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Small mammal communities in an eastern Brazilian park

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Title:
Small mammal communities in an eastern Brazilian park
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Stallings, Jody R., 1954-
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English
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ix, 200 leaves : ill. ; 28 cm.

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Subjects / Keywords:
Epiphytes ( jstor )
Forest habitats ( jstor )
Forestry ( jstor )
Forests ( jstor )
Mammals ( jstor )
Marsupials ( jstor )
Primary forests ( jstor )
Seasons ( jstor )
Species ( jstor )
Trees ( jstor )
Habitat (Ecology) -- Brazil -- Rio Doce State Forestry Park ( lcsh )
Mammals -- Brazil -- Rio Doce State Forestry Park ( lcsh )
Mammals -- Habitat ( lcsh )
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bibliography ( marcgt )
theses ( marcgt )
non-fiction ( marcgt )

Notes

Thesis:
Thesis (Ph. D.)--University of Florida, 1988.
Bibliography:
Includes bibliographical references (leaves 187-199).
General Note:
Typescript.
General Note:
Vita.
Statement of Responsibility:
by Jody R. Stallings.

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University of Florida
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University of Florida
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Copyright [name of dissertation author]. Permission granted to the University of Florida to digitize, archive and distribute this item for non-profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder.
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AA00004827_00001 ( sobekcm )

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SMALL MAMMAL COMMUNITIES
IN AN EASTERN BRAZILIAN PARK

















By

JODY R. STALLINGS


A THESIS PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN
PARTIAL FULFILLMENT OF THE REQUIREMENTS
FOR THE DEGREE OF DOCTOR OF PHILOSOPHY


UNIVERSITY OF FLORIDA


1988




SMALL MAMMAL COMMUNITIES
IN AN EASTERN BRAZILIAN PARK
By
JODY R. STALLINGS
A THESIS PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN
PARTIAL FULFILLMENT OF THE REQUIREMENTS
FOR THE DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
1988


ACKNOWLEDGEMENTS
This project could not have been carried out without the help
of numerous people. Foremost, I thank John G. Robinson for
suggesting the research project in the Rio Doce State Forestry
Park. His perception and patience aided 1n all aspects of the
project and his visit to the project site in May, 1986, was a great
boost to my morale. I would also like to thank the other members
of my committee for their suggestions during the planning stage and
for helping me to interpret the data. I thank Drs. John F.
Eisenberg, Melvin E. Sunquist, Wayne R. Marion, and Nigel J. H.
Smith. John Eisenberg was a constant source of support through the
analysis and writing stages. His continued interest 1n the results
and concern for my welfare helped to carry me through difficult
periods. Kent H. Redford provided obscure and relevant
publications. He also visited the field site at the beginning of
the project and encouraged me to sample the eucalypt forests.
Funding for this project was provided from the Program for
Studies 1n Tropical Conservation at the University of Florida,
World Wildlife Fund-US, The Organization of American States, and
the Instituto Estadual de Florestas (IEF). I thank John Robinson,
Kent Redford, Russell Mittermeier, John Eisenberg, Jose Carlos


Carvalho, and Celio Valle for helping me to obtain the funds
necessary to carry out the project.
I would like to thank Jose Carlos Carvalho, president of IEF,
for allowing me to work in the R1o Doce State Forestry Park. Dr.
Celio Valle, limar Bastos Santos, and Gustavo A. B. da Fonseca were
very instrumental in helping me to obtain my scientific expedition
visa to conduct research 1n Brazil. Celio Valle and Gustavo
Fonseca were more than helpful throughout the various stages of
fieldwork and allowed me to work in the mammalogy laboratory in
Belo Horizonte. Gustavo Fonseca was more than generous 1n sharing
his computers with me. Dr. Wilson Mayrlnk and his excellent staff
treated me in a concerned and professional manner. I thank them
for saving my nose.
The professional staff at the R1o Doce State Forestry Park
contributed to the project 1n several ways. Ademlr Camara Lopes,
the park Administrator, helped us get settled into the Park. He
offered no cost housing for the duration of the project and
contributed a modest amount of the Parks gasoline supply for my
use 1n the project. He also was able to obtain funds from IEF for
the construction of arboreal platforms and 60 wire traps.
Hermogenes Ferreira S. Neto was extremely important 1n solving
daily problems and 1n handling our correspondence. Jose Lourenco
Ladeira, the park dendrologist, identified the shrub and tree
species that occurred within each trapping post. I am very
grateful for his participation and was amazed at his knowledge of
the taxonomy.


I thank Drs. Phil Myers, Mike Carleton, Guy Musser, Al
Gardner, and James Patton for identifying the voucher specimens.
Their prompt reply enabled me to begin the analysis shortly after
returning to the states.
I relied on several workers from the R1o Doce park to
implement the field project. Trails were opened in an expert
manner by Ivanil Moreira and Waldemar Queroga dos Santos. Ivanil
later became my field assistant and proved to be an extremely
valuable asset. He never once complained because of the Incredible
numbers of ticks or because of all the small mammals that bit
chunks out of h1s fingers. I also thank Lldair Rufino for climbing
42 trees and placing the arboreal platforms.
Several students from the Departamento de Ecologia and
Zoologia participated 1n the fieldwork. I was aided greatly in the
data collection by 3 students. I thank Ludmilla Aguiar, Eduardo
Lima Sabato, and Luiz Paulo de Souza Pinto. I could not have
collected the appropriate data without their help. I also thank
Sonia Riquiera for her effort in helping us adjust to Minas Gerais,
and for participating in the initial stages of the fieldwork. She
was a pleasure to work with and always brought a positive attitude
to unpleasant working conditions.
My stay in Brazil was greatly facilitated by Gustavo and Ana
Fonseca. Their hospitality was more than generous and they always
opened their door to me and my family. Their hospitality was more
than "mineiro", it was more of true friendship. My wife and I will
iv


always be grateful to all that the Fonsecas did to help us get
adjusted in Brazil.
To my wife and best friend, Cathie, I offer my deepest
gratitude for her patience, understanding, and moral support during
the 12 month field project. It was always a pleasure to come home
from the field and be met by a smiling and positive person. I am
very appreciative for Cathies participation 1n the project and
wish that she could have had more freedom to pursue her own
interests.
v


TABLE OF CONTENTS
Page
ACKNOWLEDGEMENTS i1
ABSTRACT viii
CHAPTERS
I INTRODUCTION 1
General Background 1
Study Organization 3
II SMALL MAMMAL INVENTORIES IN AN EASTERN BRAZILIAN PARK 5
Introduction 5
Study Site 7
Methods 11
Results 22
Discussion 48
III TEMPORAL VARIATION IN TRAPPING SUCCESS OF
DIDELPHID MARSUPIALS IN AN EASTERN BRAZILIAN PARK 57
Introduction 57
Materials and Methods 58
Results 66
Discussion 81
IV SMALL MAMMAL ASSOCIATIONS AND MICROHABITAT
SELECTION IN AN EASTERN BRAZILIAN PARK 88
Introduction 88
Materials and Methods 89
Results 197
Discussion 114
V FOREST FIRE AS A DETERMINANT OF SMALL MAMMAL
DIVERSITY IN A BRAZILIAN FOREST 122
Introduction 122
Materials and Methods 126
VI


Results 138
Discussion 155
VI CONCLUSIONS AND SYNTHESIS 160
APPENDIX 165
LITERATURE CITED 187
BIOGRAPHICAL SKETCH 200
vi i


Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy
SHALL MAMMAL COMMUNITIES IN AN EASTERN BRAZILIAN PARK
By
JODY R. STALLINGS
August 1988
Chairman: John G. Robinson
Major Department: Wildlife and Range Sciences
Forest Resources and Conservation
This dissertation examines small mammal communities in native
and non-native habitats in one of the largest remaining forested
tracts of land in the Brazilian Atlantic Forest. Small mammals
were live-trapped over a period of 12 months in grass, eucalypt,
and native forested habitats. Eucalypt and native forested
habitats were rich in rodent species, but marsupials were
numerically higher in terms of relative densities. Grass habitats
were rich in rodent species and were dominated by one rodent
species. Trapping success for marsupials was observed to increase
during the sharp dry season. Paucity of food resources may be the
factor responsible for the observed increase in trapping success.
The Rio Doce State Forestry Park has been subjected to frequent
crown fire in the recent past, and relatively little primary forest
remains. Marsupials may dominate in terms of abundance in this
vi i i


Park because of the large amount of secondary forest. Small mammal
species diversity was calculated from forest stands that were
burned completely to the ground, in stands that were burned in a
mosaic fashion, and 1n primary stands. Species diversity was found
to be higher in intermediately fire disturbed native forested
habitats in comparison to heavily disturbed and primary forested
habitats. Mosaic habitats, composed of both secondary and primary
forest seres, offer suitable habitat to more species than either
secondary or primary forest stands alone.
IX


CHAPTER I
INTRODUCTION
General Background
The Atlantic Forest of eastern Brazil 1s one of the most
threatened ecosystems 1n the World due to Intense alteration of the
forested habitat (Mittermeier et al., 1982; Fonseca, 1983).
Fonseca (1983) estimates that approximately 3 5X of the native
forest remains today, with most of the remaining forest patches
occurring in protected areas and on privately owned lands. These
remaining forests are composed of mixed stands of primary, and
mostly, secondary forest seres. The Impact of these isolated and
heterogeneous forests on the wildlife communities has not been
studied. Only preliminary surveys have been conducted for the
primates that occur in this region and the remaining vertebrate
fauna is poorly known. This study focuses on small mammal
communities 1n a variety of habitats in one of the largest
remaining protected areas in the Atlantic Forest of eastern Brazil.
This study examines temporal and spatial variation in small mammal
communities in a region severely threatened by human activities.
1


2
The notion of prevailing stable environmental conditions 1n
the tropics helped to support the belief that tropical species
exhibit less temporal variation 1n reproduction, activity, and
feeding than temperate species. A current body of data reveals
that the tropics are much more dynamic than previously believed
(e.g., Leigh et al., 1982). This 1s especially true of tropical
areas that experience pronounced wet and dry seasons. The
Brazilian Atlantic region has such pronounced seasons. With this
in mind, this study Investigated the effect of time on the trapping
success of small mammals.
Habitat structure has been shown to affect the species
composition and relative abundance of small mammals in forested
(Dueser and Shugart, 1978; August, 1984) as well as 1n non-
forested systems (August, 1984; Lacher et al., in press; Lacher and
Alho, 1988). Tropical forest small mammals form a diverse group
that has a varied diet, use of vertical space, and activity time.
In addition, South American small mammals can be composed of two
large groups, rodents and marsupials, in comparison to North
American communities. These factors will be taken Into
consideration in determining how these small mammals allocate the
available space and resources in forest stands of differing
structure.
In some forest communities, disturbances can help to increase
floristic diversity (Connell, 1978; Horn, 1974). Connell (1978)
suggested that intermediate disturbances promote species diversity
in tropical forests and in coral reefs. This pattern has not been


3
observed for tropical vertebrate fauna, however, the relationship
is Intuitive. Early secondary and primary forests are
characterized as spatially homogeneous, while forests disturbed in
an intermediate fashion are spatially heterogeneous. Forest seres
characterized by a high degree of spatial heterogeneity should
provide more resources for more species in comparison to relatively
homogeneous environments. This study Investigated small mammal
communities 1n a variety of forest habitats that were affected by
forest fire.
Study Organization
This dissertation 1s divided into four major chapters. Each
chapter reads as an Independent paper, but all of the chapters have
a common theme. The theme 1s the small mammal communities in the
Rio Doce State Forestry Park 1n eastern Minas Gerais, Brazil. This
Park lies in the Rio Doce Valley and is part of the highly
endangered Atlantic Forest system of eastern Brazil. The primate
fauna of this region 1s highly endemic and very endangered
(Mittermeier et al., 1982). Inventories of other mammal species
are lacking for this region, although, preliminary species lists
indicated a highly endemic small mammal fauna.
Chapter II 1s concerned with intensive small mammal
inventories in this park. This chapter describes the major habitat
types that were sampled, the sampling methodology, the effect of
trap position and type, and the small mammals that occurred in each


4
major habitat type. Chapter II sets the stage for the subsequent
chapters.
Chapter III examines the temporal variation 1n trapping
success of marsupials. Many species of small mammals have a
"window of vulnerability" or a period of time during the year when
they are trapped with greater frequency. This chapter addresses
this question by examining the pattern of trapping success of
didelphld marsupials through time.
The fourth chapter addresses the effect of habitat structure
on the use of space by small mammals. I sampled small mammal
communities 1n 5 forested habitats. Individual species were
compared by their use of vertical space, diet, and time of
activity.
Chapter V examines the effect of forest fire in structuring
the small mammal communities 1n the R1o Doce Park. Tropical forest
fire has not been recognized as a determinant of species diversity.
In this chapter I suggest that even a low frequency fire can set
back succession 1n a forest system that may require 300 years to
reach "climax conditions. Fires in humid forests can occur
because of unseasonably dry periods that produce optimal conditions
for intense crown fires. Gaps or patches produced by fires can
affect species diversity of small mammals. Disturbances of an
intermediate fashion, such as fire mosaics, should Increase species
diversity because it produces a mixture of successional seres that
are suitable for a number of small mammal species.


CHAPTER II
SMALL MAMMAL INVENTORIES IN AN EASTERN BRAZILIAN PARK
Introduction
In comparison to its geographical size, few studies have been
conducted on small mammal communities 1n Brazil. Most published
reports on Brazilian mammals are preliminary species lists (e.g.
Av1la-P1res and Gouvea, 1977) or inventories (e.g., Vieira, 1955;
Moojen, 1952). Some Brazilian studies have focused on densities
(Emmons, 1984), others have focused on abiotic effects on small
mammals (Borchert and Hansen, 1983; Peterson, in press), while
others have addressed such diverse subjects as plantation effects
(Dietz et al., 1975) and public health needs or mammals that carry
human diseases (e.g., Laemment et al., 1946; Botelho and Linardi,
1980; Dias, 1982).
Recently, work 1n Brazil using mark-release techniques has
addressed the use of space, longevity, diversity and social habits
of small mammals (see Alho, 1982). As Alho (1982) pointed out,
most of these studies were concentrated in the xerophitic Cerrado
and Caatinga habitats. Fewer studies have been carried out 1n the
humid forests. Carvalho (1965) live trapped small mammals in a
tropical humid forest in Sao Paulo. Malcolm (pers. comm.) trapped
small mammals in tropical humid forest near Manaus, Amazonia.
5


6
Fonseca (pers. comm.) worked on small mammals in a range of
habitat types 1n eastern Minas Gerais.
The purpose of this paper 1s to report on intensive small
mammal inventories in a tropical humid Brazilian forest, the Rio
Doce State Forestry Park. This Park occurs within the geographical
limits of the highly endangered Brazilian Atlantic Forest (Fonseca,
1983). Small mammal Inventories are lacking from this park.
Gastal (1982) and Avila-P1res (1978) report preliminary results
from intermittent small mammal Inventories in the Rio Doce park.
The Brazilian Atlantic forest has a highly diverse flora and fauna,
with many endemic species of trees (Mori et al., 1981), reptiles
(Muller, 1973), and birds (Haffer, 1974). In-depth mammal
inventories in the region are lacking. The mammalian fauna is
poorly known. Mittermeier et al. (1982) and Kinzey (1982) report
on the high level of diversity and endemism found in the primates
of the region. Preliminary species lists for non-volant mammals in
this region suggest a very high diversity and endemism (Cabrera,
1957, 1961; Honacki et al., 1982; Moojen, 1952; and Vieira, 1955).
For this project, a preliminary checklist was prepared of the non
volant mammal species that probably occur in the Atlantic Forest
region (Table A-1). These data Indicate that for the region there
are at least 131 species, 50 of which (39%) are endemic.


7
Study Site
Small mammal trapping was carried out 1n the R1o Doce State
Forestry Park (19 4818" and 19 2924" south latitude and 42
3830" and 42 2818 west longitude). The Park was created in 1944
at the request of Dorn Helvecio, the bishop of the region (Gllhaus,
1986). The State Forestry Institute of the state of Minas Gerais
is the present administrative body.
The climate of the Park 1s classified as tropical humid
(Gllhaus, 1986) with a seasonal pulse of precipitation from
November through February and a pronounced dry season from June
through August. Average annual rainfall for a 20 year period
(CETEC, 1981) was 1480 mm, although the rainfall recorded during
this year of the study was considerably less (Figure 2-1). Mean
annual temperature averages 22 C (CETEC, 1981) and mean minimum
monthly temperatures vary greatly throughout the year (Figure 2-2).
The Park boundaries on the north and the east are two rivers,
the R1o P1rac1caba and the R1o Doce, respectively (Figure 2-3).
The southern and western boundaries abut plantations of Eucalyptus
spp. forests.
The predominant relief forms in the Park are hills originating
from fluvial plane dissection and valleys derived from fluvial
deposits (Gllhaus, 1986). The altitude in the Park varies from 230
to 515 m. CETEC (1981) reports that 21% of the Park is composed of
plains, 40% undulating to strongly undulating hills and 34%
strongly undulating hills to mountainous terrain.


8
WALTER AND L
CUMATC DIAGRAM
400
350 -
30C -
250 -
200
150
TOO
50 -
MONTHLY RAINFALL (MM.)
MEAN TEMPERATURE (0 )
JAN PEB MAP APR MAY jun JUL AUQ SEP 00T NOV OEC
MONTHS
AVERAGE ANNUAL RAINFALL 1400 mm
OATA rPGM :364-74
WALTER AND LEITH
CUMATSC DIAGRAM
MONTHS
ANNUAL AVERAGE RAINFALL 804 mm
3ata from iQee-taae
Figure 2-1. Walter and Leith climatic diagram characterizing
precipitation surplus and drought per month 1n Rio Doce State
Forestry Park, Minas Gerais, Brazil. A= data collected for a
twenty year period (1954-1974) and B= data collected during
study.


