. Biodiversity Abundance Index of medium and large mammals (Cont.)
Species
San Martin-3
Cashiriari-2
Collared peccary
Collection
Observed
Tracks
Scats
Hair
Bone
Feeding
Beds and dens
Totals
0
10 x 4 = 40
5 x 40 = 200
0
0
0
2x3=6
3x2=6
5x1=5
257
0
0
5 x 5 = 25
0
5 x 6 = 30
0 (bones & hair duplicate)
2x1=2
0
1x1=1
58
Common agouti
Collection
Observed
Tracks
Scats
Hair
Bone
Feeding
Beds, dens, & trails
Totals
10 x 1 = 10
0
5 x 20 = 100
0
0
0
2 x 2= 4
3 x 20= 60
0
174
0
10 x 2 = 20
5 x 8 = 40
0
0
5 x 1=5
2 x 3= 6
3 x 3= 9
0
80
Red brocket deer

Collected
Observed
Tracks
Scats
Hair
Bone
Feeding
Beds, dens, trails, wallow
Vocalizations & odor
Totals
0
1 0 x 1= 10
5 x 8= 40
0
0
0
0
3 x 1= 3
5 x 1= 5
58
0
10 x 1= 10
5 x 8= 40
0
5 x 1= 5
0
0
3 x 3= 9
0
64
331

Table 5. Biodiversity Abundance Index of medium and large mammals(Cont.)
San Martin-3
Species
Jaguar
Collected
Observed
Tracks
Scats
Hair
Bone
Feeding
Beds, dens, trails
Vocalization & odor
Totals
Puma
Collected
Observed
Tracks
Scats
Hair
Bone
Feeding
Beds, dens, trails
Vocalizations & odor
Totals
Ocelot
Collected
Observed
Tracks
Scats
Hair
Bone
Feeding
Beds, dens, trails
Totals
332
Cashiriari-2
0
0
0
0
0
0
0
3 x 1= 3
0
3
0
0
5 x 3 = 15
3 x 2=6
0
0
0
3 x 3=9
5 x 1=5
35
0
0
0
0
0
0
0
0
0
0
0
0
5 x 3=15
3 x 2=6
0
0
0
3 x 3=9
5 x 2=10
40
0
0
5 x 3= 15
3 x 1= 3
0
0
0
3 x 3= 9
0
27
0
10 x 1= 10
5 x 4= 20
0
0
0
0
3 x 4= 12
0
42
Appendix 3. Visual survey method.

Survey routes were walked
each day at least once, often twice,
from 7-9:30 a.m. and from 2:305:30 p.m. Frequent stops were
made to listen and observe mammal
activity. When prospective canopy
travel routes and well-used surface
trails were approached, the researchers stopped and carefully
observed the location for five to 10
minutes. The strategy was effective
for spotting monkeys and calling
them into identification and photography range. Two common agoutis,
one red brocket deer, two Amazon
dwarf squirrels, and one southern
Amazon red squirrel were observed
using this survey method.
As a general biodiversity monitoring method, this survey technique offers a low return on the
time invested, but is simple, convenient, and can be done as other
survey methods are being used.
Appendix 4. Vocalization and

calling survey method.
During the daily walks on the
survey routes, the researchers
would stop at locations which
offered some visibility on the forest
floor or canopy, and call using
monkey, tapir, jaguar, javelina, and
various rabbit, mouse and fawn deer
distress cries. About 4 p.m., when
other survey methods had been
completed, stands were chosen and
called for 20 minutes with a
CRITRCall to see which species
would respond.
One night, a calling and spotlight survey was attempted. Light
penetration was nil and the project
was abandoned when a jaguar
vocalized in response to a fawn
bleat at a range of about 500 m.
Without protective firearms, clothing, or escape alternatives, it was
decided that night calling in jaguarpopulated areas might be less than
intelligent.
The local guides were accomplished at mimicking monkey
vocalizations. They would hear
monkey foraging noises and vocalizations, then mimic the monkeys.
In 10 out of 12 occasions, the
monkeys responded and approached
the researchers for observation and
photographs. The technique is
practical if the local guide who is
able to accomplish the vocalizations
is employed. Or, the vocalizations
of various monkeys could be recorded for play-back on a cassette
player and used to call monkeys if
guides were unavailable.
Calling with cassette tape
recordings of prey vocalizations
from protective tree stands or other
cover would likely be effective for
day or night calling of omnivores
and predators in some areas.
333
Appendix 4. Vocalization (Cont.).

The guides had thoroughly
mastered various calls with their
voices. They also used them quite
frequently and naturally as a matter
of course in hunting. The guides
were entertained by the researcher
using commercially made American
predator calls and wanted them for
their use.
Calling and vocalization surveys have a great deal of potential
for many forest animals. The techniques should be developed further.
Ocelot, jaguar, white-lipped
peccary, collared peccary, blackchested mustached tamarin, dusky
night monkey, common squirrel
monkey, brown and white-fronted
capuchin monkeys, black spider
monkeys, and red howler monkeys
were observed using various calling
methods.
334
Appendix 5. Scent-post survey

method.
The scent-post survey utilized
was a modification of the survey
method described in the Indices of
Predator Abundance in the Western
United States1 . Rather than a
distance-defined transect route,
scent stations were placed at optimum locations to record the maximum number of species. Locations
selected included the edges of
ecotones, at intersections of manmade and wildlife trails, on saddles
between hill peaks, at stream
intersections, at intersections of
wildlife trails, and where geological
features concentrated mammal
travel lanes.
One to three circles, approximately one meter in diameter were
raked into the soil, spaced about 5
m apart in close proximity to the
mammal travel path. In the center
of the circle, a Q-tip cotton swab
was placed which contained the
amount of commercial furbearer
trapping lure the cotton would hold.
A selection of Carman brand lures
were used. See the list at the end of
this appendix which includes animals which were attracted to each
lure.
Each morning, between 7 and
9:30 am, these scent posts were
visited and the tracks identified and
recorded for each station. Stations
were swept clean of tracks and
lures replaced when needed. Unless
the lure was taken, they were left
for four days before being renewed
or replaced. Thirty-six circles were
placed at San Martin-3; 28 circles
were placed at Cashiriari-2. These
resulted in 80 track observations.
As a result of previous experience with trapping lures, researchers have observed that some species
Appendix 5. Scent-post survey method (Cont.).

are neophobic and require three to
four days to become acclimated to
foreign odors. They will approach
and display various behaviors on
the lures at that time. Each species
seems to react differently to each
lure. Some species behaviors are
fairly predictable, some are not.
The responses to these scent
stations and lures were relatively
frequent, intense, and from a large
variety of mammals. It was the best
method for making reliable observations quickly, at low cost, and with
little time invested.
Scent-post survey methodologies should be thoroughly explored
for rain forest applications. This
method offers the best potential for
statistically quantifiable data for
statistical reliability tests for many
species.
Generally, the species which
responded best to the scent-post
surveys were the ones which are
particularly hard to observe in the
rain forest: the felids, canids, and
procyonids. Tapirs, agouti, and deer
were also frequent visitors. Species
adapted to the canopy are not
generally subject to this survey
method; however, adaptation could
be made to get the scents into
canopy tracking stations. Rodents
responded to many lures, but
tracking substrate did not allow for
specific identification.
The variety of olfactory stimulants is very broad. It is possible
that every mammal exposed would
respond to some scent material. It
would also be possible to adapt the
scent applications to a few species
of selected taxa for selective observations. The knowledge of how to
make scents selective to mammal
families is available for rainforest

applications.
Olfactory scents often lack the
ingredients which attract arthropods
which degrade normal food type
meats, fruits, and other bait materials. The longevity of Carmans
scents was commonly four days or
more during the present survey. The
cost of applications for Q-tips and
scent materials was less than $0.10
each.
Anonymous. 1981. Indices of
Predator Abundance in the Western
United States. U.S. Fish and Wildlife
Service, Washington, DC..
1
335

Table 5. Results of the scent-post survey method at the Lower Urubamba
region, Cusco, Per (* lures supplied by Carmans Superior Animal Lures,
RR 2, Box 182, New Milford, PA 18834; ** not used at the site)
Species attracted to the lure*
Carmans Canine Call Lure*
Brazilian rabbit
Paca
Green acouchy
Brazilian tapir
Ocelot
Puma
Jaguar
Short-eared dog
Grey Fox Gland Lure
Common agouti
Pros Choice
Common opossum
Ocelot
Green acouchy
Puma
Jaguar
Brazilian tapir
Brazilian rabbit
Common agouti
Paca
Trophy Deer Lure
Green acouchy
Common agouti
Red brocket deer
Puma
Jaguar
336
Number of visits
San Martin-3
Cashiriari-2
2
2
2
2
2
0
0
0
5
1
0
0
3
2
1
1
1
2
**
**
1
4
1
0
0
1
1
1
0
0
5
0
2
1
1
0
0
1
1
1
4
0
0
0
0
4
2
1

Table 5. Results of the scent-post survey method (Cont.)
Species attracted to the lure
Brazilian rabbit
Common agouti
Common opossum
Green acouchy
Puma
Jaguar
Ocelot
Bobcat Gland Lure
Red brocket deer
Puma
Jaguar
Ocelot
Wind River Beaver Lure
Puma
Jaguar
Rodents
Magna Gland Lure
Rodents
Silent Partner
Rodents
Trails End
Rodents
Raccoon #1
Common opossum
Rodents
Raccoon #2
Still Water
Rodents
Three Rivers
Rodents
Midnight
Rodents
Red Fox Gland Lure
Rodents
Green acouchy
Number of visits
San Martin-3
Cashiriari-2
1
1
1
1
0
0
0
**
**
**
**
0
0
many
**
many
many
1
many
2
many
many
many
many
1
337
Appendix 6. Track, scat, and sign survey methods.