9
TEMPERATURE (C )
MONTHS
MEAN MAXIMUM TEMP MEAN MINIMUM TEMP
DATA FROM 1954 1974
Figure 2-2. Temperature graph showing pronounced decrease 1n
minimum temperature during June, July, and August in Rio Doce State
Forestry Park, Minas Gerais, Brazil.


10
RIO DOCE STATE
FORESTRY PARK
LEGEND
sites (3)
burned areas
eucalyptus
N
01234 km
Figure 2-3. Map of Rio Doce State Forestry Park, Minas Gerais,
Brazil. Numbers circled indicate trapping sites.


11
A unique feature of the R1o Doce State Forestry Park (PFERD)
1s the system of R1o Doce Valley lakes that occur there. There are
approximately 40 lakes and numerous marshes in the Park (Saijo and
Tundisi, 1985). According to Saijo and Tundlsi (1985) the lakes
were formed by damming the drainage river of the Rio Doce
watershed. The marshes in the Park are the result of the
sedimentation of previous lakes.
The vegetation of the park 1s classified as tropical semi-
deciduous (Gilhaus, 1986). Most of the emergent tree species lose
their leaves during the cool dry months (J. Stallings, per. obs.).
Forest fire has been a major threat to the vegetation and wildlife
in the park because of the Utter that accumulates during the dry
season. In 1964 and 1967, major fires burned approximately 30* of
the park (Lopes, 1982; Silva-Neto, 1984).
Gilhaus (1986) described 5 forested and 5 open/field habitats
for the Park (Table 2-1). All of the forested and open/field habitats
have been altered to some extent by fire, with the exception of the
Tall Primary Forest with Epiphytes.
Methods
I live-trapped small mammals in 4 general habitas types: (1)
homogeneous eucalypt forest, (2) eucalypt forest with native species
subcanopy, (3) wet meadow, and (4) native forested habitats. I
trapped in 5 sites within the native forested habitat type. Two of
the sites (RD/C and RD/T) were primary forests and corresponded to
the Gilhaus (1986) classification of Tall Primary Forest with


12
Epiphytes. Two other sites (RD/H and RD/M) were altered by forest
fire in 1967 and burned in an intermediate fashion which produced a
forest mosaic of short, secondary and tall forest. This habitat type
corresponded to the Medium to Tall Forest with Bamboos and
Graminoids (Gilhaus, 1986). The remaining site (RD/F) was burned
completely to the ground in 1967 and the resulting vegetation type
corresponded to the Medium Secondary Forest with Bamboos and
Graminoids (Gilhaus, 1986).
The wet meadow habitat (RD/B) corresponded to both the Low
Woodland and Low Tree and Scrub Tallgrass Savanna classified by
Gilhaus (1986). This habitat type occurs between the edge of
permanent marshes and secondary forest. Grasses from the family of
Graminae are the dominant vegetative cover and were introduced in
the region as food for cattle.
The eucalypt forest with native subcanopy habitat was planted
with Eucalyptus saligna in 1954 after the original vegetative cover
was removed. The eucalypt forest was harvested selectively in 1964
and again in 1971. However, the eucalypt forest was never clear-cut
and the native species were allowed to regenerate, largely through
coppicing, and developed into a complex native subcanopy. The result
is a homogeneous eucalypt upper canopy and a native species
subcanopy or "mata su.ia." Tall exotic grasses cover the ground.
Emergent eucalypt trees reach 20 m in height.
The homogeneous eucalypt forest was first planted with
Eucalyptus saligna in 1954 and was harvested 3 times on a 7 year
rotation. There is no native woody herbaceous subcanopy, only


13
Table 2-1. Forested and open/field habitats that occur in the Rio
Doce State Forestry Park, Minas Gerais, Brazil. Definitions and
characteristics of habitats taken from Gilhaus (1986).
HABITAT TYPE
% TOTAL
COMPARISON TO
THIS STUDY
FORESTED
TALL PRIMARY FOREST
WITH EPIPHYTES
8.4
RD/C, RD/T
PRIMARY FOREST
TALL FOREST
30.0

MEDIUM TO TALL FOREST WITH
BAMBOOS AND GRAMINOIDS
30.6
RD/M, RD/H
MOSAIC FOREST
MEDIUM SECONDARY FOREST WITH
BAMBOOS AND GRAMINOIDS
17.2
RD/F
LOW SECONDARY
FOREST
LOW SECONDARY FOREST
0.1

OPEN/FIELD
LOW WOODLAND
1.1
RD/B
WET MEADOW
LOW TREE AND SCRUB TALLGRASS
SAVANNA
0.6
RD/B
WET MEADOW
TALLFERN FIELD
0.1

EVERGREEN TALLGRASS FIELD
WITH TvDha sd.
3.0

PARTIALLY SUBMERGED
SHORTHERB FIELD
AND
AQUATIC HABITAT
8.9
100.0



14
eucalypt trees. Tall exotic grasses are the dominant ground cover.
Eucalypt trees reach approximately 15 m in height.
Individuals were snap trapped in order to obtain voucher
specimens, dietary, and reproductive information. Snap trapping was
carried out exclusively in wet meadow and secondary succession
habitats. All specimens were either preserved in 10% formalin or
made into museum study skins with corresponding complete skulls. I
also made study skins of Individuals of species of uncertain taxonomic
status that were live trapped in habitats other than those where snap
trapping was carried out.
I placed one line of terrestrial Sherman live traps 1n the
homogeneous eucalypt habitat. Twenty-five Shermans, baited with dry
oatmeal, pineapple chunks and cotton balls soaked with cod-liver oil,
were opened for 4 consecutive nights on 5 occasions through the
study. Trap stations were separated by 15 m.
In the wet meadow habitat, live trapping started in February,
1986, and continued at monthly intervals through January, 1987. I
used two parallel trapping lines in this habitat. Each line was 280 m
in length and subdivided into 20 trap stations separated by 15 m. All
traps were placed on the ground, with even numbered stations having
only one Sherman Uve trap while odd numbered stations had one
Sherman and one locally made small wire live trap. Bait was
identical to that used in the homogeneous eucalypt forest.
In the eucalypt forest with native subcanopy and the native
forested habitats, the trapping design was identical (Table 2-2). In
each area, I cut 3 parallel lines 300 m in length through the forest.


15
Table 2-2. Trapping design used in native forested and eucalypt
forest with native species subcanopy habitats. Trapping lines are
A, B, and C. Numbers 1 16 represent trapping posts. Terrestrial
medium sized live traps = ¡, arboreal medium sized live traps = *,
small arboreal Sherman live traps = S, small terrestrial Sherman
live trap = s, large terrestrial live trap = L. Trapping lines
were separated by 100 m.
TRAPPING
STATION
TRAPPING LINES
A
B
C
1 *\ ¡S
2 L;S *\
3 *| !s
4 L ¡ s *|
5 *¡ ¡s
6 l;s *¡
7 *: is
8 L| s *|
9 *| | S
10 L;S *|
11 *l Is
12 L|s *|
13 *| | S
14 L¡S *|
15 *| |s
L | S I
16


16
Sixteen trapping stations were placed along the line separated by 20
m. I used Sherman live traps and locally made small (15 X 15 X 30
cm) and large (25 X 30 X 60 cm) wire live traps. All traps were
placed within 3.5 m of the trapping post. All trapping posts had a
terrestrial small Uve trap. Odd numbered trapping posts had a small
wire live trap placed in a tree or bush. Mean arboreal live trap
height was 1.2 m. Odd numbered trapping posts had a Sherman live
trap alternating between arboreal and terrestrial positions. The
exterior trapping lines were identical with respect to number, kind
and placement of traps. I did not use the large live traps on the
interior line. Aside from the large live traps, the exterior and
interior lines were equal in trap number. However, the positions of
the Sherman and small live wire traps were reversed for the interior
line. The Sherman live traps were introduced Into each of the
forested habitats after the study was well underway in an attempt to
sample smaller bodied species. I placed Shermans in two of the
native forested sites in January, 1986, and introduced Shermans in the
remaining forested sites in May of the same year.
I also experimented with traps that were placed high in the
canopy by a pulley and platform device. This method was similar to
that developed by Malcolm (pers. comm.) for use in the Brazilian
Amazon. Mean trap height was 11.2 m. I used these arboreal traps
to sample the canopy dwelling small mammals. Traps were located at
trapping posts along the established lines. The post and exact trap
location in each tree were determined in a subjective manner. I
placed traps in trees which I thought had a high degree of canopy


17
connectivity and upper stratum vine density. Forty-two arboreal
platforms were spread across 4 native forested sites. The primary
forest sites, RD/C and RD/T, and one of the mosaic sites, RD/H,
each had 12 arboreal traps, 4 traps per line. The other mosaic site,
RD/M, had only 6 traps because I did not believe that there was
sufficient upper strata development to support canopy dwelling
species. Arboreal canopy trapping started in June, 1986, and
continued through October, 1986. Trapping coincided with the
schedule developed for the terrestrial and arboreal trapping.
I used dry oatmeal, pineapple chunks and cotton balls soaked
with cod-liver oil for bait. Traps were set during the day and
remained open for 5 consecutive nights each month for one calendar
year.
The first occasion that an individual was trapped was considered
a first capture. The first capture plus subsequent captures of each
individual were considered total captures. Minimum known alive
(MKA) was the number of individuals actually captured each month
whether the capture was the first capture or a recapture from a
different session.
Trapping success of small mammals was calculated in the
following manner. I multiplied the number of traps by the number of
nights the traps were baited and armed per site per month to
determine the number of trap nights. Trapping success was the
number of individuals, MKA or total captures of all species divided by
the number of trap nights and expressed in percentages. For
example, if 100 individuals were trapped during 1000 trap nights, the


18
trapping success would be (100/1000) X 100 = 10X. Recapture indices
were calculated by dividing total captures by first captures and
indicated the average number of times an individual of each species
was captured.
I recorded the following information from each captured small
mammal: date, location on trapping line, position of trap, species, sex,
whether the trapped animal was a juvenile or adult, its reproductive
condition, general condition, external parasitic load on a relative
scale, and behavior upon release. I also recorded standard body
measurements for each Individual: body length, tail, ear, hind foot
and mass. I placed a numbered metal eartag in the left ear of each
individual upon the initial trapping of the individual.
Taxonomic determination of questionable species was made by
taxonomists specializing 1n various small mammal groups. Voucher
specimens were brought to the United States and the taxa were
distributed to the following people: cricetid rodents and marsupials of
the genus Marmosa were sent to Dr. Phil Myers, University of
Michigan; rodents of the subgenus Oecomys were sent to Dr. Guy
Musser, American Museum of Natural History, and to Dr. Mike
Carleton, U.S. National Museum of Natural History.
I used body measurements, weight, reproductive condition and
pelage characteristics to determine the age class (juvenile or adult)
of each captured Individual. An individual was considered an adult if
it was reproductively active. Female rodents were considered
reproductively active if they 1) had a perforated vulva, 2) were
pregnant, or 3) were lactating. Marsupial females were considered


19
reproductive!y active if they 1) were lactating or 2) had young
attached to the teat field. Hale rodents were considered
reproductively active if the testes were descended. I could not
determine the reproductive status of male marsupials as the testes are
permanently descended. However, the activity state of the sternal
gland in marsupials can indicate the reproductive time of year.
Initially, I used the overall gestalt of each individual of each sex to
assign age classes. Later, I compared my initial classification with
the body measurements and mass. Body measurements and weights
were sorted for each sex of each species and plotted to determine if
recorded values could be grouped according to size. I then assigned
a body measurement value as the threshold for separating juvenile
and adult age classes. These age classes were then compared to the
initial age classes that were assigned in the field.
I used the General Linear Program (PC-SAS) ANOVA to test for
the equivalence of adult body measurements and mass means between
sexes for each species. This analysis enabled me to determine the
extent of sexual dimorphism for the external characters. Statistical
significance was set at < 0.05.
Feeding categories were determined by stomach content analysis
(Charles-Dominique et al., 1981) and from the literature. I relied
heavily on information gleaned from the literature on food
preferences of small Neotropical mammals.
The use of vertical space (1.e., terrestrial, scansorial, or
arboreal) of each species was determined from 3 data sets. I
compared the proportion of captures in trees to that of captures on


20
the ground for each species. There were more terrestrial than
arboreal traps and this bias was corrected by adjusting the number of
total trapping opportunities. For this adjustment, I divided the
number of arboreal total captures by the total arboreal opportunities
(or total arboreal trap nights). The same was done for terrestrial
total captures. Results of these two divisions were summed and each
respective result (e.g., arboreal) was divided into the sum. The result
generated adjusted percentage success of arboreal and terrestrial
captures. The sum of total captures was multiplied by the adjusted
percentage in order to generate adjusted number of captures per
trapping stratum (on the ground or in arborescent vegetation). These
adjustments also were made for each species at each site as well as
pooled adjustments across all sites. I tested the null hypothesis that
there was no difference in the proportion of arboreal and terrestrial
captures for each species across all habitats as well as within each
trapping site. Prior to using Students t-test, I arc-sin transformed
the proportions. I then compared the proportion of arboreal and
terrestrial responses upon release for each species across all habitat
types. These adjustments were made for all species in each habitat
type.
I used Students t-tests to test the null hypothesis that there
was no difference in the locomotory response upon release of each
species. These 2 data sets were then compared to determine if there
were any differences between where an individual was captured and
its locomotory behavior upon release across all habitat types. These
tests also were performed for each habitat sampled. A species was


21
considered to be arboreal 1f that species was found to have a high
proportion of arboreal captures and a high proportion of arboreal
behavior upon release. The opposite would be true for a terrestrial
species. A species would be scansorial if there were no significant
differences in the proportion of spatial captures and no significant
differences in the proportion of behaviors exhibited upon release. I
then compared my results obtained from the trapping data to the
available literature for each species.
I also recorded the presence and relative abundance of other
vertebrates while walking the trapping lines 1n each sampling area.
Upon encountering an animal, I recorded the species, distance in
meters from the trail, height in trees for arboreal species, hour of
day, group size and other natural history data. I moved at a pace of
approximately 1 km. per hour and covered a distance of 1050 m per
sampling area. These walks yielded 60 km of repeat censuses per
sampling area. This method does not allow for the computation of H
indices because it is difficult to ascertain the identity of individuals
being censused. Because of this problem, I only calculated species
richness per taxonomic group, number of encounters of each species
through time, and number of encounters of each group per linear km.
These data are presented 1n Table A-3.
I also superficially sampled the bat fauna of the Park. I
mounted mist nets in various habitat types (e.g., lake edge, secondary
habitat, primary habitats, and manmade structures) in order to
determine the species that occurred in the park. Nets were mounted
one hour before dusk and opened for approximately 4 hours each


22
night. Mist netting occurred on a sporadic basis. A species list of
bats obtained by these methods is presented in Table A-4.
Results
Species Accounts
I logged 1,308 captures of 17 species of small mammals in 40,490
trap nights. The small mammal fauna of the park consisted of 6
species of marsupials and 11 species of rodents. The diet, use of
vertical space and habitat requirements are presented in Table 2-3 for
each species captured during the study.
Marsupials - Family Didelphidae.
Didelphis marsupial is Linne. (1758). The black-eared opossum
ranges widely in South America from the Isthmus of Panama to
southern Brazil. This species occurs sympatrically with D. albiventris
throughout much of its range (Strellein, 1982). In the Rio Doce
Valley, however, D. marsupial is inhabits moister habitats, while D.
albiventris occurs in the cerrado vegetation (Valle and Varejao, 1981).
A. Gardner (pers. comm.) suggested that the form of D. marsupial is
found in eastern Brazil is distinct and should be referred to as D.
azarae. This species Inhabits brushy and forested habitats (Alho,
1982; Nowak and Paradiso, 1983). Miles et al. (1981) found this
species to be nocturnal, with a preference for nesting in tree
cavities. I captured this species in all forested habitats in the park
(Table 2-4). Adult body measurements do not indicate sexual
dimorphism (Table A-2). Females have a well developed pouch. This


23
Table 2-3. Ecological role played by each species captured during
this study in the Rio Doce State Forestry Park. GM= grasslands and
wet meadows, B= brushy areas, S= secondary forests, P= primary
forests, F= fossorial or semifossorial, SA= semiaquatic, T=
terrestrial, S= scansorial, A= arboreal, HG= herbivore-grazer; FG=
frugivore-granivore; F0= frugivore-omnivore; 10= insectivore-
omnivore. Taxa endemic to the Brazilian Atlantic rainforest or
the eastern coastal area of South America.
SPATIAL
DIETARY
SPECIES
HABITAT
ADAPTATION
CLASSIFICATION
MARSUPIALS
DidelDhis marsuDialis
B, S,
P
T, S
FO
Metachi rus nudicaudatus
s,
P
T
IO/FO
Marmosa incana*
B, S,
P
S
10
M. cinerea
B, S,
P
A
10
M. microtarsus*
s,
P
A
10
Caluromys philander
s,
P
A
FO
RODENTS
Oecomvs trinitatis
s,
P
S
FG
OrYZomvs caoito
s,
P
T
FG
0. subflavus*
GM, B,
S
T
FG
0. niqriDes*
GM,
B
S
FG
Akodon cursor*
GM, B,
S
T
10
Calomvs laucha
GM,
B
T
FG
Nectomvs sauamiDes
GM,
B
SA
HG
Abrawavaomvs ruschii*
S
T
FG?
Oxvmvcterus roberti*
GM, B,
S
F
10
Rhioidomvs mastacalis
s,
P
A
FG
Cavia fulqida*
GM,
B
T
HG
EurYZvqomatomvs SDinosus*
GM,
B
F
HG