During the survey period, the
biodiversity researchers were encourage to assist each other in
making observations and gathering
evidence of occurrence of species
from their specialties. This was very
helpful to all researchers involved.
Evidence obtained includes several
skulls, bones, hair, and visual
observations of mammals. When
possible or necessary, the lead
specialist would then follow up to
verify the observation.
In addition, the guide and
researchers would carefully inspect
the survey routes for fresh tracks,
feces, urine stations, feeding,
digging, or territorial markings,
animal parts, and other tangible
evidence that mammal species were
present. These materials or signs
were photographed or sampled for
verification of the observations.
The various evidence was identified
and assigned a numerical value and
put into the indices as described in
Appendix 1.
This method was also very
productive, particularly for finding
evidence that was not available
from other methods. This method
was used to document kinkajou,
tayra, southern two-toed sloth,
puma, jaguar, crab-eating raccoon,
nine-banded armadillo, collared
peccary, and contributed to a
confirmed occurrence index for
many other species.
The lack of track, bone, hair,
and sign keys for South American
mammals is a handicap to the
accuracy of biodiversity monitoring
and would be a worthy activity for
some funding agency to sponsor.
Track, scat, and sign surveys
presume that the researcher has
338
expertise in sign reading. Rain

forest evidence is difficult to find
and identify because the durability
of scats, body parts, and physical
disturbances is short-term due to
scavengers, insects, and rain. These
factors, however, insure that whatever evidence is found is rather
recent in origin.
Appendix 7. Trapping survey method.

Emphasis for this preliminary
monitoring effort was intentionally
placed on non-lethal sampling
methods for the following reasons:
* Larger mammal numbers are
much lower than rodents and bats
and recruitment is much less, so
removal of individuals has a greater
impact on future monitoring results.
It is best to leave them in the
population if other means are
available for monitoring.
* Most well-known large
mammals are common in collections, museums, and zoos, so there
is a diminished need.
* Larger mammal collections
require much more complicated
logistical support for preservation
of skins and utilization of specimens.
* Larger mammals are used as
food resources by local communities. It is best to have their expressed consent and participation in
collecting and then keep it to a
minimum.
* Collecting large mammal
specimens requires large blocks of
time.
* Collection permits are in
process but have not been formally
issued.
* A specific goal of the researchers was to develop a process
which would minimize the need for
collections of large mammal specimens for all of the reasons listed
above.
* Some of the public is sensitive to unnecessary collections of
large mammals.
The researchers chose to take
two sizes of Conibear traps which
are quick-kill, body-gripping traps
(220s and 110s), and four sizes of
adaptable snares including sizes #0,
#1, #2, and #4 Gregerson brand

snares. These snares were selected
for their versatility. They can be
used as humane killing devices, hair
collectors, or live-capture-andrelease devices by various and
simple modifications. Further, the
places they are set, height off the
ground, loop size, and sensitivity
with which they are set will provide
various elements of evidence about
species movement on trails and
around scents and baits.
Care must be taken to set them
selectively so unwanted animals are
not accidentally taken; e.g., puma in
a snare set for a paca. Reasons
include the following:
* short life of available bait
materials,
* low densities and long
intervals between trail use by
resident animals,
* hundreds of trail choices,
* selective placement meant
avoiding the best locations for
setting, and
* rain and corrosion of the
trap surfaces affected trap trigger
sensitivity.
In all, one common agouti, and
one common opossum were collected in these devices. Two common agoutis, one paca, and one
ocelot were missed by insensitive
snares failing to close. The
Conibears2-220s, and 6-110s
were set for 176 trap nights and
yielded one common opossum.
Snares were set for 1012 trap nights
and yielded one common agouti and
four near misses. One common
opossum was caught in a National
live trap associated with the small
mammal survey.
Gin traps or foothold traps,
combined with the scents identified
339
Appendix 7. Trapping survey method (Cont.).

as attracting target animals, were
the obvious strategy to use for
efficient trapping. However, no
foothold traps were brought for the
survey. In the researchers opinion,
foothold traps would have captured
each species which responded to
the scent posts and are listed in
Appendix 5.
Ensuring the environment is
maintained in great condition after
340
natural resources extraction is

critical for these people and equally
important for all of us.
Long-term
Monitoring
of Medium
and Large
Mammals
in the Lower
Urubamba
Region
Major Boddicker
Rocky Mountain Wildlife
Enterprises
Introduction
While participating in and then

helping to analyze the results of the
initial biodiversity monitoring
surveys in the Lower Urubamba
region during May 1997, four tasks
were identified for the next phases
of the project. They are discussed
in this paper. First, however, is a
short summary of the thinking that
led to identification of the tasks.
The present monitoring activities track what happens to the
ecosystem after planning is completed and gas exploration begins.
That is important. To ensure that
major problems resulting from
future exploration and development
do not occur, baseline monitoring
should be accomplished along with

the planning effort. Most likely, any
significant problems likely to occur
will be the result of placement of
pipelines and facilities. The pipeline
route from the gas field to Lima and
location of the gas processing
plants, pumping stations, pipeline
maintenance facilities, and access
are all critical. Biodiversity monitoring will provide useful information
so that decisions about routing and
locations can be adjusted to avoid
ecological problems.
Past experience has shown that
a lack of biodiversity monitoring
can result in ecological disasters.
Gas exploration and development in
the Lower Urubamba may have
such negative potential. Careful
research focused on preventing
damage will likely be a good investment for Shell Prospecting and
Development (Per) B.V., Per, and
the environment.
Based on such reasoning, the
following four tasks have been
delineated for large mammals during
the next phases of the Lower
Urubamba biodiversity assessment
and monitoring project. While the
341
focus is on large mammals, the

basic rationale for the tasks holds
for other taxa in the study area.
A. Conduct monitoring of
mammal diversity at San Martin-3,
associated with gas production.
B. Conduct biodiversity monitoring along the proposed pipeline
routes from Urubamba to Lima and
C. Develop training manuals
and support materials to assist
future researchers in their environmental assessment efforts; e.g.,
compile track, scat, and sign keys
and biomonitoring and sampling
technique manuals for South American mammals.
D. Train Peruvians and other
South American researchers in
for large mammals. (The level of
technology and knowledge for
large-mammal monitoring in South
America is not current with the
developments in these areas that
have taken place over the past
decade.)
Description of
Biodiversity
Monitoring Tasks
Conduct monitoring of mammal diversity at San Martin-3,

associated with natural gas production.
Explanation: This activity is
designed to monitor the large
mammal populations at San Martin,-3, Cashiriari-2, and other well
sitesincluding the test plot areas
on the Urubamba Riverover the
next five years. Researchers will
spend at least 15 days at each site
during the same time of each year.
1. Time frame: five years.
342
2. Staff needed: two field

researchers, one supervisor/researcher, two support staff (clerical).
3. Duration of field activities:
three months annually (April June).
4. Activities:
* Monitor large mammal populations;
* Identify species;
* Develop population estimates;
* Assess changes;
* Determine if changes are
occurring;
* Make remedial recommendations;
* Develop progress reports and
final reports.
Conduct biodiversity monitoring along the proposed pipeline
routes from Urubamba to Lima and
Explanation: This task will
monitor large mammals in the
different life zones along the proposed pipeline route to Lima. The
routes would be aerially
transversed, and potential areas for
monitoring would be identified.
Those areas would be checked for
large mammal population parameters. The objective is to recommend pipeline placement in habitat
that would have the least negative
impact on sensitive species; e.g.,
speckled bear and jaguar. It is
possible that shifting the pipeline
even just a few meters would help
avoid destruction of features that
are important to sensitive species.
Once the pipeline is in place, the
sites will be monitored to track
long-term impacts.
1. Time frame: five years.
2. Staff needed: three field
researchers and two support staff.

three months annually or until
completed (August - October).
4. Activities:
* Monitor sample life-zone
areas along proposed pipeline routes
to identify any large mammals that
are present;
* Estimate populations;
* Recommend pipeline placement so as to decrease the potential
for negative impacts;
* Submit progress reports,
recommendations, and final reports
to assist Shell Prospecting and
Development (Per) B.V. in planning the pipeline route;
* Compile samples of mammal
specimens for Peruvian and U.S.
natural history museums.
Develop training manuals and
support materials to assist future
researchers in their environmental
assessment efforts; e.g., compile
track, scat, and sign keys and
biomonitoring and sampling technique manuals for South American
mammals.
1. Time frame: three years.
2. Staff needed: one researcher/writer, one typist/word
processor/editor.
one month annually (January February).
4. Activities:
* Produce bone, hair, tracks,
scat, and sign manual for South
American mammals and a
biomonitoring technique manual to
supplement those already available;
* Extend and update the sampling technique manual.
Train Peruvians and other
South American researchers in
for large mammals (The level of
technology and knowledge for
large-mammal monitoring in South
America is not current with the
developments in these areas that
have taken place over the past
decade).
Explanation: South American
researchers would accompany and
assist the contracted staff during
various field projects and follow-up
laboratory work to learn large
mammal monitoring techniques.
343
344
Ecto-and
Endoparasites
in Mammals:
Biodiversity
Assessment
in the Lower
Urubamba
Region
Ricardo Guerrero
Instituto de Zoologa Tropical,
Facultad de Ciencias,
Universidad Central de Venezuela
Introduction
This paper describes the most

significant aspects of the biology of
the distinct groups of parasites
found in the Lower Urubamba
region. The systematics of the
species sampled are underway, to
the point that the absolute and
relative abundances of the parasites
found in each of the mammal
species sampled can be estimated.
This information constitutes the
baseline for long-term monitoring
studies.
The ecology and systematics of
the parasites from the study area
will provide valuable information. I
depart from the principle that the
parasite fauna is at the top of the

ecosystem trophic chain. The study
of parasites has the advantage of
detecting early changes in the
ecology of the animal species where
they live. For example, animals
under stress may suffer higher rates
of parasitism.
The identification of parasites
and their significance have several
implications. On one hand, it is
very important to identify to the
species level all parasites associated
to each of the taxa studied. This is
a process that takes time since
parasites tend to be very small and
similar in morphology. Many parasites have converged in their morphological adaptations to the
parasitic life. On the other hand,
many of the parasitic species will
be new to science, building knowledge so that comparisons among
species and different locations are
possible.
Methods
As part of the biodiversity

region, I sampled the parasites that
345
Mammals
with low
mobility,
including
species
that live on
the forest
floor and
in semiaquatic
environments and
arboreal
species
were
sampled
for
parasites
346
live in the mammal communities at

San Martin-3 and Cashiriari-2. To
maximize data gathering on the
parasites in the region, I sampled
both the parasites that live on the
external surface of the body (ectoparasites) and those that survive in
the internal organs of the mammal
(endoparasites). I sampled a number
of species of bats, rodents, and
marsupials, using the following
criteria to select the individuals that
were sampled for parasites.
Mammals with low mobility,
including species that live on the
forest floor and in semi-aquatic
environments and arboreal species. I
sampled species of rodents that
ranged from granivores (animals
that feed on plant seeds) to omnivores (animals with a wide variety
in diet) as well as marsupial species
that are mostly omnivores and
insectivores.
Mammals with high mobility,
including bats, the only group of
flying mammals. Animals in this
group have relatively high mobility
and dispersion, allowing them to
seek out those places that they
consider optimal for survival. This
means that some of these species
can abandon a site that has been
disturbed, while others can colonize
disturbed areas. Within this group
are animals exhibiting a variety of
feeding habits, which is a necessary
component in the biology of the
endoparasites that will reside in the
animals. The ability to disperse is
important to the biology of ectoparasites; where the animals rest
influences their chances of being
infected by the ectoparasites. As an
example, bats can be frugivores,
carnivores, insectivores, hematophagous, and nectarivores (see
Wilson et al. this volume) and can
roost in closed, semi-closed, and

opened areas.
Once the mammal was
sampled, it was immediately placed
in a plastic bag so the ectoparasites
could not escape. The plastic bags
and the mammals fur were then
inspected for ectoparasites under
the dissecting stereo microscope. I
inspected for internal parasites once
the specimens was processed,
looking for both mature and immature parasites in the organs. I paid
special attention to the digestive
system where many endoparasites
live. I did not consider protozoans
(unicellular organisms that can
cause diseases such as malaria) in
the study.
Results and
Discussion
At San Martin-3, I sampled 39

mammal species for ectoparasites
and 41 for endoparasites; at
Cashiriari-2, I sampled 45 species
of mammals for both groups (Appendix 1). The total number of
different species sampled was 65,
higher than the 53 species sampled
in a study at nearby Pakitza, Manu,
Per, over a similar period of time
(Guerrero, 1996). The total number
of individuals sampled in the Lower
Urubamba study area was 332. I
found that most of them had ectoand endoparasites. Only 1.3% of
the sample was not infected. I
recorded close to 800 parasites for
San Martin-3 and Cashiriari-2; it
appears that at least 25 are new
species.
The following endo- and ectoparasites were found, listed in order
of most abundant and frequent to
least abundant and frequent: 1)
endoparasitesNematoda
(Trichostrongylida, Oxyuridae,
Ochocercidae, Seuratidae,
Rictularidae, and Muscipedidae),
Trematoda, and Pentastomida
(larvae); 2) Trombiculidae,
Laelapidae, Streblidae,
Atopomelidae, Spinturnicidae,
Labidocarpidae, Macronysidae,
Mallophaga, Anoplura, Myobiidae,
Ixodidae, Nycteribiidae,
Spalaerorhynchidae, Polyctenidae,
Siphonaptera, and Argasidae.
The following conclusions may
be drawn from these findings.
The levels of parasitism and
the type of parasites are similar to
those in other natural Amazonian
environments such as the Manu
region and several Venezuelan
localities. Thus, parasite communities have structural qualities and
quantitative similarities.
The abundance and diversity of
the Trombiculidae and the
Trichostrongylida show that the
edaphic conditions at San Martin-3
and Cashiriari-2 have not been
altered so as to affect these groups.
This is important because it is
physical and chemical factors that
allow establishment, survivorship,
and development of the larvae of
these groups. This conclusion is
supported by the presence of
Siphonaptera and several species of
mites (Ixodidae and Argasidae) that
also have larval phases in the soil.
The large abundance of Trematoda in bats, rodents, and marsupials indicates that the river and
stream conditions in the study areas
are still unaltered. Trematoda larvae
need an intermediary host (usually a
snail) to complete their life cycle.
Snails are very sensitive to changes
in water quality (see Ramirez and
Cordova, this volume), so the
abundance and diversity of the
Trematoda indicates that the community of molluscs is healthy;
therefore, it is not likely that the

physical and chemical properties of
the water have been significantly
altered.
Groups such as Filarias,
Rictularids, and others are also well
represented. Parasites in these
groups need an hematophagous
(blood sucking) insect as a vector
for transmission. Their abundance
indicates that the entomofauna of
the forest has not been affected.
To this point, then, knowledge
of parasites at San Martin-3 and
Cashiriari-2 indicates that the forest
has not been functionally altered.
The material sampled in this study
will serve well as a baseline for
short- and long-term monitoring of
the biotic conditions in the Lower
Urubamba region.
The levels
of
parasitism
and
the type of
parasites
are similar
to those in
other
Amazonian
environments.
References
Guerrero, R. 1996. Estudio de los

Ectoparasitos de los murcielagos
de Pakitza, Parque Nacional Manu
(Per). Pages 643-657 in Manu:
(D. E. Wilson and A. Sandoval,
eds.) Smithsonian Institution,
Washington, DC.
347
Appendix 1. List of mammal species sampled for ecto- and endoparasites

in the Lower Urubamba region.
San Martin - 3
Ectoparasites Endoparasites
Marsupialia
Monodelphis brevicaudata
Chiroptera
Peropteryx macrotis
Saccopteryx bilineata
Saccopteryx leptura
Chrotopterus auritus
Micronycteris minuta
Mimon crenulatum
Phyllostomus hastatus
Phylloderma stenops
Tonatia bidens
Tonatia brasiliensis
Tonatia sylvicola
Anoura caudifer
Anoura latidens
Choeroniscus minor
Carollia brevicauda
Carollia castanea
Rhinophylla pumilio
Artibeus lituratus
Artibeus obscurus
Chiroderma villosum
Dermanura anderseni
Dermadura cinerea
348
3
+
+
+
25
3
+
+
+
25
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
Cashiriari - 2
1
39
+
+
+
+
+
39
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
Appendix 1. List of mammal species sampled for parasites (Cont.).

San Martin - 3
Sturnira lilium
Sturnina magna
Sturnina tildae
Uroderma bilobatum
Vampyressa bidens
Vampyressa macconnelli
Vampyressa pusilla
Vampyrodes caraccioli
Diphylla ecaudata
Myotis nigricans
Myotis riparius
Myotis ruber
Myotis simus
Eptesicus brasiliensis
Molossus ater
Molossus molossus
Promops centralis
Rodentia
Neacomys spinosus
Nectomys squamipes
Oecomys bicolor
Oecomys concolor
Oecomys roberti
Oryzomys capito
Oryzomys nitidus
Oryzomys yunganus
Oxymycterus inca
Dasyprocta punctata
Proechimys spp
TOTA L
+
+
+
+
+
+
+
+
+
+
13
+
+
+
+
+
+
+
+
+
+
+
+
+
41
11
+
+
+
+
+
+
+
+
+
+
+
39
Cashiriari - 2
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
5
+
+
5
+
+
+
+
+
+
45
+
45
349
350
Ectoparasites
in Mammals:
Biodiversity
Assessment
in the Lower
Urubamba
Region
John Chavez
Mayor de San Marcos
Introduction
Per has a great diversity of

mammals, including bats (152
species) and rodents (152 species;
Pacheco et al. 1995). Mammal
species play vital roles in the control and maintenance of the ecosystems. In particular, bats act as
pollinators and seed dispersors and
are known for their importance in
medical epidemiology (see Wilson
et al. this volume). Rodents serve
to spread micorrhyza-ridden fungus,
furthering the crucial association
between the roots of vascular
plants and specialized fungi. Rodents also help control certain
species of insects that serve as a
baseline of the nutritional chain of
vertebrates. However, little is
known about the endo- and

ectoparasites that survive on bats
and rodents (Kohls et al. 1969;
Radovsky and Furman 1969;
Brennan 1970; Goff 1984; Elliot et
al. 1985; Orozco 1990; Need et al.
1991). Such information is needed
to increase understanding of how
these species are related and to gain
more knowledge about the role that
fleas, ticks, mites, bugs, and
anopluros play in transmitting
disease (Goirirossi 1950; Parker et
al. 1954; Kohls 1955; Philip 1961,
1968; Hoogstaal 1961; Horsfall
1962; Muir-Head-Thomson 1968;
Meyer 1957; Jellison 1959;
Macchiavello 1959; Haas 1959;
Herms and James 1961; Jenkins
1964; FAO-WHO 1967; Audy
1968; Maa 1971; Marshall 1976).
Among the great diversity of
parasites in mammals, there are
several important groups including
the Streblidae. This group has
blood-feeding flies that are specific
to bats. The Spinturnicidae mites
are specific to bats as well as the
Nycteribiiadae, Macronyssidae,
Labidoarpidae, Myobiidae Trombiculidae, Laelapipdae, and
351
The bat
Platyrrhinus
brachycephalus
(subfamily
Stenodermatinae)
and the
mouse
Oryzomys
nitidus were
the most
common
species
checked for
parasites.
Argasidae (Anciaux de Faveaux