24
Table 2-4. Number of total captures and percent of total for each
species (SPP.) per habitat type. Numbers in parentheses represent
the percentage of captures per species per habitat percentages are
rounded to the nearest whole number.
SPP*
HABITAT
TYPES+
RD/F
RD/H
RD/M
RD/T
RD/C
RD/E
RD/B
DM
4 (3)
2 (1)
1 (D
7(6)
21(24)
7 (4)
-
MN
20(14)
44(16)
21(15)
30(23)
25(28)
18(11)
-
MI
25(18)
75(27)
20(14)
28(22)
6 (7)
14 (9)
3 (1)
MC
74(52)
71(26)
69(49)
38(30)
31(35)
77(49)
1 (0)
MM
-
1 (0)
-
-
-
-
-
CP
-
16 (6)
2 (1)
13(10)
3 (3)
15(10)
2 (1)
NS
-
6 (2)
9 (6)
-
-
-
5 (1)
RM
-
-
-
7 (6)
-
-
-
AC
3(2)
46(17)
2(1)
-
1 (D
25(16)
315(85)
OT
4 (3)
8 (3)
4 (8)
3 (2)
2 (2)
-
-
OC
-
4 (1)
12 (9)
2 (2)
-
2 (1)
1 (0)
OS
12 (9)
1 (0)
-
-
-
-
31 (8)
ON
-
-
-
-
-
-
4 (1)
OR
-
1 (0)
2 (1)
-
-
-
4 (1)
AR
-
1 (0)
-
-
-
-
-
CL
-
-
-
-
-
4 (1)
ES
-
-
-
-
-
-
2 (1)
142
276
142
128
89
158
373
+ RD/F= SECONDARY HABITAT BURNED COMPLETELY TO THE GROUND; RD/H=
SECONDARY HABITAT BURNED IN MOSAIC FASHION; RD/M= SECONDARY HABITAT
BURNED IN MOSAIC FASHION; RD/T= PRIMARY FOREST WITH LITTLE EFFECT
FROM FOREST FIRE; RD/C= PRIMARY FOREST; RD/E= EUCALYPT FOREST WITH
NATIVE SPECIES SUBCANOPY; RD/B= WET MEADOW.
* DM= Didelphis marsupial is: MN= Metachi rus nudicaudatus: MI=
Marmosa incana: MC= Marmosa cinerea: MM= Marmosa microtarsus: CP=
Caluromys philander: NS= Nectomvs sauamioes: RM= Rhipidomvs
mastacalis: AC= Akodon cursor: 0T= Oecomvs trinitatis: 0C=
Oryzomys capito: 0S= Orvzomvs subflavus: 0N= Orvzomvs nigrioes:
OR= Oxymycterus roberti: AR= Abrawavaomvs ruschi i: CL= Calomvs
laucha: ES= Eurvzygomatomys soinosus.


25
species 1s terrestrial. There was a significant percentage of
terrestrial captures (Table 2-5) and terrestrial behavior upon release
(Table 2-6). I did observe several juvenile individuals and one adult
climb readily. Charles-Domlnique (1983) reported that this species
exploits the lower stratum 1n forests, but can climb. This species is
basically an opportunistic feeder and feeds upon fruit and animal
matter (Charles-Domiique, 1983).
Metachi rus nudicaudatus Geoffroy (1803). The brown four-eyed
opposum has a geographical distribution similar to that of £).
marsupial is except that it is not found over much of Venezuela nor
in northeastern Brazil (Streilein, 1982). M. nudicaudatus can be
confused with Philander opposum as both have pale spots above the
eyes. In addition, there is considerable confusion over the taxonomy
of the two species. Nowak and Paradiso (1983) classified this species
as Philander nudicaudatus and Philander opossum as Metachiroos
opossum. I agree with Honacki et al. (1982) and follow their
classification. This species was captured in all forested habitats
(Table 2-4). Metachi rus nudicaudatus is sexually dimorphic in Its
mass and hind foot measurements (Table A-2). Females do not have
a pouch. This species is strongly terrestrial, rarely caught in
arboreal traps (Table 2-5) and rarely climbs upon release (Table 2-6).
Miles et al. (1981) found this species to be nocturnal and construct
nests on the forest floor or in ground hollows. There are very little
data on the feeding habits of this species due to the small numbers
that have been reported to be trapped. Preliminary data indicate


26
Table 2-5. Students t-tests between adjusted and arcsin
transformed percentages of terrestrial and arboreal captures of
small mammals in all forest types. NS = non significant.
X TERRESTRIAL % ARBOREAL
SPECIES
CAPTURES
CAPTURES
V
P <
DidelDhis marsuoialis
66.6
23.5
40
.001
Metachi rus nudicaudatus
76.3
3.9
156
.001
Marmosa incana
43.9
46.1
169
NS
Marmosa cinerea
27.8
62.2
356
.001
Caluromvs Dhilander
16.3
73.8
49
.001
Nectomvs sauamioes
69.6
21.0
17
.05
RhiDidomvs mastacalis
28.8
61.1
5
NS
Akodon cursor
83.5
6.3
390
.001
Oecomvs trinitatis
31.2
58.8
19
NS
OrYzomvs caoito
62.9
27.0
19
.05
OrYzomvs subflavus
58.0
32.5
46
.01


27
Table 2-6. Results of Students t-tests between terrestrial and
arboreal behavior upon release of small mammals captured in all
habitats. NS= non significant. Species abbreviations are
explained in Table 2-4.
SPECIES
X
N
TERRESTRIAL
BEHAVIOR
% ARBOREAL
N BEHAVIOR
V
T
P <
DM
33
73.2
3
16.7
34
3.265
0.01
MN
150
80.7
4
9.1
152
4.934
0.001
MI
95
53.5
52
36.5
145
3.448
0.001
MC
11
10.8
294
78.9
303
7.743
0.001
CP
2
18.4
18
71.6
18
2.488
0.05
NS
12
90.0
0
0.0
10
10.882
0.001
RM
1
24.0
5
67.8
4
1.393
NS
AC
70
90.0
0
0.0
68
26.282
0.001
OT
9
64.8
2
25.3
9
1.763
NS
OC
13
90.0
0
0.0
11
11.326
0.001
OS
10
90.0
0
0.0
8
9.933
0.001


28
that this species is an insectivore-omnivore (Robinson and Redford,
1986) or frugivore-omnivore (Hunsaker, 1977).
Marmosa incana Lund 1840. This mouse opposum is endemic to the
Brazilian Atlantic Rainforest (Streilein, 1982). M. incana occurs in
both secondary and primary forest habitat. This species 1s small in
size (adults, average weight = 62 g) and strongly sexually dimorphic
in body size and color. Males tend to have larger ears and hind feet
(Table A-2) while females tend to have a more rose colored venter
and less pronounced face mask (P. Myers, pers. 1itt.). Females do not
have a true pouch. This species tends to use both the ground and
arborescent vegetation. I classify the spatial adaptation of this
species as scansorial (Table 2-3). There was no significant difference
in the proportion of terrestrial and arboreal captures (Table 2-5, p >
.90, df=169), however; individuals tended to remain on the ground
upon release (Table 2-6, p < 0.001, df=145). No data exist on the
feeding category of this species. Other species of Marmosa which
have similar body mass are classified as insectivore-omnivores.
Stomach content analysis (n=3) showed 100% insects from two orders,
Coleptera and Orthoptera (Table 2-7). I classify this species as an
insect1vore-omnivore based on the relationship found between body
mass and dietary classification (Robinson and Redford, 1986).


29
Table 2-7. Stomach content analysis of small mammals captured in
Rio Doce State Forestry Park, Minas Gerais, Brazil (N= number of
stomachs analyzed).
SPECIES
FRUIT SEEDS GRASS INSECTS
Marmosa incana (n=3)
Akodon cursor (n=23)
Orvzomvs subflavus (n=1)
Orvzomvs nigripes (n=5)
Oxvmvcterus roberti (n=2)
Calomvs laucha (n=1)
Nectomvs sauamipes (n=2)
19%
19%
8%
5%
-
95%
34%
11%
21%
100%
50% 50%
100%
52%
34%
100%


30
Marmosa cinrea Temminck (1824). M. cinrea has a disjunct
geographical distribution in South America; it occurs in northern
Venezuela through the Guianas, and it occurs in the Brazilian
Atlantic Rainforest extending into Paraguay (Streilien, 1982). M.
cinerea occurs in brushy and forested habitat, ranging from secondary
to primary. This species is a large bodied Marmosa (Table A-2,
average weight= 105g) and 1s highly sexually dimorphic based on
external body measurements. Males tended to have larger body, tail,
ear and foot (Table A-2). Females do not have a pouch. This
species is strongly arboreal and exploits the high forest stratum
(Charies-Domin1que, 1983; Miles et al., 1981). Miles et al. (1981)
found this species to be nocturnal and to construct open arboreal
nests rather than use cavities. M. cinerea tended to be caught a
greater proportion of the time in arboreal traps (Table 2-5, p <0.001,
df=356) and tended to exhibit arboreal rather than terrestrial behavior
upon release (Table 2-6, p < 0.001, df=303). This species is primarily
an Insectivore-omnivore (Robinson and Redford, 1986).
Marmosa microtarsus Wagner (1842). This species is restricted to the
Brazilian Atlantic Rainforest (Streilein, 1982). M. microtarsus differs
from its congener M. ag lis by possessing a pure colored white patch
of hairs on the throat and chin (Tate, 1933). There are insufficient
data in the literature to determine if this species is terrestrial,
scansorial, or arboreal. I only recorded one capture during the study,
in an intermediately disturbed habitat (Table 2-4). However, the
species has a long prehensile tail and short wide feet which suggest


31
an arboreal lifestyle. This species is probably an Insectivore-
omnlvore.
Caluromvs philander Linne (1758). This species has a disjunct
geographical distribution with populations in Venezuela, the Guianas
and northern Brazil and in southeastern Brazil (Streilein, 1982). The
woolly opossum is classified as a forest dwelling species (Nowak and
Paradiso, 1983). This species was present in the eucalypt, mosaic,
and primary forested habitats (Table 2-4). Based upon the external
body measurements, there was no sexual dimorphism in adults (Table
A-2). Females lack a true pouch. According to Charles-Dominique
(1983) and Miles et al. (1981) this species exploits the high forest
stratum and is nocturnal. My data showed that there was a higher
proportion of arboreal captures (Table 2-5, p< 0.001, df=49) and that
this species tended to climb more than remain on the ground upon
release (Table 2-6, p < 0.05, df=18). Fruit makes up a large portion
of this species diet (Charles-Dominique, 1983; Robinson and Redford,
1986).
Rodents Family Cricetidae
Oecomys (Oryzomys) trinitatus = (Q. concolor) Wagner (1845). This
genus is in need of revision and the subgenus Oecomys is currently
being revised (P. Myers, pers. comm.). This species was previously
called Orvzomvs concolor and was known, within Brazil, as an
Amazonian species (Alho, 1982). However, Nitikman and Mares (1987)
reported trapping this species in gallery forest in the Brazilian


32
cerrado. I captured this species in all native forested habitats (Table
2-4). The species is not sexually dimorphic (Table A-2).
Gyldenstolpe (1932) and Moojen (1952) stated that this species is
"more or less adapted for arboreal life." My data suggest that this
rat is scansorlal; there were no significant differences in the
proportion of terrestrial and arboreal captures (Table 2-5, p < .10,
df=19) and no significant differences in terrestrial and arboreal
behavior upon release (Table 2-6, p < .20, df= 9). Most species of
the genus Orvzomvs are frugivore-granivores (Robinson and Redford,
1986).
Oryzomys subflavus Wagner (1842). This species is distributed
throughout the Guianas, southeastern Brazil and eastern Paraguay
(Honackl et al., 1982; Alho, 1982). In Brazil, it occurs in the
cerrado, caatinga and Atlantic Rainforest (Alho, 1982). This species
was captured in wet meadow and heavily disturbed secondary habitat
(Table 2-4). There were no differences in body measurements
between sexes 1n adult individuals (Table A-2). I classify this species
as terrestrial. This species tended to be captured more on the
ground than in the trees (Table 2-5, p < 0.01, df=46) and was never
observed to climb upon release (Table 2-6). Stomach content analysis
(n=1) showed 95% grass and 5% fruit (Table 2-7).
Oryzomys capito Olfers (1818) = Q. goeldi. laticeps and intermedius.
This species has a wide distribution throughout the Neotropics and
occurs in a variety of habitats ranging from agricultural fields


33
(Moojen, 1952) to humid forests (Alho, 1982). Orvzomvs capito was
primarily captured in humid forests ranging from intermediate levels
of disturbance to primary forests in the park (Table 2-4). There
were no significant differences in body measurements between sexes
for adults (Table A-2). My capture and release data are in
accordance with Alhos (1982) classification for this species. 0.
capito tended to be caught more on the ground than 1n trees (Table
2-5, p < 0.05, df=19) and tended to remain on the ground upon
release (Table 2-6, p < 0.001, df=11).
Orvzomvs (01iqorvzomvs) nigripes (eliurus) Wagner (1845). This
small-bodied rodent (Table A-2) occurs in grassland, wet meadow and
secondary forest habitat in northern Argentina, eastern Paraguay,
southern Brazil and the Bolivian Beni (Honacki et al., 1982). In the
park, all captures were made in the wet meadow habitat (Table 2-4).
All captures were made on the ground (n=4), however; the Individuals
climbed readily in captivity (J. Stallings, pers. obs.). The results
from the stomach analysis (n=5) revealed a wide range of foodstuffs
(Table 2-7).
Abrawayaomvs ruschii Cunha and Cruz (1979). This species is only
known from the type locality in Espirito Santo, eastern Brazil. It is
endemic to the Brazilian Atlantic Rainforest. The single capture of
this species was recorded in an intermediately disturbed forest (Table
2-4). There is very little information available regarding the ecology


34
of this species and there are only three study skins found in
museums (A. Gardner, pers. comm.).
Rhipidomvs mastacalis Lund (1840). Climbing mice range south from
Margarita and Tobago Islands to Venezuela and Guianas to
northeastern and east central Brazil (Honacki et al., 1982). This
species was only captured in a relatively undisturbed primary forest
(Table 2-4). Sample size was too small to detect any differences
between terrestrial and arboreal captures and behavior upon release.
However, as the common name implies, this species climbs readily. I
captured two individuals 1n my house in the park, a commonly cited
exotic" habitat for this species (Nowak and Paradiso, 1983).
Nectomvs sauamipes Brants (1827). The neotropical water rat occurs
in aquatic habitats either in grasslands and wet meadows or in
forests. This species distribution ranges from the Guianas to
Colombia to Peru and in Brazil, Paraguay and northeastern Argentina
(Honacki et al., 1982). This species tended to be caught more on the
ground (Table 2-5, p < 0.05, df=17) and exhibited a significant
tendency to remain on the ground upon release (Table 2-6, p < 0.001,
df=10). Stomach content analysis (n=2) showed 50% grass and stems
and 50% fruit (Table 2-7).
Akodon cursor Winge (1887). This species occurs in several habitat
types from southeastern and central Brazil to Uruguay, Paraguay and
northern Argentina. The wet meadow habitat in the park was the


35
primary habitat to capture this species (Table 2-4). A. cursor was
formerly Included in A. arviculoides (Honacki et al., 1982). This
species is sexually dimorphic in tail (p < 0.01) and body (p < 0.008)
length (Table A-2). A. cursor is strongly terrestrial (Table 2-5 and
Table 2-6). Analysis of stomach contents (n=23) revealed a high
proportion of insects, seeds and fruit (Table 2-7).
Ca lorn vs laucha Olfers (1818).. This species occurs 1n grassland and
wet meadows in southern Bolivia, southeastern Brazil, Paraguay,
central Argentina and Uruguay. I only captured this species in the
wet meadow habitat 1n the Park (Table 2-4). The results from one
stomach sample revealed 100% seeds (Table 2-7).
Oxymycterus roberti Thomas (1901). The burrowing mouse occurs in
a variety of habitats but is usually associated with moist substrate in
open or brushy habitats. I captured this species in wet meadow and
secondary habitats in the Park (Table 2-4). This species is endemic
to eastern Brazil. This semifossorial mouse is described as an
insectivore (Nowak and Paradisio, 1983). Stomach analysis (n=2)
revealed 100% insects (Table 2-7).
Family Caviidae
Cavia fulgida Wagler (1831). This species of cavy is endemic to the
open grasslands and wet meadows of the Atlantic Rainforest of
eastern Brazil (Honacki et al., 1982; Nowak and Paradiso, 1983). I
captured this species in grassland and wet meadow habitats in the


36
Park, however; it was not trapped in site RD/B. For this reason this
species does not appear in Table 2-4. Cavies are terrestrial and are
herbivore-grazers (Nowak and Paradiso, 1983).
Family Echimyidae
Eurvzvgomatomvs soinosus Fischer (1814). The single species of
guiara is endemic to southeastern Brazil, Parguay and northeastern
Argentina (Honacki et al., 1982). I captured this species in the wet
meadow habitat (Table 2-4). This species inhabits open grasslands
and wet meadows, is considered terrestrial or semifossorial (Alho,
1982), and is most probably a herbivore-grazer.
Trapping results
Tables 2-8, A-5, and A-6 present the capture results by species
for the three main habitat types: native forested, eucalypt with
native species subcanopy and wet meadow habitats, respectively. As
a group, marsupials represented 79.2% and 83.3% of first and total
captures, respectively, in native forested sites, and 67.7% and 82.9% in
eucalypt forest with native species subcanopy. Rodents represented
97.3% and 98.4% of the first and total captures, respectively, in the
wet meadow habitat.
In the native forested habitat, Marmosa cinerea represented over
40% of the marsupial captures (Table 2-8), while in the eucalypt
forest, this species represented more than 58% of the marsupial
captures (Table A-5). Akodon cursor was the major contributor to
the rodent captures in all three habitats. This species only