1971-1976; Kohls 1962; Wenzel et
al. 1966; Wenzel 1976; Webb and
Loomis 1977; Marshall 1981-1982;
Nutting 1985; Guerrero 1992;
Guerrero 1993). A few incidental
and routine studies in Per have
helped describe new species, primarily on ectoparasites of bats and
one study for the rodent group
Siphonapera. But for the most part,
little has been done to study the
parasite fauna in Per. More research is especially needed for
groups such as the Polycteniade and
the Labidocarpidae (Veshima 1972;
Guerrero 1992)
This paper presents results
from an initial survey conducted in
the vicinity of Shell Prospecting
and Development (Per) B.V. well
sites San Martin-3 and Cashiriari-2
in the Lower Urubamba region,
Department of Cusco, Per. Here,
I present a list of all the ectoparasites identified to date with a
reference to the host where they
were captured. I also give a brief
description of the characteristics of
each group found in the area.
Methods
This work was conducted in the

month of May, the last month of
the rainy season in the study area.
Both well sites were covered by a
dense vegetation. The bamboo
Guadua sp. dominated San Martin-3,
while at Cashiriari-2 we found
primary forest with some secondary
forest in the areas cleared for the
well (see Alonso et al. this volume).
The two well sites are 12 km apart
(10 min. helicopter flight) near the
Camisea River.
Bats were captured with mist
nets as described in Wilson et al.
(this volume), and Voss and
Emmons (1996). Nets were placed
352
along trails and within the

biodiversity plots established at the
well sites. Rodents were trapped by
Solari and Rodriguez (this volume),
and I sampled the parasites within
them. Material was removed by
hand with entomological forceps,
knives soaked in alcohol, fine
combs, and brushes; sometimes a
stereoscopic microscope was employed. All sampled materials were
placed in vials with 70% alcohol
and transported to the laboratory
for analysis. The specimens will be
mounted in permanent slides for
their taxonomic identification. In
the laboratory, the specimens were
processed and mounted in Hoyer
solution. The specimens were then
dried at 56 C, and identified to
genus and species using a compound microscope and the taxonomic keys of Fairchild et al.
(1966), Wenzel et al. (1966),
Wenzel (1976), Guerrero (1990,
1993, 1996) among others.
Results and
Discussion
Of the 428 individual bats

sampled by colleagues, I reviewed a
total of 207 specimens from 32
species most of them within the
family Phyllostomatidae (Appendices 1 and 2). I found a 68.5% rate
of infestation. I also reviewed 30
individual rodents, mostly of the
family Muridae, out of the 97
captured specimens and registered a
93.3% rate of infestation. Some
species were free of ectoparasites.
The bat Platyrrhinus brachycephalus
(subfamily Stenodermatinae) and
the mouse Oryzomys nitidus were the
most common species checked for
parasites.
I sampled a total of 155 specimens of ectoparasites, most of
them collected for the first time to

the Camisea Region (Appendix 2).
The Streblidae were the parasites
most frequently found, in particular
the genus Trichobius (Gervais 1844).
The Nycteribiidae, genus Basilia
(Ribeiro 1903) were found very
unfrequently (Table 1). The following families (order, author, year )
have been identified: Streblidae
(Diptera) Kolenati 1863,
Nycteribiidae (Diptera) Samonelle
1819, Spinturnicidae (Acarina)
Oudemans 1902, Macronyssidae
(Acarina) Oudemans 1936,
Labidocarpidae (Acarina) Gunther
1942, Trombiculidae (Acarina)
Ewing 1944, and Rhopalopsyllidae
(Siphonaptera) Baker 1905 (see
Appendix 3).
The Streblidae (STR) family is
composed by a diverse group of
blood feeding flies specialized on
bats mostly from the
Phyllostomidae family. All these
flies are ectoparasites with the
exception of females of the genus
Ascodipteron from the old world
(Wenzel 1970). There are 223
known species classified in five
subfamilies, three from the new
world with 147 species (Guerrero
1992). I sampled 86 individuals of
this well known family to Per
(Wenzel 1970; Koepcke 1987;
Guerrero 1992). The first species
for Per was described by

Townsend (1913). The most frequently found genus was Trichobius.
The Nycteribiidae (NYC)
family is also composed by blood
feeding apterous flies that feed on
bats within the Vespertilionidae,
Molossidae, and Phyllostomidae
family. There are genera for the new
world: Basilia (Ribeiro 1903), and
Hershkovitzia (Guimaraes and
DAndretta 1956). Only five specimens were sampled. Guimaraes and
DAndretta (1956) revised the
family for the species found in Per
(Guerrero 1996).
The Spinturnicidae (SPI) family
is composed by parasitic mites of
bats. Three genera have been
described for the new world. It is a
well represented group in Per, but
was not considered in the taxonomic revisions by Rudnick (1960)
and Webb and Loomis (1977). I
found 26 specimens.
The Trombiculidae (TRO)
family is composed by mites that
parasite a diverse group of vertebrates including reptiles, amphibians, birds, and mammals. The
larval stage is the parasitic phase
that in some cases can be found in
humans (Jenkins 1948; Jones 1950).
I found 14 specimens.
The Ixodidae (IXO) family is
composed by mites of the genera
Table 1. Frequency of collection of ectoparasites per family and their most predominant genus
Family
Streblidae (Diptera)
Nycteribiidae (Diptera)
Spinturnicidae (Acarina)
Trombiculidae (Acarina)
Ixodidae (Acarina)
Pulicidae (Siphonaptera)
Rhopalopsyllidae (Siphonaptera)
Stephanocircidae (Siphonaptera)
Frequency (%)
55.4
3.22
16.7
9.03
5.80
3.87
3.87
1.93
Genus
Trichobius
Basilia
Periglischrus
Eutrombicula
Amblyomma
Xenopsylla
Polygenis
Sphinctopsylla
Frequency (%)
74.4
100.0
100.0
57.1
55.6
100.0
66.7
100.0
353
Amblyomma and Ixodes. These mites

are found in the hard surfaces of
the body. These parasites have a
dorsal shield in all stages, as well as
a capitulum that can be seen dorsally. These parasites stay in their
host for long periods and they have
medical importance since they
transmit patogens that can cause
toxonemy, paralisis, and skin irritations (Cooley 1946; Farichild 1966;
Hoogstraal 1967; Jones et al. 1972;
Faust et al. 1974). I found nine
specimens.
The Pulicidae (PUL; genus
Xenopsylla Glinkiewiex 1907), the
Rhopalopsyllidae (RHO; genus
Rhopalopsyllus (Baker 1905), and the
Stephanocircidae (STE; genus
Sphinctopsylla Jordan 1931), within
the Siphonaptera group are widely
distributed and well known for the
medical importance as they are
parasites of mice (Ryckman 1971).
This group has not been studied in
Per systematically except by
Maihuay et al. (1994). Fifteen
species were sampled.
Species accounts
Trichobius joblingi (Wenzel 1966)

was frequently found in the bat
Carollia perspicillata, but it was also
sampled from C. brevicauda, C.
castanea, Artibeus lituratus, and
Diphylla ecaudata. Guerrero (1996)
collected this parasite in C.
brevicauda, C. castanea, and C.
perspicillata and Phyllostomus elongatus
from Pakitza, and suggests a possible new species from C. castanea.
Wenzel (1970) found it in C.
perspicillata in Junin, Loreto and
Puno, in Per, while Orozco (1990)
found it in P. elongatus in
Tambopata, Madre de Dios. This is
the first collection for T. joblingi in
Camisea and D. ecaudata for Per.
354
T. dugesii (Townsend 1891) was

registered by Gerrero (1996) in Per
from Glossophaga soricina. Chavez
and Mendoza (1994) found it in Ica
(southern Per), and Wenzel et al.
(1976) sampled it in Junin in the
same host. There is great association between this parasite and
Glossophaga, although it has been
collected also from other bats.
Here, I report for the first time this
parasite in C. perspicillata and C.
brevicauda in Camisea.
T. persimilis (Wenzel 1976) was
found for the first time in Per in C.
perspicillata. Wenzel found this
species in Venezuela mostly in C.
brevicauda. This parasite is very
similar to T. joblingi.
T. parasiticus (Gervais 1844)
was recorded previously from
Cusco, Ica, Piura, Iquitos and
Loreto, Per, by Wenzel (1976)
from the blood feeding bat Desmodus
rotundus, and by Elliot et al. (1985)
from Huaraz (Per) from the same
bat species. Orozco (1990) reported
this parasite from Sturnira lilium
from Tambopata Madre de Dios,
Per. I found it on C. brevicauda.
Trichobius sp. 1, Trichobius sp. 2,
and Trichobius sp. 3 were sampled
from C. perspicillata, C. castanea, and
C. brevicauda, all very similar to T.
persimilis, T. joblingi y T. dugesii but
they present large wings, transversal
suture mesonotal little angulated,
reduced mesonotal quetotaxia, with
a more or less large row of setae in
the ante-escutelar region, some
specimens lack the metastaernal
lobule. I could not evaluate some
abdominal characteristics due to the
state of preservation, but they are
clearly of animals within Trichobius.
Speiseria ambigua (Kessel 1925)
was registered in Loreto and
Lambayeque by Wenzel (1970) from
an unidentified guest, although the
same author and Guerrero (1994,