37
Table 2-8. Capture results from native forested plots in Rio Doce
State Forestry Park, Minas Gerais, Brazil. RECAP INDEX= total
captures/first captures, and represents the average number of times
that an individual of species X is captured. Numbers in
parentheses represents percent of contribution of capture per
species per taxonomic group. Species abbreviations are explained
in Table 2-4.
TOTAL FIRST RECAP
SPECIES
CAPTURES
% TOTAL
CAPTURES
% TOTAL
INDEX
MARSUPIALS
DM
35
4.5 ( 5.4)
32
7.8 ( 9.9)
1.1
MN
140
18.0 (21.6)
91
22.3 (28.1)
1.5
MI
154
19.8 (23.8)
90
22.1 (27.8)
1.7
MC
283
36.4 (43.7)
92
22.5 (28.4)
3.1
MM
1
0.1 ( 0.2)
1
0.2 ( 0.3)
1.0
CP
34
0.4 ( 5.3)
18
4.4 ( 5.5)
1.9
647
83.2(100.0)
324
79.4(100.0)
RODENTS
NS
15
1.9 (11.5)
9
2.2 (10.7)
1.7
RM
7
0.9 ( 5.4)
3
0.7 ( 3.6)
2.3
AC
52
6.7 (40.0)
27
6.6 (32.1)
1.9
OT
21
2.7 (13.8)
19
4.7 (22.6)
1.1
OC
18
2.3 (13.8)
15
3.7 (17.9)
1.2
OS
13
1.7 (10.0)
7
1.7 ( 8.3)
1.9
OR
3
0.4 ( 2.3)
3
0.7 ( 3.6)
1.0
AR
1
0.1 ( 0.81
1
0.2 ( 1.2)
1.0
130
16.8(100.0)
84
20.6(100.0)


38
Table 2-9. Trapping success of small mammals calculated by habitat
type in Rio Doce
State Forestry Park,
Minas
Gerais, Brazil.
HABITAT TYPE
NUMBER OF
TRAP NIGHTS
NUMBER OF
CAPTURES % SUCCESS
NATIVE FOREST
(EXCLUDING PLATFORMS)
30,960
710
2.3
NATIVE FOREST
PLATFORMS ONLY
1,050
66
6.3
WET MEADOW
1,980
373
18.8
EUCALYPT FOREST
W/NATIVE SPECIES
SUBCANOPY
6,000
158
2.6
EUCALYPT FOREST
W/NO SUBCANOPY
TOTALS
500
40,490
1
1,308
0.0


39
represented about 7% of the total captures in the native forested
habitat, but 40* of the rodent captures (Table 2-8). In the eucalypt
forest, A. cursor represented about 16* of the total captures and 93*
of the rodent captures (Table A-5). This rodent was the dominant
species captured in the wet meadow habitat, representing about 85*
of both the total and of the rodent captures.
In both the native and eucalypt forested habitats, marsupials in
general were recaptured at a high rate. Didelphis marsupial is and
Marmosa microtarsus both showed low recapture rates and reflect the
small sample size. Especially noteworthy was the relatively high
recapture rate of Marmosa cinerea (Tables 2-8 and A-5). No
individuals of other species, neither rodent nor marsupial, were
recaptured as frequently as individuals of this species. In the native
forested habitat, individuals of M. cinerea were recaptured on the
average 3.1 times, while in the eucalypt forest, individuals of this
species were recaptured on the average 7.7 times.
Akodon cursor was the only rodent that had a relatively high
number of captures and recapture rate (Tables A-5 and A-6). This
species had a recapture rate of 1.6 and 3.0 in the eucalypt and wet
meadow habitats, respectively.
Trapping success
Table 2-9 presents the trapping success by habitat type.
Trapping success was calculated for small mammals in the native
forested habitat. The platform trapping data was excluded. It must
be kept in mind that the sampling effort in each general habitat
category was different, however, comparisons of trapping success are


40
PERCENT SUCCESS
MONTHS
EUCALYPT -+- WET MEADOW NATIVE FOREST
Figure 2-4. Comparison of trapping success of small mammals in
three habitat types: eucalypt forest, wet meadow, and native
forest in the Rio Doce State Forestry Park, Minas Gerais, Brazil.


41
the result of the number of captures relative to the number of
trapping opportunities or nights. Overall, the wet meadow habitat
yielded the highest trapping success (18.8%) while the homogeneous
eucalypt habitat generated the lowest trapping success (0.02%).
Figure 2-4 compares the progression of the trapping successes of
the native forested habitat (without the platform data), the wet
meadow habitat and the eucalypt forest with subcanopy habitat.
Although these three habitats have unequal sampling effort and
trapping design, these habitats were sampled for the period of one
year and show important temporal trends. From the overall gross
comparison portrayed in Figure 2-4 and the percent success presented
in Table 2-9, in contrast to the wet meadow habitat, it appears that
the forested habitats, both native and exotic, were quite similar in
overall percent success and monthly trapping success. The wet
meadow trapping success fluctuated greatly throughout time, from a
high approaching 45% in March and April, to a crash lower than 10%
in May, October, and November.
I plotted the total captures, minimum known alive and first
captures through time for each of the three habitats. Figure 2-5
shows the capture curves for the native forested sites. Trapping
success was relatively low and stable from November through April,
with a noticeable increase in June, July and the early part of August.
This peak in trapping success dropped back to the levels observed
prior to the increase. Figures 2-6 and 2-7 compare the trapping
success of the eucalypt forest and the wet meadow habitat,
respectively. The highest number of captures at any one time in the


NUMBER OF CAPTURES
Figure 2-5. Small mammal capture curves for all native forested
sites in Rio Doce State Forestry Park, Hinas Gerais, Brazil.
Capture curves include total captures, minimum known alive (MKA),
and first captures.


43
RD/E EUCALYPT Sill
NUMBER OF CAPTURES
MONTHS
TOTAL CAPTURES MKA CAPTURES FIRST CAPTURES
Figure 2-6. Small mammal capture curves for eucalypt forest 1n
the Rio Doce State Forestry Park, Minas Gerais, Brazil. Capture
curves include total captures, minimum known alive (MKA), and
first captures.


44
NUMBER OF CAPTURES
MONTHS
TOTAL CAPTURES MKA CAPTURES FIRST CAPTURES
Figure 2-7. Small mammal capture curves for wet meadow habitat
in the Rio Doce State Forestry Park, Minas Gerais, Brazil.
Capture curves include total captures, minimum known alive (MKA),
and first captures.


45
Table 2-10.
T rapping
success by trap type for all species in all
habitat types.
, Trap
types are arranged according
to trapping
location: terrestrial or arboreal. Trap types are as follows: 1ST=
small terrestrial Sherman live trap; 3MT=
medium sized
terrestrial
live trap; 5LT= large
terrestrial live
trap; 2SA= small arboreal
Sherman live trap; 4MA
arboreal platform trap.
= medium sized
arboreal live
trap; 6PA=
NO.
NO.
PERCENT
TRAP TYPE
CAPTURES
TRAP NIGHTS
SUCCESS
TERRESTRIAL
1ST
370
4080
9.1
3MT
553
18000
3.1
5LT
42
5760
0.7
965
30480
3.2
ARBOREAL
2SA
48
2640
1.8
4MA
228
8640
2.6
6 PA
66
1050
6.3


46
wet meadow habitat was double the highest in the eucalypt forest.
However, the trends were similar. Both habitats showed two
pronounced peaks in their respective capture curves that corresponded
to the same months throughout the year. There was a peak in
February, March, and April followed by a crash, and another peak in
June, July, and August followed by another crash.
T.rap Types
Overall, trapping success was higher with terrestrial traps than
with arboreal ones (Table 2-10). Caution should be used in these
comparisons as the number of trap nights are unequal; almost three
times the number of terrestrial trap nights as the number of arboreal
ones. However, I feel that some degree of comparison can be drawn
from this analysis based upon the number of captures relative to the
number of trap nights.
Trapping success by trap type varied considerably (Table 2-10).
Small terrestrial Sherman live traps were the most successful (9.1%),
medium terrestrial traps represented the trap with the greatest
number of captures and trapping opportunities, and large terrestrial
live traps were relatively unproductive.
Arboreal trap type success varied. Small arboreal Shermans
were the least productive arboreal trap type with only 1.8% success.
The arboreal platform traps had a trapping success of 6.3%.
Table 2-11 reports the number of total captures of each species
and the number of captures per trap type. Odd numbered traps
represent terrestrial traps and even numbered ones represent arboreal


47
Table 2-11. Trap response by species across all habitat types.
Trap types are explained in Table 2-10. Species abbreviations are
explained in Table 2-4.
SPECIES
TRAP
TYPES
TOTAL
1ST
2SA
3MT
4MA
5LT
6AP
DM
42
0
0
24
2
15
1
MN
158
0
0
129
3
26
0
MI
171
13
14
106
38
0
0
MC
361
2
29
145
150
0
35
CP
51
0
0
9
13
0
29
MM
1
0
0
0
1
0
0
NS
19
4
0
13
1
1
0
RM
7
0
0
3
3
0
1
AC
392
312
2
78
0
0
0
OT
21
1
3
9
8
0
0
OC
21
8
0
11
2
0
0
OR
10
7
0
3
0
0
0
AR
1
0
0
0
1
0
0
OS
44
18
0
20
6
0
0
CL
2
2
0
0
0
0
0
ON
4
3
0
1
0
0
0
EG
2
0
0
2
0
0
0
TOTALS
1307
370
48
553
228
42
66
PERCENTAGES
28.3
3.7
42.3
17.4
3.2
5.0


48
traps. In general, the number of captures per species in terrestrial
and arboreal traps represented the spatial adaptation for each species.
For marsupials, Didelphis marsupial is and Metachi rus nudicaudatus
were captured principally in terrestrial medium live traps (3MT),
while Marmosa incana was captured in all trap types except for the
large terrestrial and arboreal platform traps. Marmosa cinerea was
captured in all trap types except the large terrestrial traps.
Caluromys philander was trapped principally in arboreal medium live
traps and arboreal platform traps. The sample size for rodents was
too small to allow for a clear trapping trend. Akodon cursor is the
clear exception. The small terrestrial Sherman live trap was most
effective for this species. This was mostly a consequence of the
approximately 80% of all Akodon captures made in the wet meadow
habitat (Table 2-10). Orvzomvs subflavus was trapped principally in
terrestrial small shermans and medium live traps.
Discussion
The trapping success realized in this study for neotropical humid
forests falls within the range of observed success rates (Table 2-12).
However, one striking difference between this and other neotropical
small mammal studies was the high number of marsupial captures
relative to rodent captures (Table 2-12). All reported studies show
rodent biases, and usually high captures of rodents relative to
marsupials. Only Emmons (1984) reported a marsupial to rodent
capture ratio approaching equality (Table 2-12). Fonseca (pers.
comm.) reported marsupial biased trapping results from a variety of


49
Table 2-12. Percent capture by taxonomic group, trapping success
and number of trap nights by habitat type for this study compared
to other Neotropical field studies. % M= percent of total
marsupial captures; % R= percent of total rodent captures; % T.S.=
percent trapping success; # T.N.= mumber of trap nights.
STUDY
SITE
% M
% R
% T.S.
# T.N.
NATIVE FOREST
THIS STUDY
BRAZIL
83.2
16.8
2.3
30,960
DIETZ ET AL., 1975
BRAZIL
9.3
90.7


CARVALHO, 1965
BRAZIL
0.3
99.7
3.6
10,080
EMMONS, 1984
PERU
48.0
52.0
6.9
2,987
EMMONS, 1984
PERU


7.0
4,390
EMMONS, 1984
BRAZIL


0.8
434
DIAS, 1982
BRAZIL
2.3
97.7


NITIKMAN & MARES,
1987
BRAZIL
30.7
69.3
6.0
12,170
LAEMMENT ET AL.,
1946
BRAZIL
31.0
69.0
10.0
30,000
AUGUST, 1984
VENEZUELA
25.0
75.0
0.9
30,269
DAVIS, 1945
BRAZIL
17.0
83.0


FLEMING, 1972
1974
PANAMA
19.0
81.0
16.0
24,732
OCONNELL, 1979
VENEZUELA
12.0
88.0


WET MEADOW/SAVANNA/PANTANAL
THIS STUDY
BRAZIL
2.0
98.0
18.8
1,980
AUGUST, 1984
VENEZUELA
0.0
100.0
1.9
3,660
AUGUST, 1984
VENEZUELA
25.0
75.0
0.1
4,400
LACHER & ALHO,
IN PRESS
BRAZIL
0.0
100.0
4.2
3,582
BORCHERT & HANSON,
1983
BRAZIL
0.0
100.0
3.5
4,173
OCONNELL, 1981
VENEZUELA
10.0
90.0


HOMOGENEOUS EUCALYPT
FOREST
DIETZ ET AL., 1975
BRAZIL
0.0
100.0
THIS STUDY
BRAZIL
0.0
100.0
0.2
500
EUCALYPT FOREST WITH
I NATIVE SUBCANOPY
THIS STUDY
BRAZIL
83.0
17.0
2.6
6,000


50
native forest sites in eastern Minas Gerais. Avila-Pires (1978)
captured 245 rodents and 40 marsupials from the R1o Doce Park.
Gastal (1982) reported that 5 species of marsupials were captured in
the Park but gave no comparative data for rodents. Dias (1982)
trapped more rodents than marsupials in the Rio Doce Valley in
Minas Gerais.
Hunsaker (1977) stated that marsupials require considerable
effort to trap. Perhaps one explanation for the observed high
marsupial/rodent capture ratio could be due to the habitat type found
in the Park. There is very little primary habitat in the Park relative
to secondary habitat (Table 2-1). Most of the forest habitat in the
Park has been altered by fire in the recent past (Lopes, 1982). The
primary forest plots that I sampled yielded the lowest species
richness and absolute captures of didelphid marsupials relative to the
other secondary forested habitats (J. Stallings, in prep.). Charles-
Dominique (1983) suggested that didelphid marsupials can reach high
local densities in areas of abundant food resources. He postulated
that these species are r-strategists and are adapted to the "unstable
environment of secondary forests." In Panama, Didelphis marsupial is
tended to occur at higher densities in primary forest, while
Caluromvs and Philander were found at higher densities in secondary
forest than primary forest (Fleming, 1972).
Marsupials were recaptured with great frequency in this study,
especially Metachi rus nudicaudatus. Marmosa incana and Marmosa
cinerea. My recapture data on marsupials agree with data reported
by Fleming (1972, 1973), August (1984) and to some degree with that


51
found by OConnell (1979). My data do not agree with Hunsaker
(1977) who stated that didelphid marsupials are difficult to recapture.
For example, individuals of Marmosa cinerea were captured on the
average 7.7 times in the eucalypt forest with native subcanopy
habitat. This high recapture rate could be explained by the fact that
this habitat in effect was surrounded by habitat insuitable for
arboreal species. In essence, this habitat was a forested island. On
one side there was a monoculture of Eucalyptus saligna with no
subcanopy, on another a marsh turned into a rice field, and the other
two sides of the forest were bounded by pasture. Arboreal species
should spend more time in arborescent vegetation than on the ground.
These species should be more hesitant to disperse than terrestrial
ones.
Another obvious difference between this study and other
inventories conducted in neotropical native forests was the absence of
echymid rodents. Species of the genus Proechimvs are the most
widespread taxa of the family Echymidae in the neotropics
(Hershkovitz, 1969). These forest species are terrestrial and usually
appear on species lists from forest inventories. In Panama,
Proechimys was a common forest capture (Glanz, 1982; Eisenberg and
Thorington, 1973). Handley (1976) reported Proechimvs as a common
species in Venezuela. Emmons (1984) and Terborgh et al. (1986)
reported captures of Proechimvs from forested sites in Peru and
Ecuador. There are several reports of Proechimvs captures in
Brazilian tropical moist forests. Laemmert et al. (1946), Emmons
(1984), Carvalho (1965), Miles et al. (1981) and Malcolm (pers. comm.)


52
reported Proechimvs in their inventories from the Brazilian Amazon.
In the Atlantic Forest, Davis (1945) and Fonseca (pers. comm.)
reported two species of Proechimvs from their studies in the states
of Rio de Janeiro and Minas Gerais, respectively. Dias (1982) trapped
one species of Proechimvs in 3 study areas in the Rio Doce Valley.
I did not capture one individual of Proechimvs from the Park in
approximately 35,000 trap nights from 1985-86, nor from an additional
30,000 trap nights from 1986-87 (J. Stallings, in prep.). One
explanation could be the presence of predators in the Park.
Eisenberg (1980) speculated that the abundance of rodents in some
neotropical sites and the paucity of rodents in other sites could be
the result of the presence or absence of top predators. Hershkovitz
(1969) stated that species of the genus Proechimvs "are the basic
source of protein for lowland predators in the Brazilian subregion."
The felid community in the Park is intact. All of the felids have
been observed by field workers in the recent past. I saw spoor from
jaguar, puma, ocelot, Geoffroys cat, and jaguarundi.
In the wet meadow habitat my findings were consistent with the
literature (Table 2-12). In every other study, rodent captures were
higher than marsupial captures. In this study, Akodon cursor was the
dominant species in terms of absolute numbers and captures.
OConnell (1981) reported that Zvqodontomvs was the dominant rodent
in grass habitat in Venezuela and represented 85% of the total rodent
captures. One difference between this study and others was the
observed high trapping success. As can be observed from Table 2-4,


53
Akodon cursor made up 85% of the captures and was responsible for
the high trapping success.
The eucalypt forest with native subcanopy habitat yielded
surprising results. I did not expect to find many small mammals in
this habitat because of "plantation effects." However, I captured 7
species of small mammals, most of these being marsupials. In fact,
the marsupial/rodent capture ratio was similar to that observed in the
native forested habitat (Table 2-12). I could not find any studies in
the literature to compare to the results from this habitat. Perhaps
the study that sampled homogeneous eucalypt forest would be the
most appropriate (Dietz et al., 1975). Dietz et al. (1975) captured
two species of terrestrial rodents, Orvzomvs nigripes and Akodon
cursor, in the homogeneous plantations with grass/bamboo
undergrowth. The authors caught a total of 5 species, only one of
which was not strictly terrestrial, in the two native forested habitats.
The plantation habitats in this and my study are similar in that they
both were eucalypt plantations of similar age and that the terrestrial
substrate of both was covered by grass. The major difference was
the native species subcanopy. I captured a relatively high number of
terrestrial rodents and marsupials and a high number of arboreal
marsupials. I did not capture any arboreal rodents. The native
species subcanopy could be considered a secondary forest sere, if the
emergent eucalypt stratum is ignored. Following Charles-Dominiques
(1983) hypothesis of increased didelphid marsupial abundances in
secondary habitat, it is not surprising that I captured marsupials at a
rate consistent with the native forested habitat.


54
The temporal capture results suggested that there were two
systems operating in the Park. The trapping data showed a
pronounced peak in the total number of captures, mka and first
captures for the native forested habitat during the cool, dry winter.
Davis (1945) reported a similar trend in the trapping results and
suggested that this trend was the result of more younger individuals
present 1n the trapping pool or because of a paucity of natural food
items during this time of year. My data do not support the
hypothesis that more younger individuals explain the pronounced
increase; rather 1t appears that food resource paucity results in the
increase (Chapter III).
The eucalypt and wet meadow habitats showed similar trapping
trends with a peak in the cool, dry winter, and a peak in late
summer. These two habitats might have yielded similar capture curves
because of the grass substrate. Akodon cursor was an important
component of both habitats and is an insectivore/omnivore that is
reported to use a high proportion of grass and grass seed in its diet
(Nowak and Paradiso, 1983). Harmosa incana and Metachi rus
nudicaudatus are insectivore/omnivores and frugivore/omnivores,
respectively, and perhaps track insect availability in grass substrate.
The grass species did not produce seeds until late May. Thus,
perhaps the peak observed in February, March and April can
be explained by the lack of food for both rodents and marsupials.
The second peak, which occurred in June, July, and August, could
also be explained in terms of a decrease in food availability. Insect
and fruit availability are usually low during the hibernal period in


55
seasonal neotropical forests (e.g., Janzen and Schonener, 1968). The
marsupial species rely heavily on these food resources. The results
of a preliminary stomach content analysis on Akodon. suggest that
insects are important items in this species diet (Stallings, in prep.).
Graminoids in this habitat were dry and seeds were not as readily
available as they were during April and May.
The trap type data analysis revealed that terrestrial small
Sherman Uve traps were very productive in the wet meadow habitat
but yielded relatively few captures in the forested habitats. The
same results were obtained for arboreal small Shermans in the
forested habitats. Large terrestrial Uve traps were unproductive 1n
the forested habitats. The most productive trap types were the
medium sized terrestrial and arboreal traps and the arboreal platform
traps. Some individuals of species that are considered terrestrial
were captured in arboreal traps. This can be explained by some low
arboreal traps that were connected to the ground by either a vine or
log.
The use of the arboreal platform traps did not allow me to trap
additional species that were not already trapped using the terrestrial
and low arboreal traps. However, I was able to increase the
frequency of capture of the highly arboreal marsupial Caluromvs.
Malcolm (per. comm.) obtained similar results with platform traps in
Manaus. Perhaps the first use of arboreal platforms for trapping
small mammals were in the studies described by Davis (1945) and
Laemmert et al. (1946). Unfortunately, I could not determine the
success of these traps nor the species captured from these studies.


56
In total, I logged 49 captures of this Caluromvs in both the eucalypt
with native species subcanopy and the native forested habitats. In
the latter habitat, I only trapped this species 5 times in the
terrestrial and low arboreal traps. I trapped 29 Caluromvs in the
native forested habitat by using the arboreal platforms. I would have
underestimated the presence of this species had I not used the
platform traps. Marmosa cinerea was trapped also with relative high
frequency in this trap type.
I was surprised to find such a high trapping frequency of
Caluromvs in the eucalypt forest with native species subcanopy.
Although this species can be quite common in native species forested
habitats, I found it unusual that this highly arboreal frugivore would
be inhabiting an exotic monoculture plantation. There obviously was
sufficient food resources available from the native species subcanopy.
The fact that this species was captured in terrestrial and low
arboreal traps, without using platforms, suggests that this species was
using the subcanopy.
The results of this small mammal inventory suggest that
marsupials play an important role in the community structure of small
mammals in one of the largest remaining native tracts of Atlantic
forest in Brazil. Wet meadow habitat in this region is speciose in
rodents, and perhaps dominated by one or two species. Eucalypt
forests with native species subcanopy can play an Important role in
the conservation of small mammal communities in a region greatly
altered by monocultura! plantations.


CHAPTER III
TEMPORAL VARIATION IN TRAPPING SUCCESS OF
DIDELPHID MARSUPIALS IN AN EASTERN BRAZILIAN PARK
Introduction
Tropical rainforests are classified as stable and evergreen, with
little seasonal change in the flora and fauna (Richards, 1952; Smith,
1974). However, tropical seasonal and semideciduous forests exhibit
seasonal changes 1n floral phenology (e.g., Augspurger, 1982; Foster,
1982; Garwood, 1982) and consequently variation in faunal populations
and activities (e.g., Howe, 1982; Worthington, 1982; Smythe et al.,
1982; Glanz et al., 1982). These changes have been related to
variation in the abiotic environment, such as changes in photoperiod,
precipitation, and temperature (Sinclair and Norton-Griffiths, 1979;
Leigh et al., 1982).
Tropical forests that exhibit a pronounced dry season seem to
have a peak of fruiting at the onset of the wet season immediately
following the dry season (Foster, 1980). While the reasons for this
peak in the fruiting period are not clear, the result is that fruit is
more abundant at certain times of the year. This Increase in the
availability of fruit drastically affects the animals that use these
resources for food.
Population fluctuations interpreted by trapping results of small
mammals must be carefully analyzed to determine if the population is
57


58
actually fluctuating or if animals are responding to trap bait during
periods when food is available (Hunsaker, 1977). Previous research
on didelphid marsupials in the neotropics (Davis, 1945; OConnell,
1979; Fleming, 1972, 1973; August, 1984) have found that trapping
success of marsupials increased during the hibernal and prevernal
months. This increase in trapping success has been attributed to the
paucity of food resources during this period. Another interpretation
of the results could be linked to reproduction, such as greater
movements of males or a large number of immatures. These might be
related to the initiation or the cessation of reproductive activity.
One objective of this paper is to test the null hypothesis that
trapping success of marsupials does not vary with season in the
Atlantic Forest of eastern Brazil. Another objective is to test the
null hypothesis that any peak in trapping success was a consequence
of an increase in the population or reproductive activities.
Materials and Methods
Small mammals were trapped from October, 1985, through
September, 1986, in the Rio Doce State Forestry Park, Minas Gerais
state, Brazil. The Park lies between the coordinates 19 4818" and 19
2924 south latitude and 42 3830" and 42 2818" west longitude. The
climate of the Park is classified as tropical humid with a rainy season
between the months of November and February and drought during
the months of June, July, and August. Mean annual temperature is
approximately 22 C, with mean minimum monthly temperatures
fluctuating greatly throughout the year (Figure 3-1). Both the mean


TEMPERATURE (C )
SUMMER SEASON MONTHS WINTER SEASON
MEAN MAXIMUM TEMP * MEAN MINIMUM TEMP
DATA FROM 1954 1974
Figure 3-1. Temperature graph showing both minimum and maximum
mean temperatures per month in the Rio Doce State Forestry Park,
Hinas Gerais, Brazil. Winter and summer seasons are indicated
along the X axis.


60
maximum and minimum monthly temperatures reach their respective
lows during the drought season.
For the purposes of this paper, I divided the year into two
seasonal periods. This division is based on the monthly mean
minimum temperature. If the mean monthly minimum temperature was
above 17 C. the month was classified as summer, otherwise it was
considered the winter period. A comparison of my seasonal
classifications and those of Davis (1945) shows thermal and intuitive
consistencies (Figure 3-1). My winter season corresponds to the
period from April September and the summer season from October -
March. My classification corresponds to those of Davis in the
following manner: my winter season includes the autumnal, hibernal
and prevernal seasons, while my summer season includes the vernal,
aestival and sertina! seasons.
The phytogeographical domain of the Park has been classified by
several authors. For example, according to Ab Saber (1977, in
Gilhaus, 1986), the Park occurs within the Tropical Atlantic Domain.
Rizzinis (1963, in Gilhaus, 1986) classification defines the Parks
domain as the Atlantic Province, Austro-Oriental sub-province,
Cordilheira sector. Gilhuis (1986) correctly pointed out that both
classification schemes locate the Park in a transitional zone bordering
the more humid Tropical Atlantic Domain and the drier Cerrados
Domain. I found tree species from both domains to be present in the
Rio Doce Park. Trees are more deciduous than would be expected in
Tropical Atlantic Domain forest. Most of the emergent tree species
and some others in the upper strata lose their leaves during the cool


61
and dry hibernal and prevernal seasons (Gilhuis, 1986; Stallings, pers.
obs.). The forest of the park is classified as Tropical Semideciduous
by Alonso (1977, in Gilhaus, 1986), Tropical Broadleaved by Azevedo
(1969, in Gilhaus, 1986) and Tropical Pluvial Seasonal by Lima (1966,
in Gilhaus, 1986).
Small mammals were live-trapped at 5 forested sites within the
Park boundary. Additional trapping was conducted in and around the
Park and the methodology is described in Chapter II. For the
purposes of testing the above mentioned hypotheses, I recorded the
following observations on each captured individual. The initial
capture of each individual was a first capture and a numbered ear tag
was placed on the left ear of all animals to identify individuals that
were captured previously. Total captures were first captures plus
subsequent captures of the same individual. The minimum known
alive (MKA) were the number of individuals captured on the first
occasion and those individuals recaptured only on one occasion during
each trapping period. Sex, age and reproductive information was also
recorded.
The probability of capturing a small mammal is affected by 3
factors: 1) temporal effects, 2) behavorial effects, and 3)
heterogeneity effects (White et al., 1982). The effects of
heterogeneity (e.g., number and placement of traps in an animals
home range or variation due to social dominance) vary among animals,
but capture probabilities for each animal remain constant per
occasion. In contrast, temporal (e.g., variation over time because of
environmental conditions or trapping effort) and behavior (e.g., trap


62
avoidance or fascination) effects can alter capture probabilities after
the initial capture of an Individual.
First capture data are more affected by heterogeneity and
temporal effects than behavioral effects (White et al., 1982). An
increase in the number of juveniles in first capture data through time
suggests that the population is increasing. However, if young
individuals do not account for an increase in first capture data, then
an alternative explanation 1s that the increase reflects individuals
present in the population that were not previously captured.
Total capture and minimum known alive data are influenced by
temporal, behavioral, and heterogeneity effects. Total capture data
are the result of all captures per occasion or through time, inclusive
of first captures, successive recaptures, and recapture of individuals
separated by time. These data should not be interpreted as a good
estimator of population abundance because they are strongly affected
by behavioral and temporal factors. An increase in recapture data
through time can be interpreted as a change in the capture
probabilities of marked individuals due to trap fascination, rather
than trap shyness which would reduce capture probabilities.
The minimum known alive data are the best population estimator
of the three. These data incorporate first captures and only the first
recapture of an individual captured in a previous occasion for each
trapping period. This estimator, in effect, uses the number of new
captures plus survivorship of animals captured previously to obtain an
estimate of abundance.


63
I only used marsupial capture data to test the null hypotheses
that trapping success does not vary by season and that any expected
increase is due to reproduction. Marsupials comprised 83% of the
total small mammal captures and 80* of the first captures (Chapter
II). Marsupial species included in the analysis were Didelphis
marsupial is (azarae). Metachi rus nudicaudatus. Marmosa incana.
Marmosa cinerea. and Caluromvs philander. Fruit and insects are
important compontents of this groups diet (Chapter II).
I used correlation analysis and simple regression to determine if
there was any association between trapping success and mean
minimum monthly temperature. I quantified trapping success of first
captures, minimum known alive and total captures for each season.
Chi-square tests for goodness of fit were then used to test for
significant deviation from expected capture frequencies between
seasons.
I looked at (1) an increase in the number of juveniles during
the winter season, (2) an increase in the number of males between
seasons, (3) a male biased sex-ratio during the winter season, and (4)
an increase in the number of lactating/pregnant females during the
winter season to test the null hypothesis of an expected increase due
to reproduction. For this analysis I used Chi-square tests goodness
of fit to test for significant deviation from the above expected
results.
Individuals were placed into either juvenile or adult subjective
age classes. Initially, the age class placement was based on the
overall morphological gestalt of each individual per species. I then


64
used the recorded body measurements (Chapter II) to determine the
threshold between the two age classes and I compared field
determined ages to ages derived from body measurements. Body mass
and hind foot measurements were relied on heavily to determine age
class.
Age class categories of the 5 species of marsupials are presented
in Table 3-1 and are taken from Chapter III. For Didelphis
marsupialis. juvenile and adult age categories were based on the hind
foot size. This species can attain a body mass of 2 kg. The hind
foot would grow slower in comparison to smaller bodied species and
body mass would provide a better indicator of age. Animals with a
hind foot of 51 mm or smaller were classified as juveniles (average
weight 417 gm). Lengths greater than 51 mm (average weight 1049
gm) were classified as adults. I used body mass to determine the
threshold for Metachi rus nudicaudatus. Marmosa incana, and Marmosa
cincera. These species are relatively small and foot size reaches its
maximum at an early age. Body mass provided the better estimator
because mass increased as the individuals grew older. Metachlrus
nudicaudatus was placed into age categories based upon a threshold
weight of 90 gms. Age class placement was determined by a weight
of 35 gm and 50 gm for Marmosa incana and M. cinerea. respectively.
All Caluromys philander that were captured were considered adults.
The number of individuals of each sex were sorted for first,
MKA, and total capture data between and within seasons.
Comparisons were made for sex biased trapping success between


65
Table 3-1. Criteria used to place marsupials into either juvenile
or adult age categories. Individuals per species were classified
as juveniles if designated measurements were less than cut-of
measurements used in table. Species were not sexually dimorphic
based on body measurements listed.
SPECIES
BODY
MEASUREMENT USED
CUT-OFF
MEASUREMENT
DidelDhis marsuDialis
HIND FOOT
51 MM
Metachi rus nudicaudatus
MASS
90 G
Marmosa incana
MASS
35 G
Marmosa cinerea
MASS
50 G


66
seasons for all capture data. In addition, comparisons were made for
sex biased trapping success within seasons.
Female marsupials were examined to determine if they were
lactating and for the presence of young attached to the teat field.
Females with young attached were considered pregnant. Frequencies
of lactating/pregnant females were compared on a seasonal basis.
Results
There was a significant negative association observed between
mean monthly temperature and trapping success through time. This
association is significant only if we look at the number of total
captures (f=28.444, p < .0003) (Figure 3-2) or the minimum known
alive (f=9.939, p < .007) (Figure 3-3). First capture data did not
show a significant association (Figure 3-4).
The first capture data (0.05 0.001) and total captures (p< 0.001) of marsupials were greater than
expected in the winter season than in the summer season (Table 3-2).
Juveniles were observed throughout the year. Juveniles were
represented from four of the five species of marsupials (Table 3-3).
There was no significant difference in the minimum known alive or
first captures of juveniles between seasons. A significant difference
(p < .001) was observed in the total captures of juveniles between
seasons. More total captures of juveniles were observed in the
winter season than expected.
The number of juveniles captured per species between seasons
was fairly constant (Table 3-3) from minimum known alive and first


67
NUMBER OF CAPTURES
Figure 3-2. Relationship between temperature and total captures
of didelphid marsupials in the Rio Doce State Forestry Park,
Minas Gerais, Brazil.


68
Figure 3-3. Relationship between temperature and minimum known
alive captures (MKA) of didelphid marsupials in the Rio Doce
State Forestry Park, Minas Gerais, Brazil.


69
NUMBER OF CAPTURES
0 5 10 15 20
TEMPERATURE (C )
Figure 3-4. Relationship between temperature and first captures of
didelphid marsupials in the R1o Doce State Forestry Park, Minas
Gerais, Brazil.