1996) had found it in C. perspicillata.
Speiseria peytonae (Wenzel 1976)
was registered for the first time on
C. brevicauda in Venezuela. Guerrero
(1994) also found it in Venezuela,
and Pakitza, Per, (1996).
Pseudostrebla riberoi (Costa Lima
1921) was registered in Loreto, Per
(Wenzel 1970) from Tonatia silvicola.
Wenzel (1976) found it in
Amazonas, Venezuela found it on
the same host. Guerrero (1994,
1996) registered in Venezuela and
Pakitza, all on the same host. I
found two male specimens on the
same species of bat.
Paratrichobius longicrus (Miranda
Ribeiro 1907) was registered in
Loreto, Per by Wenzel (1970), and
by Wenzel (1976) in Venezuela
from the bat Artibeus lituratus (190
individuals). Guerrero (1996)
revised material sampled from the
same species of bat collected in
Per, Venezuela, Bolivia, and
Brazil. I found this species for the
first time in Camisea mostly from
A. lituratus, but it was also sampled
from Carollia perspicillata and
Chirodema villosum.
Paratrichobius dunni (Curran
1935) was sampled from several
locations in Venezuela (Wenzel
1976) most frequently from
Uroderma bilobatum followed by U.
magnistrum. Guerrero (1996)
sampled one female and one male
from U. bilobatum, in Pakitza, Per.
Guerrero (1994) revised material
from Venezuela, Brazil, and Bolivia,
all sampled from the same species
of bat.
Paratrichobius cf. salvini (Wenzel
1966) described from the bat
Chiroderma salvini. Wenzel (1976)
reported it from Venezuela parasitizing the same species of bat,
although he also found it in other
hosts and determined as

Paratrichobius species A and B. After
revising specimens from Venezuela
and Brazil, Guerrero (1994) found
differences in male genitalia among
individuals coming from different
bat species (e.g. Uroderma
bilobatum), but he considered all
morphotypes within the same
species. In this study I followed his
criteria since I found differences
among individuals coming from U.
bilobatum, Chiroderma villosum, and
C. trinitatum bats.
Paratrichobius sp. 1 was sampled
from Artibeus lituratus (two females
and two males), and Vampyressa
bidens (one female). It is
charcarterized by seven large spines
in the pre-femure, being the most
apical curved. It also has a parallel
row of short setae and fine posterior legs not long that makes it
resemble P. sanchezi. More material
needs to be analyzed before they
can be classified.
Anatrichobius sp. 1 was sampled
from Myotis riparius, and it has
similar morphology to A. scorzai in
its form, in the quetotoaxia of the
mesonoto, the form and size of the
wings, as well as the genitalia.
Megistopoda aranea (Coquillet
1899) was previously found in
Lambayeque and Huanuco, Per
(Wenzel 1970). Wenzel (1976)
found 545 individuals in Venezuela
most of them from Artibeus
jamaicensis. Guerrero (1996) found it
in Pakitza, Per, in the same bat
species and on female from
Uroderma bilobatum. Guerrero (1994)
determined material from Venezuela, mostly from Artibeus jamaicensis,
while material fromBolivia and
Brazil material came from A.
lituratus, and a female found in
Diphylla ecaudata from Mxico. I
found two specimens in A. lituratus.
355
Megistopoda proxima (Seguy

1926) was previously found in
Madre de Dios, Per in Sturnira
lilium (Jobling 1952). Guerrero
(1996) found 10 individuals in the
same species of bat in Pakitza, and
Orozco (1990) reports it from
Tambopata Madre de Dios.
Guerrero (1994) registered in
several localities and hosts in
Venezuela, Panama, Martinica,
Dominica, and Bolivia mostly on
Sturnira, and he argues that all these
specimens may belong to an species
complex rather than to a single
species. I also sampled from S.
lilium.
Asidoptera phyllostomatis (Perty
1833) was previously sampled in
Venezuela mostly from Artibeus
jamaicensis, although it has also
being sampled from Anoura caudifer,
Artibeus lituratus, A. fuliginosus,
Glossophaga soricina, Phyllostomus
hastatus, Pternotus parnelli, and
Uroderma bilobatum (Wenzel 1966).
The same author concludes that
Asidoptera busckii is synonimus to A.
phyllostomatis. Guerrero (1996) found
10 individuals in A. jamaicensis, and
two from A. obscurus. Here, I found
A. phyllostomatis in A. planirostris.
Asidoptera sp. 1 was sampled
from Sturnira lilium and S. tildae
(two females and one male). It is
similar to A. falcata and A. delatorrei
registered in Venezuela by Wenzel
(1976). Guerrero (1996) registered
A. falcata to Per sampled from S.
lilium, and comments the works of
Koepcke (1988) who sampled A.
delatorrei from S. lilium in Huanaco,
Per. This can be a misidentification since A. delatorrei has
only being sampled from Panama,
Guatemala, Mexico and Venezuela,
but not in the Amazon Basin.
Similarly, Orozco (1990) possibly
misidentified an specimen that he
356
named A. delatorrei sampled from S.

lilium in Tambopata, Madre de Dios,
Per.
Neotrichobius delicatus
(Machado-Allison 1966) was
sampled from Vampyresa pusilla from
Venezuela. Wenzel (1966) also
found this parasite in Artibeus spp.,
A. cinereus, and Rhinophylla pumilio
but with different measurements;
thus the author only considered as
N. delicatus those sampled from V.
pusilla. Guerrero (1996) revised
samples from Venezuela sampled
from V. pusilla, as well samples
from Pasco, Per, and from the bat
R. pumilio, and concludes that this
parasite seems to have large distributions and can have several species as hosts, similar to the genus
Paratrichobius. It is thus recommended to do a taxonomic revision
of this complex. Here, I sampled
two females from V. pusilla.
Neotrichobius cf. stenopterus
(Wenzel and Aitken 1966) was
described for the first time in
Panama sampled from Uroderma
bilobatum. Guerrero (1996) considers this species as confused, since it
has been reported from Panama,
Trinidad, Suriname, Brazil and
Colombia (Wenzel and Aitken
1966; Wenzel 1970). Wenzel
(1976), however, determined
material from Suriname as N.
bisetosus, and Trinidad as N.
delicatus, and considered N. delicatus
and N. stenopterus as synonymous. I
sampled one specimen from U.
bilobatum. It has many similarities to
N. stenopterus, but we believe that
this group should be revised
taxonomically.
Metelasmus pseupdopteros
(Coquillet 1907) was described
from Artibeus lituratus from Paraguay. Wenzel (1966) reported it
from several species of bats includ-
ing A. lituratus, Chiroderma villosum,

and Phyllostomus hastatus from
Venezuela. A. lituratus was the most
frequent species. I found it in A.
lituratus and Platyrrhinus
brachycephalus, two females and one
male respectively, which are first
records for Camisea.
Strebla guajiro (Garcia and Casal
1965) has been sampled in Venezuela in the fish eating bat Noctilio
labialis. Wenzel (1966), however,
registered it from 76 localities in
Venezuela mostly from Carollia
perspicillata (89.2%) and C.
brevicauda (8.7%), and concluded
that Carollia is the genus that this
parasite is found rather than the fish
eating bat. Moreover, Guerrero
(1996) reported it for Pakitza in C.
brevicauda (more frequent) and C.
perspicillata. In this study, I found
the first records for Camisea being
three females in C. brevicauda, two
males in C. perspicillata, and one
female in Carollia sp.
Strebla alvarezi (Wenzel 1966)
was registered in Panama from
Micronycteris megalotis. Wenzel et al.
(1966) reported it also from
Micronycteris and Saccopteryx bilineata
from Panama. Guerrero (1996)
reported the first record, one female, for Per in Lonchophylla
thomasi from Pakitza. I found two
females in L. thomasi, and one male
from Platyrrhinus infuscus.
Strebla cf. mirabilis (Waterhouse
1879) was sampled in Loreto and
Junin, Per, from Phyllostomus
hastatus, P. elongatus, and Trachops
cirrhosus, and in Venezuela from T.
cirrhosus (91.6%) and P. hastatus
(Wenzel et al. 1966). Guerrero
(1996) found seven specimens in T.
cirrhosus, and one male from P.
elongatus in Pakitza. I found a male
in the vampire bat Diphylla ecaudata
that represents the first record for

Camisea.
Strebla sp. 1 is represented by
one individual from Tonatia silvicola.
It has morphological characteristics
such as separate big and rectangular
plates from Frontoclipeo, form and
quetotaxia of the Mesonoto, occipital plates with strong spiny setae,
and inclinated and angular
postgonitos that could be considered as S. kohlsi, reported from
Venezuela from T. silvicola (Wenzel
1976). Guerrero (1996) also found
it in T. silvicola and Phylloderma
stenops.
Pseudostrebla riberoi (Lima 1921)
was registered originally from Brazil
from Tonatia amblyotis (sin.
silvicola). It was reported also to
Per from Loreto (Jobling 1939),
and from Venezuela (three females)
from T. silvicola (Wenzel 1976).
Guerrero (1996) found one male in
T. silvicola from Pakitza, and I found
two males from the same bat species.
Paradyschiria parvula (Falcioz
1931) was registered from Noctilio
labialis in Brazil. Wenzel (1966)
found it more frequently in N.
labialis, but also in Molossus aztecus
and M. ater from Venezuela.
Guerrero (1996) reported it from a
sample of 462 individuals all
sampled from N. albiventris. I found
two specimens in Promops centralis, a
possibly occasional host.
Basilia myotis (Curran 1935) was
described from a specimen sampled
from Myotis nigricans in Panama. This
species has a wide distribution and
has been found in Panama, Guatemala, Colombia, Per, Venezuela,
and British Guayana. In Per, three
individuals were sampled in Loreto
(Guimaraes and Dandretta 1956).
This species seems to be synonymous to B. bellardii (Rondani 1878),
357
and with B. ferrisi (Theodor 1967),