70
Table 3-2. Comparison of the first captures, minimum known alive
(MKA) and total captures of five species of didelphid marsupials
between seasons in the Rio Doce State Forestry Park, Minas Gerais,
Brazil. Significant at 0.05-0.01. ** Significant at 0.01
0.001.
SEASONS
WINTER
SUMMER
CHI-SQUARE
FIRST CAPTURES
206
151
4.23*
MKA CAPTURES
276
157
16.35**
TOTAL CAPTURES
392
190
35.05**


71
Table 3-3. The number of first captures (FIRST), minimum known
alive (MKA) and total captures (TOTAL) of juvenile didelphid
marsupials per season. N.S= Non significant.
SUMMER SEASON WINTER SEASON
FIRST MKA TOTAL FIRST MKA TOTAL
SPECIES
D. marsupial is
M. nudicaudatus
M. incana
M. cinerea
TOTALS
FIRST CAPTURES X2
MKA CAPTURES X2
TOTAL CAPTURES X2
3
8
10
7
5
14
15
17
26
8
10
19
33
40
69
: 0.00,
N.S.
: 0.55,
N.S.
14.38,
P < 0.
001
6
6
7
5
5
21
14
18
35
8
21
85
33
50
148


72
capture data. The total capture data per species indicated that there
was a recapture bias during the winter season, although the
difference was not significant. Figure 3-5 shows that first captures
of adults, not juveniles, clearly represented the majority of captures
during May, June, and July.
The number of captures of each sex between seasons is
presented in Table 3-4. There were no significant differences
between the number of males captured between seasons or the number
of females captured between seasons using the first capture data.
Minimum known alive and total captures data showed significant
differences in the expected number of males or females between
seasons. More males and more females were captured during the
winter season than during the summer season, with a higher
proportion of males being captured during the winter season. For the
first capture data, minimum known alive and total capture data there
were no significant differences in the male/female sex ratio within
each season (Table 3-5).
More lactating females, whether carrying young or not, were
captured during the summer season than the winter season (Table
3-6). However, this difference, although approaching significance,
was not statistically significant (x2 =2.89; .10 < p <.05). One
striking observation was the obvious decrease in the occurrence of
lactating/pregnant females during the months of June, July and
August (Figure 3-6). This decrease is also observed when comparing
number of juveniles and lactating females per species through time
(Figures 3-7, 3-8, and 3-9).


>
73
OF ADULTS AND YOUNG BY MONTH
NUMBER OF CAPTURES
MONTHS
cm NO. YOUNG GO NO. ADULT
Figure 3-5. Comparison of first captures of adult and young
didelphid marsupials per month in the Rio Doce State Forestry Park,
Minas Gerais, Brazil.


74
Table 3-4. Between season comparisons of the number of first
captures (FIRST), minimum known alive (MKA) and total captures
(TOTAL) of didelphid marsupials per sex. Significant at 0.05-
0.01. ** Significant at 0.01-0.001.
BETWEEN SEASON COMPARISON
SUMMER SEASON WINTER SEASON
X2
NUMBER OF MALES
FIRST CAPTURES
62
83
1.52
MKA CAPTURES
78
152
*
*
o
o>
f
TOTAL CAPTURES
97
215
22.30**
NUMBER OF FEMALES
FIRST CAPTURES
66
70
0.05
MKA CAPTURES
79
124
4.98*
TOTAL CAPTURES
92
174
12.63**


75
Table 3-5. Within season comparison of the number of first
captures, minimum known alive captures (MKA) and total captures of
didelphid marsupials per sex. N.S.= Non significant. Significance
level set at p < 0.05.
SEX
X2
SUMMER SEASON
MALES
FEMALES
FIRST CAPTURES
62
66
0.06 N.S.
MKA CAPTURES
78
79
0.003 N.S.
TOTAL CAPTURES
97
92
0.06 N.S.
WINTER SEASON
MALES
FEMALES
FIRST CAPTURES
83
70
0.55 N.S.
MKA CAPTURES
152
124
1.42 N.S.
TOTAL CAPTURES
215
174
2.16 N.S.


76
Table 3-6. The frequency
marsupials trapped per season in
Minas Gerais, Brazil. N.S.= Non
of lactating/pregnant didelphid
the Rio Doce State Forestry Park,
Significant.
SPECIES
SEASONS
SUMMER WINTER
D. marsupial is 4
M. nudicaudatus 11
M. incana 9
M. cinerea 12
Caluromvs philander 1
1
12
0
3
3
TOTALS
37
19


CO h- CD lO ^ CO CM
77
MONTHS
EB Dldelphls H:.:::.l Metachirua Ml m. incana
EE¡3 M. clnerea 1... I Caluromys
Figure 3-6. Comparison of lactating females of five species of
didelphid marsupials per month in the Rio Doce State Forestry Park,
Minas Gerais, Brazil.


78
NUMBER
M. nudicaudatua
Figure 3-7. Comparison of the number of captures of young and
lactating females of the didelphid marsupial Metachi rus
nudicaudatus in the Rio Doce State Forestry, Minas Gerais, Brazil.


79
NUMBER
M. incana
Figure 3-8. Comparison of the number of captures of young and
lactating females of the didelphid marsupial Marmosa incana in the
Rio Doce State Forestry Park, Minas Gerais, Brazil.


80
CY
AND LACTATING FEMALE-
"2)
NUMBER
MONTHS
CZI YOUNG LACTATING FEMALES
M. cinerea
Figure 3-9. Comparison of the number of captures of young and
lactating females of the didelphid marsupial Marmosa cinerea in the
Rio Doce State Forestry Park, Minas Gerais, Brazil.


81
Discussion
There is obviously a connection between animal population
numbers and the availability of food resources (Leigh et al; 1982).
An intuitive model is that population increases should track and peak
with maximum production of food resources. By timing reproduction
prior to the peak of available food resources, individuals maximize
their opportunities for the survival of their offspring.
There was a clear peak in the trapping success of marsupials
during the winter months in the Rio Doce State Forestry Park.
There was a significant negative association between the number of
total captures or minimum known alive each month and mean
minimum monthly temperature. However, the first capture data did
not support these previous findings. These data suggest that the
recapture of individuals plays an important role in the trapping
success numbers and that the numbers of new individuals did not
increase during the winter season.
August (1984) found an increase in the capture success of
Marmosa robinsoni during the dry season in Venezuela, while
OConnell (1979) reported an increase in the minimum known alive for
Marmosa robinsoni. M. fuscata. and Didelphis marsupial is during the
same period in Venezuela. Fleming (1972) observed an increase in
the number of captures of three species of marsupials during the dry
season in Panama. Davis (1945) reported more captures and
recaptures of two species of didelphid marsupials, Didelphis


82
marsupialis and Caluromvs philander, during the dry, cold season in
Teresopolis, Brazil.
Research on neotropical marsupial mating behavior suggests that
typically males are more active prior to and during the reproductive
or mating season (Atramentowicz, 1982; Hunsaker, 1977). This may be
a consequence of the polygynous mating systems observed for these
small mammals (Eisenberg, 1981). There was no significant difference
between seasons in the number of first captures of male marsupials.
The same results were obtained for female first captures. There were
more male and female minimum known alive captures during the
winter season than the summer season. The same results were
obtained for the total captures of male and female marsupials during
the winter season, but at a higher level of significance. These data
again suggest that there is a recapture bias in the trapping data, and
that observed and significant differences in the number of captures
and individuals of each sex between seasons is a function of being
recaptured and not related directly to reproduction. However, the
minimum known alive and total capture data show male captures in
the winter season to be more significant than females captures during
the same period. These findings could be interpreted as increased
male activity over female activity at the onset of the reproductive
season.
A direct measure of population growth is the number of young
observed at any point in time. My results indicate that during the
winter season the number of young individuals captured was not
greater than in the summer season, but more young individuals were


83
recaptured during the this season than in the summer season.
OConnell (1979) reported an increase in the number of individuals of
Didelphis marsupial is during the wet season in Venezuela. She
attributed this increase to the first captures of juveniles.
The last measure that was used to determine population growth
was that of the occurrence of lactating/pregnant females through
time. Statistically, there was no difference in the occurrence of
reproductively active females between seasons, although there was a
suggestion of more reproductively active females in the summer
season than in the winter season. OConnell (1979) found that most
of the species she studied to be sexually inactive during the dry
season.
It is clear from the previous analyses that there was a
pronounced increase 1n the number of animals known to be alive and
the overall trapping success during the hibernal and prevernal seasons
(Figure 3-10). However, the data do not support the hypothesis that
this significant increase is due to more juveniles present in the
population, nor does it result from increased male activity or more
lactating/pregnant females. I conclude that population growth of
marsupials was constant if we consider only first capture data.
Clearly, the trapping success curve was being driven by adult
individuals; individuals that were being recaptured more during a
certain period in time than at another.
I did not measure the availability of fruit or insects during the
study. Phenological studies (CETEC, 1981) in the park from 1977-
1981 found that trees were in flower and fruit throughout the year,


NO. CAPTURES
MONTHS GROUPED ACCORDING TO SEASON
SEASONS GROUPED BY MONTHS (DAVIS, 1945)
Figure 3-10. Frequency of captures of didelphid marsupials by
season in the Rio Doce State Forestry Park, Minas Gerais, Brazil.


85
with a noticeable, but not statistically significant increase from
October through December (Figure 3-11). These studies also indicated
that the months of June, July and August represented the lowest
incidence of tree species with flowers and fruits. There was also a
low incidence of tree species with flower and fruits in January.
There are no quantitative data available on the seasonality of insect
density in the park. However, studies in other Neotropical areas
report a positive association between increased insect abundance and
precipitation and temperature (Janzen and Schonener, 1968; Wolda,
1978, in August, 1984; Davis, 1945). It is possible that such a trend
occurs in the park due to the high seasonality of rainfall and
temperature.
In other areas were neotropical didephids have been studied
(Fleming, 1972; OConnell, 1979; August, 1984; Charies-Dominque,
1983), the dry season represented the period of the year when food
resources were scarce. In contrast, the three month period from
October December in the park showed more species in fruit and
flower than any other three month period. These trends are
interesting when compared to the low and peak three month trapping
success periods. The highest trapping success coincides with the
lowest number of tree species in fruit and flower and likewise the
lowest period of trapping success coincides with the maximum period
of fruit and flower production.
However, it must be stated that trapping success remained
relatively constant for the period of December and January when the
number of tree species with fruit and flower were observed to be


86
ENCY OF FRUIT
AND FLOWERS
NO. SPECIES IN FRUIT
SEASONAL PERIODS IN MONTHS
Figure 3-11. Frequency of species of trees and shrubs in fruit and
flower per seasonal period in the Rio Doce State Forestry Park,
Minas Gerais, Brazil.


87
low. This two month period has the highest mean temperature and
precipitation of the year. If insect density is high at this time, then
perhaps marsupials are able to switch to insect prey or other foods
in response to the unavailability of fruit.
I would interpret my results of a significant increase in total
trap success as a response to the lack of fruit during the subtropical
winter and not due to (1) greater movement of males (i.e., more
captures of males than females during this period) nor (2) a
significant increase 1n the number of juveniles captured nor (3)
because of more reproductively active females during this period.
Marsupials fell into traps easier during this period because they were
searching for food. This interpretation is consistent with the
findings of Atramentowicz (1982), Charles-Dominique et al. (1981),
Charles-Dominique (1983), OConnell (1979), Fleming (1972, 1973),
Davis (1945) and August (1984) and suggests that reproduction and
weaning of young marsupials occur at the end or after the dry season
when food resources are more abundant.


CHAPTER IV
SMALL MAMMAL ASSOCIATION AND MICROHABITAT
SELECTION IN AN EASTERN BRAZILIAN PARK
Introduction
There are several studies on small mammal habitat selection in
temperate zones (e.g., Dueser and Porter, 1986; Dueser and Shugart,
1978, 1979; Hallett, 1982; Rosenzweig and Winakur, 1969), but there
are relatively few small mammal habitat studies conducted in the
neotropics. August (1983, 1984) studied the relationship between
small mammals and habitat structure in Venezuela. Lacher and Alho
(in press) and Lacher et al. (1988) reported habitat selection by small
mammals in grassland habitats in southern Brazil. Nitikman and
Mares (1987) reported microhabitat preferences of small mammals in a
gallery forest in central Brazil. The purpose of this paper is to
examine the effect of habitat structure on habitat use by small
mammals in an eastern Brazilian forest, the Rio Doce State Forestry
Park.
Preliminary analysis of the habitat in this Park suggested that
there are distinct differences in the forest structure across several
forested habitats (Gilhaus, 1986). Forest fire, in the form of intense
crown fire, has played an important role, at least in the recent past,
in structuring the forest. This study tested the hypothesis that small
88


89
mammal abundances vary with habitat and that species select
microhabitats within each habitat.
Materials and Methods
This study was conducted in the Rio Doce State Forestry Park
(Rio Doce), Minas Gerais, Brazil (19 48 18 and 19 29 24 south
latitude and 42 38 30" and 42 28 18" west longitude). The Park
contains over 35,000 ha and elevation ranges from 230 to 515 m. The
mean annual precipitation for the Park was 1480 mm. from 1954 to
1974, however annual precipitation during the study amounted to 947
mm. (Figure 4-1). Mean monthly maximum and minimum temperatures
vary appreciably throughout the year (Figure 4-2).
Study Sites
There are several distinct forested and open/field habitats in the
Park (Gilhaus, 1986). Slope, soil quality and moisture, and elevation
all affect habitat type. However, forest fire has played the major
role affecting the forested habitat in this Park. The vegetation of
the Park is classified as tropical semi-deciduous and most of the
emergent tree species lose their leaves during the cool dry months.
In 1964 and 1967, major forest fires burned approximately 30% of the
forest (Lopes, 1982; Silva-Neto, 1984). Fire is important because leaf
Utter accumulates during the dry season.
During the course of this study, I surveyed small mammal
communities in 5 distinct forested sites which represent 3 habitat
types in the Park (Figure 4-3). The following is a brief description
of each study site.


90
WALTER AND LEITH
CLIMATIC DIAGRAM
WALTER AND LEITH
CLIMATIC DIAGRAM
MONTHLY RAINFALL (MM.)
MEAN TEMPERATURE (0 )
JAN FEB MAR APR MAY JUN JUL AUG SEP OOT NOV 060
MONTHS
AVERAGE ANNUAL RAINFALL U80 mm
MONTHLY RAINFALL MAA)
MEAN TEMPERATURE (O )
MONTHS
ANNUAL AVERAGE RAINFALL BOA mm
QUA FROM 1864-1874
DATA FROM mm
A.
5.
Figure 4-1. Walter and Leith climatic diagram characterizing
surplus precipitation and drought by month in the Rio Doce State
Forestry Park, Minas Gerais, Brazil.


Full Text
SMALL MAMMAL COMMUNITIES
IN AN EASTERN BRAZILIAN PARK
By
JODY R. STALLINGS
A THESIS PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN
PARTIAL FULFILLMENT OF THE REQUIREMENTS
FOR THE DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
1988

ACKNOWLEDGEMENTS
This project could not have been carried out without the help
of numerous people. Foremost, I thank John G. Robinson for
suggesting the research project in the Rio Doce State Forestry
Park. His perception and patience aided 1n all aspects of the
project and his visit to the project site in May, 1986, was a great
boost to my morale. I would also like to thank the other members
of my committee for their suggestions during the planning stage and
for helping me to interpret the data. I thank Drs. John F.
Eisenberg, Melvin E. Sunquist, Wayne R. Marion, and Nigel J. H.
Smith. John Eisenberg was a constant source of support through the
analysis and writing stages. His continued interest 1n the results
and concern for my welfare helped to carry me through difficult
periods. Kent H. Redford provided obscure and relevant
publications. He also visited the field site at the beginning of
the project and encouraged me to sample the eucalypt forests.
Funding for this project was provided from the Program for
Studies 1n Tropical Conservation at the University of Florida,
World Wildlife Fund-US, The Organization of American States, and
the Instituto Estadual de Florestas (IEF). I thank John Robinson,
Kent Redford, Russell Mittermeier, John Eisenberg, Jose Carlos

Carvalho, and Celio Valle for helping me to obtain the funds
necessary to carry out the project.
I would like to thank Jose Carlos Carvalho, president of IEF,
for allowing me to work in the R1o Doce State Forestry Park. Dr.
Celio Valle, limar Bastos Santos, and Gustavo A. B. da Fonseca were
very instrumental in helping me to obtain my scientific expedition
visa to conduct research 1n Brazil. Celio Valle and Gustavo
Fonseca were more than helpful throughout the various stages of
fieldwork and allowed me to work in the mammalogy laboratory in
Belo Horizonte. Gustavo Fonseca was more than generous 1n sharing
his computers with me. Dr. Wilson Mayrlnk and his excellent staff
treated me in a concerned and professional manner. I thank them
for saving my nose.
The professional staff at the R1o Doce State Forestry Park
contributed to the project 1n several ways. Ademlr Camara Lopes,
the park Administrator, helped us get settled into the Park. He
offered no cost housing for the duration of the project and
contributed a modest amount of the Park’s gasoline supply for my
use 1n the project. He also was able to obtain funds from IEF for
the construction of arboreal platforms and 60 wire traps.
Hermogenes Ferreira S. Neto was extremely important 1n solving
daily problems and 1n handling our correspondence. Jose Lourenco
Ladeira, the park dendrologist, identified the shrub and tree
species that occurred within each trapping post. I am very
grateful for his participation and was amazed at his knowledge of
the taxonomy.

I thank Drs. Phil Myers, Mike Carleton, Guy Musser, Al
Gardner, and James Patton for identifying the voucher specimens.
Their prompt reply enabled me to begin the analysis shortly after
returning to the states.
I relied on several workers from the R1o Doce park to
implement the field project. Trails were opened in an expert
manner by Ivanil Moreira and Waldemar Queroga dos Santos. Ivanil
later became my field assistant and proved to be an extremely
valuable asset. He never once complained because of the Incredible
numbers of ticks or because of all the small mammals that bit
chunks out of h1s fingers. I also thank Lldair Rufino for climbing
42 trees and placing the arboreal platforms.
Several students from the Departamento de Ecologia and
Zoologia participated 1n the fieldwork. I was aided greatly in the
data collection by 3 students. I thank Ludmilla Aguiar, Eduardo
Lima Sabato, and Luiz Paulo de Souza Pinto. I could not have
collected the appropriate data without their help. I also thank
Sonia Riquiera for her effort in helping us adjust to Minas Gerais,
and for participating in the initial stages of the fieldwork. She
was a pleasure to work with and always brought a positive attitude
to unpleasant working conditions.
My stay in Brazil was greatly facilitated by Gustavo and Ana
Fonseca. Their hospitality was more than generous and they always
opened their door to me and my family. Their hospitality was more
than "mineiro", it was more of true friendship. My wife and I will
iv

always be grateful to all that the Fonsecas did to help us get
adjusted in Brazil.
To my wife and best friend, Cathie, I offer my deepest
gratitude for her patience, understanding, and moral support during
the 12 month field project. It was always a pleasure to come home
from the field and be met by a smiling and positive person. I am
very appreciative for Cathie’s participation 1n the project and
wish that she could have had more freedom to pursue her own
interests.
v

TABLE OF CONTENTS
Page
ACKNOWLEDGEMENTS i1
ABSTRACT viii
CHAPTERS
I INTRODUCTION 1
General Background 1
Study Organization 3
II SMALL MAMMAL INVENTORIES IN AN EASTERN BRAZILIAN PARK . 5
Introduction 5
Study Site 7
Methods 11
Results 22
Discussion 48
III TEMPORAL VARIATION IN TRAPPING SUCCESS OF
DIDELPHID MARSUPIALS IN AN EASTERN BRAZILIAN PARK 57
Introduction 57
Materials and Methods 58
Results 66
Discussion 81
IV SMALL MAMMAL ASSOCIATIONS AND MICROHABITAT
SELECTION IN AN EASTERN BRAZILIAN PARK 88
Introduction 88
Materials and Methods 89
Results 197
Discussion 114
V FOREST FIRE AS A DETERMINANT OF SMALL MAMMAL
DIVERSITY IN A BRAZILIAN FOREST 122
Introduction 122
Materials and Methods 126
VI

Results 138
Discussion 155
VI CONCLUSIONS AND SYNTHESIS 160
APPENDIX 165
LITERATURE CITED 187
BIOGRAPHICAL SKETCH 200
vi i

Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy
SHALL MAMMAL COMMUNITIES IN AN EASTERN BRAZILIAN PARK
By
JODY R. STALLINGS
August 1988
Chairman: John G. Robinson
Major Department: Wildlife and Range Sciences
Forest Resources and Conservation
This dissertation examines small mammal communities in native
and non-native habitats in one of the largest remaining forested
tracts of land in the Brazilian Atlantic Forest. Small mammals
were live-trapped over a period of 12 months in grass, eucalypt,
and native forested habitats. Eucalypt and native forested
habitats were rich in rodent species, but marsupials were
numerically higher in terms of relative densities. Grass habitats
were rich in rodent species and were dominated by one rodent
species. Trapping success for marsupials was observed to increase
during the sharp dry season. Paucity of food resources may be the
factor responsible for the observed increase in trapping success.
The Rio Doce State Forestry Park has been subjected to frequent
crown fire in the recent past, and relatively little primary forest
remains. Marsupials may dominate in terms of abundance in this
vi i i

Park because of the large amount of secondary forest. Small mammal
species diversity was calculated from forest stands that were
burned completely to the ground, in stands that were burned in a
mosaic fashion, and in primary stands. Species diversity was found
to be higher in intermediately fire disturbed native forested
habitats in comparison to heavily disturbed and primary forested
habitats. Mosaic habitats, composed of both secondary and primary
forest seres, offer suitable habitat to more species than either
secondary or primary forest stands alone.
IX

CHAPTER I
INTRODUCTION
General Background
The Atlantic Forest of eastern Brazil 1s one of the most
threatened ecosystems 1n the World due to Intense alteration of the
forested habitat (Mittermeier et al., 1982; Fonseca, 1983).
Fonseca (1983) estimates that approximately 3 - 5X of the native
forest remains today, with most of the remaining forest patches
occurring in protected areas and on privately owned lands. These
remaining forests are composed of mixed stands of primary, and
mostly, secondary forest seres. The Impact of these isolated and
heterogeneous forests on the wildlife communities has not been
studied. Only preliminary surveys have been conducted for the
primates that occur in this region and the remaining vertebrate
fauna is poorly known. This study focuses on small mammal
communities 1n a variety of habitats in one of the largest
remaining protected areas in the Atlantic Forest of eastern Brazil.
This study examines temporal and spatial variation in small mammal
communities in a region severely threatened by human activities.
1

2
The notion of prevailing stable environmental conditions 1n
the tropics helped to support the belief that tropical species
exhibit less temporal variation 1n reproduction, activity, and
feeding than temperate species. A current body of data reveals
that the tropics are much more dynamic than previously believed
(e.g., Leigh et al., 1982). This 1s especially true of tropical
areas that experience pronounced wet and dry seasons. The
Brazilian Atlantic region has such pronounced seasons. With this
in mind, this study Investigated the effect of time on the trapping
success of small mammals.
Habitat structure has been shown to affect the species
composition and relative abundance of small mammals in forested
(Dueser and Shugart, 1978; August, 1984) as well as 1n non-
forested systems (August, 1984; Lacher et al., in press; Lacher and
Alho, 1988). Tropical forest small mammals form a diverse group
that has a varied diet, use of vertical space, and activity time.
In addition, South American small mammals can be composed of two
large groups, rodents and marsupials, in comparison to North
American communities. These factors will be taken Into
consideration in determining how these small mammals allocate the
available space and resources in forest stands of differing
structure.
In some forest communities, disturbances can help to increase
floristic diversity (Connell, 1978; Horn, 1974). Connell (1978)
suggested that intermediate disturbances promote species diversity
in tropical forests and in coral reefs. This pattern has not been

3
observed for tropical vertebrate fauna, however, the relationship
is Intuitive. Early secondary and primary forests are
characterized as spatially homogeneous, while forests disturbed in
an intermediate fashion are spatially heterogeneous. Forest seres
characterized by a high degree of spatial heterogeneity should
provide more resources for more species in comparison to relatively
homogeneous environments. This study Investigated small mammal
communities 1n a variety of forest habitats that were affected by
forest fire.
Study Organization
This dissertation 1s divided into four major chapters. Each
chapter reads as an Independent paper, but all of the chapters have
a common theme. The theme 1s the small mammal communities in the
Rio Doce State Forestry Park 1n eastern Minas Gerais, Brazil. This
Park lies in the Rio Doce Valley and is part of the highly
endangered Atlantic Forest system of eastern Brazil. The primate
fauna of this region 1s highly endemic and very endangered
(Mittermeier et al., 1982). Inventories of other mammal species
are lacking for this region, although, preliminary species lists
indicated a highly endemic small mammal fauna.
Chapter II 1s concerned with intensive small mammal
inventories in this park. This chapter describes the major habitat
types that were sampled, the sampling methodology, the effect of
trap position and type, and the small mammals that occurred in each

4
major habitat type. Chapter II sets the stage for the subsequent
chapters.
Chapter III examines the temporal variation 1n trapping
success of marsupials. Many species of small mammals have a
"window of vulnerability" or a period of time during the year when
they are trapped with greater frequency. This chapter addresses
this question by examining the pattern of trapping success of
didelphld marsupials through time.
The fourth chapter addresses the effect of habitat structure
on the use of space by small mammals. I sampled small mammal
communities 1n 5 forested habitats. Individual species were
compared by their use of vertical space, diet, and time of
activity.
Chapter V examines the effect of forest fire in structuring
the small mammal communities 1n the R1o Doce Park. Tropical forest
fire has not been recognized as a determinant of species diversity.
In this chapter I suggest that even a low frequency fire can set
back succession 1n a forest system that may require 300 years to
reach "climax" conditions. Fires in humid forests can occur
because of unseasonably dry periods that produce optimal conditions
for intense crown fires. Gaps or patches produced by fires can
affect species diversity of small mammals. Disturbances of an
intermediate fashion, such as fire mosaics, should Increase species
diversity because it produces a mixture of successional seres that
are suitable for a number of small mammal species.

CHAPTER II
SMALL MAMMAL INVENTORIES IN AN EASTERN BRAZILIAN PARK
Introduction
In comparison to its geographical size, few studies have been
conducted on small mammal communities 1n Brazil. Most published
reports on Brazilian mammals are preliminary species lists (e.g.
Avila-Pires and Gouvea, 1977) or inventories (e.g., Vieira, 1955;
Moojen, 1952). Some Brazilian studies have focused on densities
(Emmons, 1984), others have focused on abiotic effects on small
mammals (Borchert and Hansen, 1983; Peterson, in press), while
others have addressed such diverse subjects as plantation effects
(Dietz et al., 1975) and public health needs or mammals that carry
human diseases (e.g., Laemment et al., 1946; Botelho and Linardi,
1980; Dias, 1982).
Recently, work 1n Brazil using mark-release techniques has
addressed the use of space, longevity, diversity and social habits
of small mammals (see Alho, 1982). As Alho (1982) pointed out,
most of these studies were concentrated in the xerophitic Cerrado
and Caatinga habitats. Fewer studies have been carried out 1n the
humid forests. Carvalho (1965) live trapped small mammals in a
tropical humid forest in Sao Paulo. Malcolm (pers. comm.) trapped
small mammals in tropical humid forest near Manaus, Amazonia.
5

6
Fonseca (pers. comm.) worked on small mammals in a range of
habitat types 1n eastern Minas Gerais.
The purpose of this paper 1s to report on intensive small
mammal inventories in a tropical humid Brazilian forest, the Rio
Doce State Forestry Park. This Park occurs within the geographical
limits of the highly endangered Brazilian Atlantic Forest (Fonseca,
1983). Small mammal Inventories are lacking from this park.
Gastal (1982) and Avila-P1res (1978) report preliminary results
from intermittent small mammal Inventories in the Rio Doce park.
The Brazilian Atlantic forest has a highly diverse flora and fauna,
with many endemic species of trees (Mori et al., 1981), reptiles
(Muller, 1973), and birds (Haffer, 1974). In-depth mammal
inventories in the region are lacking. The mammalian fauna is
poorly known. Mittermeier et al. (1982) and Klnzey (1982) report
on the high level of diversity and endemism found in the primates
of the region. Preliminary species lists for non-volant mammals in
this region suggest a very high diversity and endemism (Cabrera,
1957, 1961; Honacki et al., 1982; Moojen, 1952; and Vieira, 1955).
For this project, a preliminary checklist was prepared of the non¬
volant mammal species that probably occur in the Atlantic Forest
region (Table A-1). These data Indicate that for the region there
are at least 131 species, 50 of which (39%) are endemic.

7
Study Site
Small mammal trapping was carried out 1n the R1o Doce State
Forestry Park (19 48’18" and 19 29’24" south latitude and 42
38’30" and 42 28’18“ west longitude). The Park was created in 1944
at the request of Dorn Helvecio, the bishop of the region (Gllhaus,
1986). The State Forestry Institute of the state of Minas Gerais
is the present administrative body.
The climate of the Park 1s classified as tropical humid
(Gllhaus, 1986) with a seasonal pulse of precipitation from
November through February and a pronounced dry season from June
through August. Average annual rainfall for a 20 year period
(CETEC, 1981) was 1480 mm, although the rainfall recorded during
this year of the study was considerably less (Figure 2-1). Mean
annual temperature averages 22 C (CETEC, 1981) and mean minimum
monthly temperatures vary greatly throughout the year (Figure 2-2).
The Park boundaries on the north and the east are two rivers,
the R1o P1rac1caba and the R1o Doce, respectively (Figure 2-3).
The southern and western boundaries abut plantations of Eucalyptus
spp. forests.
The predominant relief forms 1n the Park are hills originating
from fluvial plane dissection and valleys derived from fluvial
deposits (Gllhaus, 1986). The altitude in the Park varies from 230
to 515 m. CETEC (1981) reports that 21* of the Park is composed of
plains, 40* undulating to strongly undulating hills and 34*
strongly undulating hills to mountainous terrain.

8
WALTER AND L
CL3MAT3C DIAGRAM
MONTHLY RAINFALL (MM.)
MEAN TEMPERATURE (0 )
JAN PEB MAP APR MAY jun JUL AUG SEP OOT NOV OEC
MONTHS
AVERAGE ANNUAL RAINFALL 1*00 mm
OATA rPGM 1364->874
WALTER AND LEITH
CLIMATIC DIAGRAM
MONTHLY RAINFALL MM)
MEAN TEMPERATURE iC )
MONTHS
ANNUAL AVERAGE RAINFALL BOA mm
3ATA FROM 1885-1388
Figure 2-1. Walter and Leith climatic diagram characterizing
precipitation surplus and drought per month 1n Rio Doce State
Forestry Park, Minas Gerais, Brazil. A= data collected for a
twenty year period (1954-1974) and B= data collected during
study.

9
TEMPERATURE (C )
MONTHS
MEAN MAXIMUM TEMP —MEAN MINIMUM TEMP
DATA FROM 1954 - 1974
Figure 2-2. Temperature graph showing pronounced decrease 1n
minimum temperature during June, July, and August in Rio Doce State
Forestry Park, Minas Gerais, Brazil.

10
RIO DOCE STATE
FORESTRY PARK
LEGEND
sites (3)
burned areas
eucalyptus
N
01234 km
Figure 2-3. Map of Rio Doce State Forestry Park, Minas Gerais,
Brazil. Numbers circled indicate trapping sites.

11
A unique feature of the R1o Doce State Forestry Park (PFERD)
1s the system of R1o Doce Valley lakes that occur there. There are
approximately 40 lakes and numerous marshes in the Park (Saijo and
Tundisi, 1985). According to Saijo and Tundlsi (1985) the lakes
were formed by damming the drainage river of the Rio Doce
watershed. The marshes in the Park are the result of the
sedimentation of previous lakes.
The vegetation of the park 1s classified as tropical semi-
deciduous (Gilhaus, 1986). Most of the emergent tree species lose
their leaves during the cool dry months (J. Stallings, per. obs.).
Forest fire has been a major threat to the vegetation and wildlife
in the park because of the Utter that accumulates during the dry
season. In 1964 and 1967, major fires burned approximately 30* of
the park (Lopes, 1982; Silva-Neto, 1984).
Gilhaus (1986) described 5 forested and 5 open/field habitats
for the Park (Table 2-1). All of the forested and open/field habitats
have been altered to some extent by fire, with the exception of the
Tall Primary Forest with Epiphytes.
Methods
I live-trapped small mammals in 4 general habitas types: (1)
homogeneous eucalypt forest, (2) eucalypt forest with native species
subcanopy, (3) wet meadow, and (4) native forested habitats. I
trapped in 5 sites within the native forested habitat type. Two of
the sites (RD/C and RD/T) were primary forests and corresponded to
the Gilhaus (1986) classification of Tall Primary Forest with

12
Epiphytes. Two other sites (RD/H and RD/M) were altered by forest
fire in 1967 and burned in an intermediate fashion which produced a
forest mosaic of short, secondary and tall forest. This habitat type
corresponded to the Medium to Tall Forest with Bamboos and
Graminoids (Gilhaus, 1986). The remaining site (RD/F) was burned
completely to the ground in 1967 and the resulting vegetation type
corresponded to the Medium Secondary Forest with Bamboos and
Graminoids (Gilhaus, 1986).
The wet meadow habitat (RD/B) corresponded to both the Low
Woodland and Low Tree and Scrub Tallgrass Savanna classified by
Gilhaus (1986). This habitat type occurs between the edge of
permanent marshes and secondary forest. Grasses from the family of
Graminae are the dominant vegetative cover and were introduced in
the region as food for cattle.
The eucalypt forest with native subcanopy habitat was planted
with Eucalyptus saligna in 1954 after the original vegetative cover
was removed. The eucalypt forest was harvested selectively in 1964
and again in 1971. However, the eucalypt forest was never clear-cut
and the native species were allowed to regenerate, largely through
coppicing, and developed into a complex native subcanopy. The result
is a homogeneous eucalypt upper canopy and a native species
subcanopy or "mata su.ia." Tall exotic grasses cover the ground.
Emergent eucalypt trees reach 20 m in height.
The homogeneous eucalypt forest was first planted with
Eucalyptus saligna in 1954 and was harvested 3 times on a 7 year
rotation. There is no native woody herbaceous subcanopy, only

13
Table 2-1. Forested and open/field habitats that occur in the Rio
Doce State Forestry Park, Minas Gerais, Brazil. Definitions and
characteristics of habitats taken from Gilhaus (1986).
HABITAT TYPE
% TOTAL
COMPARISON TO
THIS STUDY
FORESTED
TALL PRIMARY FOREST
WITH EPIPHYTES
8.4
RD/C, RD/T
PRIMARY FOREST
TALL FOREST
30.0
—
MEDIUM TO TALL FOREST WITH
BAMBOOS AND GRAMINOIDS
30.6
RD/M, RD/H
MOSAIC FOREST
MEDIUM SECONDARY FOREST WITH
BAMBOOS AND GRAMINOIDS
17.2
RD/F
LOW SECONDARY
FOREST
LOW SECONDARY FOREST
0.1
—
OPEN/FIELD
LOW WOODLAND
1.1
RD/B
WET MEADOW
LOW TREE AND SCRUB TALLGRASS
SAVANNA
0.6
RD/B
WET MEADOW
TALLFERN FIELD
0.1
—
EVERGREEN TALLGRASS FIELD
WITH TvDha sd.
3.0
—
PARTIALLY SUBMERGED
SHORTHERB FIELD
AND
AQUATIC HABITAT
8.9
100.0
—

14
eucalypt trees. Tall exotic grasses are the dominant ground cover.
Eucalypt trees reach approximately 15 m In height.
Individuals were snap trapped in order to obtain voucher
specimens, dietary, and reproductive information. Snap trapping was
carried out exclusively in wet meadow and secondary succession
habitats. All specimens were either preserved in 10% formalin or
made into museum study skins with corresponding complete skulls. I
also made study skins of Individuals of species of uncertain taxonomic
status that were live trapped in habitats other than those where snap
trapping was carried out.
I placed one line of terrestrial Sherman live traps 1n the
homogeneous eucalypt habitat. Twenty-five Shermans, baited with dry
oatmeal, pineapple chunks and cotton balls soaked with cod-liver oil,
were opened for 4 consecutive nights on 5 occasions through the
study. Trap stations were separated by 15 m.
In the wet meadow habitat, live trapping started in February,
1986, and continued at monthly intervals through January, 1987. I
used two parallel trapping lines in this habitat. Each line was 280 m
in length and subdivided into 20 trap stations separated by 15 m. All
traps were placed on the ground, with even numbered stations having
only one Sherman Uve trap while odd numbered stations had one
Sherman and one locally made small wire live trap. Bait was
identical to that used in the homogeneous eucalypt forest.
In the eucalypt forest with native subcanopy and the native
forested habitats, the trapping design was identical (Table 2-2). In
each area, I cut 3 parallel lines 300 m in length through the forest.