and it is closely related to B.
speiserie, to whom it differs by
having a large number of setae in
targite number 1, as well as in the
quetotaxia, the form of the lateral
sclerites of the esternite 6. B. myotis
has been considered a subspecies of
B. speiserie. I sampled three males
found in Myotis riparius.
Basilia manu (Guerrero 1992)
was described from a specimen
sampled from Myotis riparius from
Pakitza, Manu National Park, Per.
This species presents similarities
with Basilia junquiensis (Guimaraes
1946) and B. anceps (Guimaraes and
DAndretta 1956), but B. manu has
short spiny seate in the lateral
abdominal conexivio. Guerrero
(1992) also pointed out that males
present 11 spines in the margin of
the sternite V, while B. anceps has
only eight. I found one female
specimen of this parasite in M.
riparius.
Basilia cf. peruvia (Guimaraes
and DAndretta 1956) has been
found in Marcapata, Quince Mil
(Cusco), and Huanuco (Pozuzo),
Per, from the bats Myotis nigricana
and Tadarida Braziliensis. This
parasite is similar to B. travassosi
regarding the lack of setae in the
posterior border of the sterna 3, as
well as the tergite 2 that reaches the
lateral borders of the abdomen.
They can be separated since the
tergite 1 is bigger than B. travassosi,
and because the discal setae are
localized in the lateral borders. I
found one individual male.
Periglischrus imeringi (Oudemans
1902) was found previously in
Brazil living on Vampyrops lineatus,
but it was later registered as P.
meridensis (Hirst 1927) based on a
specimen sampled in Artibeus
jamaicensis from Venezuela. Radford
358
(1951) registered it as Spinturnix

artibiensis from Trinidad living in the
bat Artibeus lituratus. It has been
sampled also in Cuba and Puerto
Rico living on A. jamaicensis, in
Paraguay on Vampyrops sp. 1, in
Honduras parasitizing A. lituratus
and Uroderma bilobatum, and in
Colombia living on A. lituratus
(Rudnick 1960). Machado-Allison
(1965) challenged Rudnick proposing different subspecies. The same
author found this parasite in Venezuela living on A. lituratus, Artibeus
sp. 1, A. cinereus, Urodema
bilobatum, and Vampyrops vittatus.
Chavez and Mendoza (1994) found
in Ica, Per, P. iheringi in Artibeus
fraterculus.
Periglischrus caligus (Kolenati
1957) has been sampled in Brazil
and Surinam living on Glossophaga
soricina, and likes to parasite species
of the genus Glossophaga (Furman
1966; Herrin and Tipton 1975;
Machado-Alisson 1965; and
Rudnick 1960). Since the original
description can not permit comparisons with P. setosus,
Machado-Alisson and Antequera
(1975) support Furman (1966)
suggestion that P. caligis and P.
setosus are synonymous, and that P.
vargasi (Hoffmann) are closely
related. Chavez and Mendoza
(1994) found in Ica, Per, in G.
soricina. I found only three individuals on the same species of bat.
Periglischrus cf. acutisternus
(Machado-Allison 1965) has been
found in Phyllostomus elogatus, P.
hastatus, and P. concolor in Venezuela. Furman (1966) described this
parasite species as P. tiptoni living
on Phylostomus hastatus panamensis in
Panama, since he did not know
Machado-Allisons description.
Furman in a later paper found it
Trinidad living on Phyllostomus
discolor; in Trinidad and Panama on

Trachops cirrhosus and Myotis
chiloensis, and on P. hastatus and P.
elogatus in Colombia, and concludes
that P. tiptoni is synonymous of P.
acutisternus. I found one specimen.
Periglischrus ojasti
(Machado-Allison 1964) was the
second most frequent parasite
sampled from Sturnira lilium, S.
tildae and S. magna. This parasite is
closely related to P. iheringi, specially the females since the form of
the sternal plate is piriforme and
presents posteriorventral setae in
the femur, patela and tibia IV. They
are different in that the first pair of
propodesomal seate are bigger in P.
ojasti than in P. iheringi.
Spelaerhynchus sp. 1 was sampled
(two individuals) from the genital
region of Carollia perspicillata. This
is the first record for Per.
Eutrobicula sp. 1 was sampled 8
times in larval stages from Carollia
brevicauda, Oryzomys capito, O. nitidus,
and Oryzomys sp. 1 (Ewing 1938). It
was reported from Venezuela by
Brennan and Reed (1972). This
genus has medical importance since
it transmits pathogen such as
rickettsias. It has close to 22 parasitic species of mice and marsupials
being the most common
Eutrombicula goeldii. This is the first
record of this parasite to Per.
Parasecia sp. 1 was sampled
once in a larval stage from Mimon
crenulatum. It had a particular position when it was mounted but
characteristics such as the totaxia
of the palps, morphology and
quetotoxia of the shield, segmentation, and quetotoxia of the legs,
permitted classified it in the genus.
It was registered before in Per as
P. manueli by Brennan and Reed
(1972) and the same authors described five species and eight to be
described. This is the first record

for Camisea and probably the
second for Per.
Odontacarus was sampled from
Platyrrhinus helleri, Proechimys spinosus
and Oecomys bicolor (Ewing 1929).
Brennan and Reed (1972) recognized 12 species in Venezuela
mostly as parasites of mice, marsupials and bats, and reported in Per
Odontacarus australis as a parasite of
lizards.
Amblyomma has been sampled
from the mice Proechimys spinosus and
Oryzomys macconnelli. Most of the
specimens were larval stages and
one nymph. This genus was described by Koch (1844), it has wide
distribution and it is represented by
19 species in Panama (Fairchild et
al. 1965 ) and by 25 species in
Venezuela (Jones et al. 1972). This
is the first report for Per.
Ixodes was sampled from the
mice Oryzomys nitidus and represented by four nymphs. This genus
and the previous are widely distributed and parasite several species in
several orders. It is represented by
11 species in Panama mostly form
mice (Jones et al 1972). The genus
was described by Latreille (1795).
This is the first report of the genus
to Camisea.
Polygenis roberti (Fox 1947) was
registered in Panama from Philander
opossum, Marmosa robiinsoni,
Metachirus nudicaudatus, Didelphis
marsupialis among others. In Per it
has been sampled from Dasypus
novemcinctus, Oligoryzomys
longicaudatus, among others. In
Venezuela has been found in
Oryzomys fulvescens, Oecomys trinitus,
Heteromys anomalus, Neacomys
spinosus, Sigmodon hsipidos, and
Rattus rattus. Tiotin and MachadoAllison (1972) sampled 152 females
and 114 males mostly from Didel-
359
phis marsupialis. Maihuay et al.

(1993) registered several specimens
form the highlands of Madre de
Dios, Per, between 2250-3350 m,
including P. thormanii and Polygenis
sp. 1 sampled from the mice
Thomasomys aureus, Akodon aerosus,
and Oryzomys keaysi, and have been
associated to transmitting the
plague (Neyra 1987). I sampled two
individuals from the mice Oryzomys
capito at an altitude of 613 m,
representing the first record to the
area.
Polygenis cf. occidentalis (Jordan
and Rothschild 1923) was registered
from Venezuela living on Sturnira
lilium, but also from Trinidad from
marsupials, mice such as Oryzomys,
and in Venezuela from del Didelphis
marsupialis, and Sciurus griseogena. On
their study on fleas, Tipton and
Machado -Allison (1972) found 18
specimens (nine females and nine
males) from which more than 80%
were sampled from Rhipidomys
venustus, Didelphis marsupialis,
Sturnira lilium and Sciurus granatensis,
and the rest of the material was
taken from Proechimys semispinosus
and Rhipidomys couesi. They found
differences in the male diagrams
and the keys by Johnson (1957)
since they found a lower number of
setae in the metatibias (commonly
18-19). I took two samples, one
male from the mouse Oryzomys
nitidus, and one female from
Oryzomys sp. 1. The female sample
is not in good condition, so it will
be risky to classify the male as P.
occidentalis due to the large variability found in the species.
Rhopalopsyllus sp. 1 was sampled
from Proechimys spinosus (two males).
This genus was described by Baker
(1905). This flea species has been
reported from Panama, Venezuela,
Mexico, Colombia, Guatemala,
360
Bolivia, Brazil, and Per. Tipton

and Machado-Allison (1972) reported up to four species parasitizing mostly rodents and marsupials.
Maihuay et al. (1993) reported
material from the highlands of
Madre de Dios, Per, sampled from
Akppdpn torques and Oximicterus
paramensis.
Xenopsylla cheopsis (Rothschild
1903) is an species widely distributed that transmits the plague. I
sampled six individuals, females and
males, from Oryzomys capito,
Oryzomys sp.1, Oecomys bicolor, and
Proechimys spinosus. Tipton and
Machado-Allison (1972) found six
individuals mostly from Rattus
rattus, Oryzomys albigularia, and
Sigmodon hispidus in Venezuela. They
commented that after three years of
sampling, they only collected three
specimens of X. cheopsis, and argue
that the use of insecticides may be
affecting these fleas. Maihuay et al.
(1993) did not find this flea his
study in Per.
Sphinctopsylla sp. 1 is represented by three females found in
Oryzomys macconnelli. Tipton and
Machado-Allison (1972) reported
the genus with the species
Sphinctopsylla tolmera from Venezuela
with 112 individuals both females
and males where more than 87%
was sampled from members of the
genus Oryzomys, and reporting that
most of these fleas have an Andean
distribution and are found below
3000 m.
References
Brennan, J., and Reed. J. 1972. A

list of Venezuela chiggers particularly of small mammalian hosts
(Acarina: Trombiculidae). Brigham
Young Univ. Sci. Bull. Biol. Series
XVII (6).
Caceres, I., A. Elliot, R. Orozco,

and L. Campones. 1992.
Identificacin de especies de
Trhicobius (Gervais 1844) (Diptera:
Streblidae), en murcielagos de la
Reserva Nacional de Tambopata,
Madre de Dios Per. Rev. Per. Med.
Trop. U.N.M.S.M 6: 99-102.
Cooley, R. A. 1946. The genera
Boophilus, Rhipicephalus and Haemaphysalis (Ixodoidea) of the New
World. Natl Inst. HLTH Bull.
(187).
Chavez, J., and L. Mendoza. 1994.
Identificacin de ectoparatistos y
endoparasitos en algunas especies
de Quiropteros en el Valle de Ica.
Tesis Biologo UNSLG.
Doreste, E. 1984. Acarologa. Boletn
del IICA. San Jos de Costa Rica.
Elliot, A., I. Caceres, and
S. Macedo. 1985. Ectoparasitos del
Desmodus Rotunds I. Trichobius
parasiticus Gervais 1844 (Diptera:
Streblidae) Bol. Inst. Med. Trop.
U.N.M.S.M. 4: 18-21
FAO/WHO Expert Committee on
Zoonoses. 1967. Third Report
World Health Organization Technical Report Series No. 378.
Geneva.
Fairchild, G., G. Kohls, and
V. J. Tipton. 1965. The ticks of
Panama (Acarina: Ixodoidea). In
The Ectoparasites of Panama. R.
R. Donnelley and Sons Co., Chicago.
Furman, D. P. 1966. The
Spinturnicidae mites of Panama.
Pages 125-166 in Ectoparasites of
Panama. (R. L. Wenzel and
V. J. Tipton, eds.) Field Museum of
Natural History, Chicago.
Gettinger, D., and R. Gribel. 1989.
Spinturnicid mites (Gamasida:
Spinturnicidae) associated with
bats in central Brazil. J. Med.
Entomol 26(5): 491-493.
Goff, M., J. Whitaker, and

L. Barkley. 1984. A new species of
Loosimia (Acarina: Trombiculidae)
from a Peruvian bat. J. Med.
Entomol. 21(1): 80-81.
Gorirossi, F. E. 1950. The mouth
parts of the adult female tropical
rat mite (Liponyssus Bacoti) with
observations on the feeding
mechanism. J. Parasitol. 36:
301-318.
Guerrero, R. 1992. Streblidae
(Diptera: Pupipara) parasitos de
los murcielagos de Pakitza, Parque
Nacional de Manu (Per). Pages
627-641 in Manu: Biodiversity of
and A. Sandoval, eds.) Smithsonian Institution, Washington, DC.
Guerrero, R. 1993. Catalogo de los
Streblidae (Diptera: Pupipara)
parasitos de Murcielagos (Mammalia: Chiroptera) del nuevo Mundo
I. Clave para los generos de
Nycterophillinae. Acta Biol. Venez.
14(4): 61-75.
Guerrero, R. 1994. Catalogo de los
Streblidae (Diptera: Pupipara)
parasitos de murcielagos (Mammalia: Chiroptera) del Nuevo Mundo
IV. Trichobiinae con alas
desarrolladas. Bol. Entomol. Venez.
9(2): 161-192.
Guerrero, R. 1996. The Basilia
junquiensis species group (Diptera:
Nycteribiidae) with description of
a new species from Pakitza, Per.
Pages 665-674 in Manu:
Washington, DC.
Guerrero, R. 1996. Estudio de los
ectoparasitos de los murcielagos de
Pakitza, Parque Nacional Manu
(Per). Pages 643-657 in Manu:
361

Washington, DC.
Guimares, L. R., and V. D.
DAndretta. 1956. Sinopse dos
Nyc-Teribiidae (Diptera) do Novo
Mundo. Arquivos de Zoologia do
Estado de Sao Paulo 10: 1-184
Haas, V. H. 1959. When bubonic
plague came to Chinatown. Amer.
J. Trop. Med. & Hyg. (8): 141-147.
Herrin, C. S., and V. J. Tipton.
1975. Spinturnicidae mites of
Venezuela (Acarina:
Spinturnicidae). Brigham Young
Univ. Sci. Bull. Biol. Series 27 (3):
1-58.
Hopkins, G., and M. Rothschild.
1956. An Illustrated Catalogue of
the Rothschild Collection of Fleas
(Siphonaptera) in the British
Museum (Natural History) Vol. 2:
Coptopsyllidae, Vermipsyllidae,
Stephanocircidae, Ischnopsyllidae
[Macropsyllidae],
Hypsophthalmidae And
Xiphiopsyllidae. British Museum
(Natural History), London.
Hoogstraal, H. 1961. Tick and tickborne diseases: some international
problems and cooperation in their
study. Intl. Rev. Trop. Med. (1):
247-267.
Hoogstraal, H. 1967. Ticks in
relation to human diseases caused
by Rickettsia species. Ann. Rev.
Entomol. 12: 377-420
Horsfall, W. R. 1962. Medical
Entomology. The Ronald Press,
New York.
Herms, W., and M. James. 1961.
Medical Entomology. 5th. Edition.
The Machimillan Co., New York.
Jenkins, D. W. 1948. Trombiculid
mites affecting man: bionomics
with references to epidemiology in
the United States. Amer. J. Hyg. 48:
22-35.
Jones, B. M. 1950. The penetration
of the host tissue by the harvest
362
mite (Trombicula autummalis: Shaw).

Parasitology 40: 247-260.
Jones, E., C. Clifford, J. Keirans,
and G. Kohls. 1972. Ticks of
Venezuela (Acarina: Ixodoidea)
with a key to the species of
Amblyommma in the Western Hemisphere. Brigham Young Univ. Sci.
Bull. Biol. Series XVII (3): 1-40.
Jellison, W. L. 1959. Fleas and
disese. Ann. Rev. Entomol. 4:
389-414
Kohls, G. M. 1955. Colorado tick
fever discovered in California.
California Vector Views 2(4): 1.
Kohls, G. M., and R. E. Ryckman.
1962. New distributional records
of ticks associated with cliff
swallows Petrochelidon spp. in the
United States. J. Parasitol 48 (3):
507-508.
Krantz, G. W. 1978. A Manual Of.
Acarology. 2nd. ed. Oregon State
University Book Stores, Inc.,
Corvallis, Oregon.
Machado-Allison, C. E. 1965.
Notas sobre mesostigmata
neotropicales III. Cameroni e ta
Thomasi: nuevo genero y nueva
especie parasita de Quiroptera
(Acarina, Spinturnicidae). Acta
Biol. Venez. 4(10): 243-258.
Machado-Allison, C. E. 1965a. Las
especies venezolanas del genero
Periglischrus Kolenati 1857
(Acarinaa: Mesostigmata,
Spinturnicidae) Acta Biol. Venez.
4(11): 259-348.
Machado-Allison, C. E. and
R. Antequera. 1971. Notes on
neotropical mesostigmata: four
new Venezuelan species of the
genus Periglischrus (Acarina:
Spinturnicidae) Smithsonian Contrib.
Zool. 93: 1-16
Macchiavello, A. 1959. Estudios
sobre pre-Selvtica en Amrica del
sur I: concepto y clasificacin de la
peste. Bol. Ofic. Sani. Panama 39:

339-349.
Maihuay, C., V. Pacheco, and
S. Solari. 1993. Siphonaptera
(insecta en roedores Silvestres del
Cuzco). Rev. Per. Ento. (36): 27-28.
Morales-Malacara, J., and R. Lopez.
1990. Epizoic fauna of Plecotus
mexicanus (Chiroptera:
Vespertilionidae) in Tlaxcala,
Mexico. J. Med. Entomo. Vol. 27(4):
440-445.
Marshall, A. G. 1976. Host specificity amongst arthropods ectoparasitic upon mammals and birds in
the New Hebrides. Ecol. Entomo. 1:
189-199.
Marshall, A. G. 1982. Ecology of
insects ectoparasites on bats. Pages
369-401 in Ecology of Bats. (T.
Kunz, ed.)
Need, T., W. Dale, J. Keirans, and
G. Dasch. 1991. Annotated list of
ticks (Acari: Ixodidae, Argasidae)
reported in Per. J. Med. Entomol.
28(5): 590-597.
Orozco, R. 1990. Dipteros
Hematofagos de la Reserva
Nacional de Tambopata. Tesis
Biologo UNMSM.
Parker, R., R. Menon, A. Merideth,
M. Snyder, and T. Woodward.
1954. Persistence of Rickettsi
rickettsii in a patient recovered
from Rocky Mountain spotted
fever. J. Immunol. 73: 383-386.
Philip, C. B. 1961. Arthropod
vectors in relation to the reservoir
mechanism of microbial agent of
animal diseases. Acta Trop. (18):
257-262.
Philip, C. B. 1968. A review of
growing evidence that domestic
animals may be involved in cycles
of Rickettsial zoonoses. Bakt.
Parasitenk, Infektionskr. Hyg. 206:
343-353.
Ross, A. 1961. Biological studies on
bat ectoparasites in the genus
Trichobius (Diptera: Streblidae) in

North America, north of Mexico.
Wasman J. Biol (19): 229-246.
Rudnick, A. 1960. A. revision of
the mites of the familiy
Spinturnicidae (Acarina). Univ.
Calif. Pub. Entomol. 17(2): 157-283.
Tipton, V. J., and C. E. MachadoAllison. 1972. Fleas of Venezuela.
Brigham Young Univ. Sci. Bull. Biol.
Series XVII (6): Sept. 1972.
Theodor, O. 1967. An Illustrated
Catalogue of the Rothschild
Collection of Nycteribiidae
(Diptera). British Museum (Natural History), London.
Webb, J. P., and R. B. Loomis. 1977.
Ectoparasites. Pages 57-119 in
Biology of Bats of the New World
Family Phyllostomidae. (R. J.
Baker, J. K. Jones, and D. C.
Carter, eds.) Part. II Spec. Publ.
Mus. Texas Tech. Univ. (13): 1-364
Wenzel, R., V. J. Tipton, and
A. Kiewliez. 1966. The Streblidae
batflies of Panama (Diptera,
Calipterae: Streblidae. In Ectoparasites of Panama. (R. Wenzel
and V. J. Tipton, eds.) Field Museum of Natural History, Chicago.
Wenzel, R. L. 1970. A Catalogue of
the Diptera of the Americas South
of the United States. Mus. Zool.
Univ. Sao Paulo; Form. Dep. Zool.
Secretaria Da Agricultura.
Wenzel, R. L. 1976. The Streblid
batflies of Venezuela (Diptera:
Streblidae) Brigham Young Univ. Sci.
Bull. 20: 1-177.
363
Appendix 1. Total bats and rodents inspected for parasites (SM(1) = San
Martin plot 1; SM(2) = San Martin plot 2; SM(a) = Near San Martin; CA(3) =
Cashiriari-2 plot 3; CA(4) = Cashiriari-2 plot 4; CA(a) = near Cashiriari-2; * =
free of ectoparasites).
SM(1) SM(2) SM(a) CA(3) CA(4) CA(a) Total
Bats
Phyllostomidae
Artibeus planirostris*
Artibeus obscurus*
2
Artibeus lituratus
Sturnira lilium
Sturnira tildae
Sturnira magna
Chiroderma villosum*
Artibeus planirostris* 1
Uroderma bilobatum
Dermanura anderseni
Dermanura glauca
Vampyressa macconne
Vampyressa bidens
Rhinophylla pumilio
Carollia perspicilla
Carollia castanea
1
Carollia brevicauda 2
Anoura caudifer
1
Choeroniscus minor
Mimon crenulatum
1
Tonatia silvicola
Diphylla ecaudata
1
Emballonuridae
Saccopteryx leptura*
Vespertilionidae
Myotis riparius
1
Molossidae
Molossus ater*
Promops centralis
Rodents Muridae
Oryzomys nitidus
7
Oryzomys capito
Oryzomys macconne
Oryzomys sp. 1
1
Oecomys bicolor*
Neacomys spinosus
2
Echimyidae
Proechimys spinosus 2
364
4
1
2
1
1
1
6
6
1
1
7
1
4
2
1
4
7
3
2
6
4
1
3
3
4
1
10
6
7
2
2
1
1
1
3
5
9
4
3
7
5
6
4
1
1
4
3
1
4
4
2
1
2
1
2
2
1
1
1
1
1
4
3
6
13
7
18
7
1
13
1
21
8
2
12
8
2
3
4
8
1
18
12
12
2
1
1
2
1
1
1
2
1
13
1
1
1
5
6
3
Appendix 2. Total number of ectoparasites of the most common families

sampled in bats and rodents (STR = Streblidae [Diptera: flies]; NYC =
Nycteribiidae [Diptera: flies]; SPI = Spinturnicidae [Acarina: mites]; TRO =
Trombiculidae [Acarina: mites]; IXO = Ixodidae [Acarina: mites]; PUL =
Pulicidae [Siphonaptera: fleas]; RHO = Rhopalopsyllidae [Siphonaptera:
fleas]; STE = Stephanocircidae [Siphonaptera: fleas]).
Ectoparasites
STR NYC SPI TRO IXO PUL RHO STE
Bats
Phyllostomidae
5
3
Artibeus obscurus
4
2
Artibeus lituratus
8
3
Sturnira lilium
5
1
Sturnira tildae
1
1
Sturnira magna
3
2
Chiroderma villosum
1
3
2
Platyrrhinus brachicephalus
2
2
1
2
1
Uroderma bilobatum
3
1
Dermanura Anderseni
1
Dermanura glauca
2
Vampyressa bidens
2
Vampyressa macconne
6
Rhinophylla puhilio
Rhinophylla castanea
15
3
5
1
Carollia castanea
6
3
3
Carollia brevicauda
2
Glossophaga soriccina
Anoura caudifer
1
Choeroniscus minor
1
Mimon crenulatum
2
Tonatia silvicola
2
Diphylla ecaudata
Emballonuridae
Saccopteryx leptura
Vespertilinoidae
1
5
Myotis riparios
Molossidae
Molossus ater
2
Promops centralis
Rodents Muridae
Oryzomys nitidus
1
2
4
Oryzomys capito
2
2
2
Oryzomys macconne
3
2
Oryzomys sp. 1
1
1
1
Oecomys bicolor
2
1
Echimyidae
Proechimys spinosus
1
2
2
3
Host
365
Appendix 3. Parasite-host species list for bats and rodents sampled in

Camisea. N = nymph; L = larva; A = adult; CF = species to confirm.
ORDEN QUIRPTERA
Phyllostomidae
Artibeus obscuros
Periglischrus iheringi (A) (N)
Trichobius joblingi (A)
Trichobius persimilis (A)
Trichobius sp. 1 (A)
Speiseria ambigua (A)
Paratrichobius longicrus (A)
Strebla guajiro (A)
Spelaeorhynchus sp. 1 (A)
Rinophylla pumilio
Paratrichobius cf. salvini (A)
Periglischrus iheringi (A)
Carollia castanea
Carollia brevicauda
Trichobius parasiticus (A)
Speiseria peytonae (A)
Strebla guajiro (A)
Eutrombicula sp. 1 (L)
Periglischrus iheringi (N) (A)
Neotrichobius cf. stenopterus (A)
Carollia sp. 1
Strebla guajiro (A)
Artibeus lituratus
Paratrichobius sp. 1 (A)
Megistopoda aranea (A)
Metelasmus pseudopterus (A)
Asidoptera phyllostomatis (A)
366
Uroderma bilobatum
Paratrichobius dunni (A)
Paratrichobius cf. salvini(A)
Neotrichobius cf. stenopterus (A)
Diphylla ecaudata
Strebla mirabilis (A)
Anoura caudifer
Trichobius dugesii (A)
Periglischrus caligus (A)
Strebla alvaresi (A)
Mimom crenulatum
Parasecia sp. 1 (L)
Tonatia silvicola
Pseudostrebla riberoi (A)
Strebla sp. 1 (A)
Chiroderma villosum
Odontacarus sp. 1 (L)
Appendix 3. Parasite-host species list for bats and rodents (Cont.).

Strebla alvaresi (A)
Sturnira lilium
Megistopoda proxima (A)
Dermanura anderseni
Molossidae
Uroderma bilobatum
Paratrichobius dunni (A)
Neotrichobius cf. ctenopterus (A)
Vampyressa bidens
Paratrichobius sp. 1 (A)
Vampyressa pusilla
Neotrichobius delicatus (A)
Vampyressa pusilla
Neotrichobius delicatus (A)
Metelasmus pseudopterus (A)
Asidoptera sp. 1 (A)
Periglischrus acutisternus (A)
Periglischrus ojastii (A) (N)
Sturnira magna
Sturnira tildae
Asidoptera sp. 1 (A)
Vespertilionidae
Myotis riparius
Anatrichobius Sp. (A)
Dermanura glauca
Promops centralis
Paradyschiria parvula (A)
Echimyidae sp. 1
ORDEN RODENTIA
Muridae
Oryzomys nitidus
Ixodes sp. 1 (L)
Polygenis cf. occidentalis (A)
Oryzomys capito
Polygenis cf. roberti (A)
Xenopsylla cheopis (A)
Amblyomma sp. 1 (L) (N)
Sphictopsylla sp. 1 (A)
Oryzomys sp. 1
Eutrombicula sp. 1(L)
Polygenis cf. occidentalis (A)
Oecomys bicolor
Xenospsylla cheopis (A)
Basilia myotis (A)
Basilia cf. peruvia (A)
Basilia manu (A)
Basilia sp. 1 (A)
Proechimys spinosus
Amblyomma sp. 1 (L) (L) (L)
Rhopalopsyllus sp. 1 (A)
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Biodiversity Assessment and Long-Term Monitoring, Lower Urubamba Region, Perú 1997

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Biodiversity

The Instituto Nacional de Recursos

Biodiversity Assessment & Monitoring, SI/MAB Series #1 (1997)

gas and condensate resources.

cluded representatives of government institutions, non-governmental organizations, the scientific

Machiguenga communities assisted

The Well Sites

The San Martin-3 and

SPDP & Smithsonian/MAB Biodiversity Program, Lower Urubamba, Per

April of 1997, at the same time that

Overall, we found that the

habitat losses, 2) changes in the

Biodiversity Assessment & Monitoring, SI/MAB Series #1 (1997)

spiders, and snails, among others).

dium- and large-sized mammals

Based on our findings to date,

SPDP & Smithsonian/MAB Biodiversity Program, Lower Urubamba, Per

The Longer Term

The 170,000-ha study area

and Camisea) is located east of the

Biodiversity Assessment & Monitoring, SI/MAB Series #1 (1997)

distribution of this landscape is in

groups, most of whom are

SPDP & Smithsonian/MAB Biodiversity Program, Lower Urubamba, Per

Figure 1. Adaptive management cycle for developing biodiversity assessment

Aspects of the adaptive management processsetting objectives, carrying out an assessment

ment gives managers the flexibility

Biodiversity Assessment & Monitoring, SI/MAB Series #1 (1997)

Figure 2. SI/MAB framework for forest biodiversity monitoring.

which all future changes may be

SPDP & Smithsonian/MAB Biodiversity Program, Lower Urubamba, Per

related taxa are changing, and early

The Lower Urubamba project

monitored because of cost constraints, but scientists and project

Biodiversity Assessment & Monitoring, SI/MAB Series #1 (1997)

Figure 3. Configuration of SI/MAB biodiversity assessment and monitoring test plots.

plots will be needed in other strata

SPDP & Smithsonian/MAB Biodiversity Program, Lower Urubamba, Per

The flora of southeastern Per

were set up at San Martin-3; plot #3

Biodiversity Assessment & Monitoring, SI/MAB Series #1 (1997)

quadrats. Two team members stand

mapped in the plot. Information

SPDP & Smithsonian/MAB Biodiversity Program, Lower Urubamba, Per

species that are commonly found in

boo down and that this in turn had

Biodiversity Assessment & Monitoring, SI/MAB Series #1 (1997)

In discussions of animal diversity, most people tend to think of

Total Basal Area

Urubamba and SI/MAB plots in the

SPDP & Smithsonian/MAB Biodiversity Program, Lower Urubamba, Per

Figure 8. The known global biota.

Most arthropod species are

Biodiversity Assessment & Monitoring, SI/MAB Series #1 (1997)

it, the understory layer, and the first

tors. Butterflies are highly dependent on the microclimate conditions

SPDP & Smithsonian/MAB Biodiversity Program, Lower Urubamba, Per

regions in Per for nocturnal butterflies.