15
Table 2-2. Trapping design used in native forested and eucalypt
forest with native species subcanopy habitats. Trapping lines are
A, B, and C. Numbers 1 - 16 represent trapping posts. Terrestrial
medium sized live traps = ¡, arboreal medium sized live traps = *,
small arboreal Sherman live traps = S, small terrestrial Sherman
live trap = s, large terrestrial live trap = L. Trapping lines
were separated by 100 m.
TRAPPING
STATION
TRAPPING LINES
A
B
C
1 *\ ¡S
2 L;S *\
3 *! ! s
4 l ¡ s *¡
5 *¡ ¡s
6 l;s *¡
7 *1 Is
8 L;s *1
9 *¡ IS
10 l;s *;
11 *l Is
12 L|s *|
13 *! | s
14 L: S *\
15 *¡ Is
L | s *!
16

16
Sixteen trapping stations were placed along the line separated by 20
m. I used Sherman live traps and locally made small (15 X 15 X 30
cm) and large (25 X 30 X 60 cm) wire live traps. All traps were
placed within 3.5 m of the trapping post. All trapping posts had a
terrestrial small Uve trap. Odd numbered trapping posts had a small
wire live trap placed in a tree or bush. Mean arboreal live trap
height was 1.2 m. Odd numbered trapping posts had a Sherman live
trap alternating between arboreal and terrestrial positions. The
exterior trapping lines were identical with respect to number, kind
and placement of traps. I did not use the large live traps on the
interior line. Aside from the large live traps, the exterior and
interior lines were equal in trap number. However, the positions of
the Sherman and small live wire traps were reversed for the interior
line. The Sherman live traps were introduced Into each of the
forested habitats after the study was well underway in an attempt to
sample smaller bodied species. I placed Shermans in two of the
native forested sites in January, 1986, and introduced Shermans in the
remaining forested sites in May of the same year.
I also experimented with traps that were placed high in the
canopy by a pulley and platform device. This method was similar to
that developed by Malcolm (pers. comm.) for use in the Brazilian
Amazon. Mean trap height was 11.2 m. I used these arboreal traps
to sample the canopy dwelling small mammals. Traps were located at
trapping posts along the established lines. The post and exact trap
location in each tree were determined in a subjective manner. I
placed traps in trees which I thought had a high degree of canopy

17
connectivity and upper stratum vine density. Forty-two arboreal
platforms were spread across 4 native forested sites. The primary
forest sites, RD/C and RD/T, and one of the mosaic sites, RD/H,
each had 12 arboreal traps, 4 traps per line. The other mosaic site,
RD/M, had only 6 traps because I did not believe that there was
sufficient upper strata development to support canopy dwelling
species. Arboreal canopy trapping started in June, 1986, and
continued through October, 1986. Trapping coincided with the
schedule developed for the terrestrial and arboreal trapping.
I used dry oatmeal, pineapple chunks and cotton balls soaked
with cod-liver oil for bait. Traps were set during the day and
remained open for 5 consecutive nights each month for one calendar
year.
The first occasion that an individual was trapped was considered
a first capture. The first capture plus subsequent captures of each
individual were considered total captures. Minimum known alive
(MKA) was the number of individuals actually captured each month
whether the capture was the first capture or a recapture from a
different session.
Trapping success of small mammals was calculated in the
following manner. I multiplied the number of traps by the number of
nights the traps were baited and armed per site per month to
determine the number of trap nights. Trapping success was the
number of individuals, MKA or total captures of all species divided by
the number of trap nights and expressed in percentages. For
example, if 100 individuals were trapped during 1000 trap nights, the

18
trapping success would be (100/1000) X 100 = 10X. Recapture indices
were calculated by dividing total captures by first captures and
indicated the average number of times an individual of each species
was captured.
I recorded the following information from each captured small
mammal: date, location on trapping line, position of trap, species, sex,
whether the trapped animal was a juvenile or adult, it’s reproductive
condition, general condition, external parasitic load on a relative
scale, and behavior upon release. I also recorded standard body
measurements for each Individual: body length, tail, ear, hind foot
and mass. I placed a numbered metal eartag in the left ear of each
individual upon the initial trapping of the individual.
Taxonomic determination of questionable species was made by
taxonomists specializing 1n various small mammal groups. Voucher
specimens were brought to the United States and the taxa were
distributed to the following people: cricetid rodents and marsupials of
the genus Marmosa were sent to Dr. Phil Myers, University of
Michigan; rodents of the subgenus Oecomys were sent to Dr. Guy
Musser, American Museum of Natural History, and to Dr. Mike
Carleton, U.S. National Museum of Natural History.
I used body measurements, weight, reproductive condition and
pelage characteristics to determine the age class (juvenile or adult)
of each captured Individual. An individual was considered an adult if
it was reproductively active. Female rodents were considered
reproductively active if they 1) had a perforated vulva, 2) were
pregnant, or 3) were lactating. Marsupial females were considered

19
reproductive!y active if they 1) were lactating or 2) had young
attached to the teat field. Hale rodents were considered
reproductively active if the testes were descended. I could not
determine the reproductive status of male marsupials as the testes are
permanently descended. However, the activity state of the sternal
gland in marsupials can indicate the reproductive time of year.
Initially, I used the overall gestalt of each individual of each sex to
assign age classes. Later, I compared my initial classification with
the body measurements and mass. Body measurements and weights
were sorted for each sex of each species and plotted to determine if
recorded values could be grouped according to size. I then assigned
a body measurement value as the threshold for separating juvenile
and adult age classes. These age classes were then compared to the
initial age classes that were assigned in the field.
I used the General Linear Program (PC-SAS) ANOVA to test for
the equivalence of adult body measurements and mass means between
sexes for each species. This analysis enabled me to determine the
extent of sexual dimorphism for the external characters. Statistical
significance was set at <. 0.05.
Feeding categories were determined by stomach content analysis
(Charles-Dominique et al., 1981) and from the literature. I relied
heavily on information gleaned from the literature on food
preferences of small Neotropical mammals.
The use of vertical space (1.e., terrestrial, scansorial, or
arboreal) of each species was determined from 3 data sets. I
compared the proportion of captures in trees to that of captures on

20
the ground for each species. There were more terrestrial than
arboreal traps and this bias was corrected by adjusting the number of
total trapping opportunities. For this adjustment, I divided the
number of arboreal total captures by the total arboreal opportunities
(or total arboreal trap nights). The same was done for terrestrial
total captures. Results of these two divisions were summed and each
respective result (e.g., arboreal) was divided into the sum. The result
generated adjusted percentage success of arboreal and terrestrial
captures. The sum of total captures was multiplied by the adjusted
percentage in order to generate adjusted number of captures per
trapping stratum (on the ground or in arborescent vegetation). These
adjustments also were made for each species at each site as well as
pooled adjustments across all sites. I tested the null hypothesis that
there was no difference in the proportion of arboreal and terrestrial
captures for each species across all habitats as well as within each
trapping site. Prior to using Student’s t-test, I arc-sin transformed
the proportions. I then compared the proportion of arboreal and
terrestrial responses upon release for each species across all habitat
types. These adjustments were made for all species in each habitat
type.
I used Student’s t-tests to test the null hypothesis that there
was no difference in the locomotory response upon release of each
species. These 2 data sets were then compared to determine if there
were any differences between where an individual was captured and
its locomotory behavior upon release across all habitat types. These
tests also were performed for each habitat sampled. A species was

21
considered to be arboreal 1f that species was found to have a high
proportion of arboreal captures and a high proportion of arboreal
behavior upon release. The opposite would be true for a terrestrial
species. A species would be scansorial if there were no significant
differences in the proportion of spatial captures and no significant
differences in the proportion of behaviors exhibited upon release. I
then compared my results obtained from the trapping data to the
available literature for each species.
I also recorded the presence and relative abundance of other
vertebrates while walking the trapping lines 1n each sampling area.
Upon encountering an animal, I recorded the species, distance in
meters from the trail, height in trees for arboreal species, hour of
day, group size and other natural history data. I moved at a pace of
approximately 1 km. per hour and covered a distance of 1050 m per
sampling area. These walks yielded 60 km of repeat censuses per
sampling area. This method does not allow for the computation of H’
indices because it is difficult to ascertain the identity of individuals
being censused. Because of this problem, I only calculated species
richness per taxonomic group, number of encounters of each species
through time, and number of encounters of each group per linear km.
These data are presented in Table A-3.
I also superficially sampled the bat fauna of the Park. I
mounted mist nets in various habitat types (e.g., lake edge, secondary
habitat, primary habitats, and manmade structures) in order to
determine the species that occurred in the park. Nets were mounted
one hour before dusk and opened for approximately 4 hours each

22
night. Mist netting occurred on a sporadic basis. A species list of
bats obtained by these methods is presented in Table A-4.
RQS.u.lts
Species Accounts
I logged 1,308 captures of 17 species of small mammals in 40,490
trap nights. The small mammal fauna of the park consisted of 6
species of marsupials and 11 species of rodents. The diet, use of
vertical space and habitat requirements are presented in Table 2-3 for
each species captured during the study.
Marsupials - - Family Didelphidae.
Didelphis marsupial is Linne. (1758). The black-eared opossum
ranges widely in South America from the Isthmus of Panama to
southern Brazil. This species occurs sympatrically with D. albiventris
throughout much of its range (Strellein, 1982). In the Rio Doce
Valley, however, D. marsupial is inhabits moister habitats, while D.
albiventris occurs in the cerrado vegetation (Valle and Varejao, 1981).
A. Gardner (pers. comm.) suggested that the form of D. marsupial is
found in eastern Brazil is distinct and should be referred to as D.
azarae. This species Inhabits brushy and forested habitats (Alho,
1982; Nowak and Paradiso, 1983). Miles et al. (1981) found this
species to be nocturnal, with a preference for nesting in tree
cavities. I captured this species in all forested habitats in the park
(Table 2-4). Adult body measurements do not indicate sexual
dimorphism (Table A-2). Females have a well developed pouch. This

23
Table 2-3. Ecological role played by each species captured during
this study in the Rio Doce State Forestry Park. GM= grasslands and
wet meadows, B= brushy areas, S= secondary forests, P= primary
forests, F= fossorial or semifossorial, SA= semiaquatic, T=
terrestrial, S= scansorial, A= arboreal, HG= herbivore-grazer; FG=
frugivore-granivore; F0= frugivore-omnivore; 10= insectivore-
omnivore. * Taxa endemic to the Brazilian Atlantic rainforest or
the eastern coastal area of South America.
SPATIAL
DIETARY
SPECIES
HABITAT
ADAPTATION
CLASSIFICATION
MARSUPIALS
DidelDhis marsuDialis
B, S,
P
T, S
FO
Metachi rus nudicaudatus
s,
P
T
IO/FO
Marmosa incana*
B, S,
P
S
10
M. cinerea
B, S,
P
A
10
M. microtarsus*
s,
P
A
10
Caluromys philander
s,
P
A
FO
RODENTS
Oecomvs trinitatis
s,
P
S
FG
OrYZomvs caoito
s,
P
T
FG
0. subflavus*
GM, B,
S
T
FG
0. niqriDes*
GM,
B
S
FG
Akodon cursor*
GM, B,
S
T
10
Calomvs laucha
GM,
B
T
FG
Nectomvs sauamiDes
GM,
B
SA
HG
Abrawavaomvs ruschii*
S
T
FG?
OxvmYcterus roberti*
GM, B,
S
F
10
Rhioidomvs mastacalis
s,
P
A
FG
Cavia fulqida*
GM,
B
T
HG
EurYZvqomatomvs SDinosus*
GM,
B
F
HG

24
Table 2-4. Number of total captures and percent of total for each
species (SPP.) per habitat type. Numbers in parentheses represent
the percentage of captures per species per habitat percentages are
rounded to the nearest whole number.
SPP*
HABITAT
TYPES+
RD/F
RD/H
RD/M
RD/T
RD/C
RD/E
RD/B
DM
4 (3)
2 (1)
1 (D
7(6)
21(24)
7 (4)
-
MN
20(14)
44(16)
21(15)
30(23)
25(28)
18(11)
-
MI
25(18)
75(27)
20(14)
28(22)
6 (7)
14 (9)
3 (1)
MC
74(52)
71(26)
69(49)
38(30)
31(35)
77(49)
1 (0)
MM
-
1 (0)
-
-
-
-
-
CP
-
16 (6)
2 (1)
13(10)
3 (3)
15(10)
2 (1)
NS
-
6 (2)
9 (6)
-
-
-
5 (1)
RM
-
-
-
7 (6)
-
-
-
AC
3(2)
46(17)
2(1)
-
1 (D
25(16)
315(85)
OT
4 (3)
8 (3)
4 (8)
3 (2)
2 (2)
-
-
OC
-
4 (1)
12 (9)
2 (2)
-
2 (1)
1 (0)
OS
12 (9)
1 (0)
-
-
-
-
31 (8)
ON
-
-
-
-
-
-
4 (1)
OR
-
1 (0)
2 (1)
-
-
-
4 (1)
AR
-
1 (0)
-
-
-
-
-
CL
-
-
-
-
-
4 (1)
ES
-
-
-
-
-
-
2 (1)
142
276
142
128
89
158
373
+ RD/F= SECONDARY HABITAT BURNED COMPLETELY TO THE GROUND; RD/H=
SECONDARY HABITAT BURNED IN MOSAIC FASHION; RD/M= SECONDARY HABITAT
BURNED IN MOSAIC FASHION; RD/T= PRIMARY FOREST WITH LITTLE EFFECT
FROM FOREST FIRE; RD/C= PRIMARY FOREST; RD/E= EUCALYPT FOREST WITH
NATIVE SPECIES SUBCANOPY; RD/B= WET MEADOW.
* DM= Didelphis marsupial is: MN= Metachi rus nudicaudatus: MI=
Marmosa incana: MC= Marmosa cinerea: MM= Marmosa microtarsus: CP=
Caluromys philander: NS= Nectomvs sauamioes: RM= Rhipidomvs
mastacalis: AC= Akodon cursor: 0T= Oecomvs trinitatis: 0C=
Oryzomys capito: 0S= Orvzomvs subflavus: 0N= Orvzomvs nigrioes:
OR= Oxymycterus roberti: AR= Abrawavaomvs ruschi i: CL= Calomvs
laucha: ES= Eurvzygomatomys soinosus.

25
species 1s terrestrial. There was a significant percentage of
terrestrial captures (Table 2-5) and terrestrial behavior upon release
(Table 2-6). I did observe several juvenile individuals and one adult
climb readily. Charles-Domlnique (1983) reported that this species
exploits the lower stratum 1n forests, but can climb. This species is
basically an opportunistic feeder and feeds upon fruit and animal
matter (Charles-Domiñique, 1983).
Metachi rus nudicaudatus Geoffroy (1803). The brown four-eyed
opposum has a geographical distribution similar to that of £).
marsupial is except that it is not found over much of Venezuela nor
in northeastern Brazil (Streilein, 1982). M. nudicaudatus can be
confused with Philander opposum as both have pale spots above the
eyes. In addition, there is considerable confusion over the taxonomy
of the two species. Nowak and Paradiso (1983) classified this species
as Philander nudicaudatus and Philander opossum as Metachiroos
opossum. I agree with Honacki et al. (1982) and follow their
classification. This species was captured in all forested habitats
(Table 2-4). Metachi rus nudicaudatus is sexually dimorphic in Its
mass and hind foot measurements (Table A-2). Females do not have
a pouch. This species is strongly terrestrial, rarely caught in
arboreal traps (Table 2-5) and rarely climbs upon release (Table 2-6).
Miles et al. (1981) found this species to be nocturnal and construct
nests on the forest floor or in ground hollows. There are very little
data on the feeding habits of this species due to the small numbers
that have been reported to be trapped. Preliminary data indicate

26
Table 2-5. Student’s t-tests between adjusted and arcsin
transformed percentages of terrestrial and arboreal captures of
small mammals in all forest types. NS = non significant.
X TERRESTRIAL % ARBOREAL
SPECIES
CAPTURES
CAPTURES
V
P <
DidelDhis marsuoialis
66.6
23.5
40
.001
Metachi rus nudicaudatus
76.3
3.9
156
.001
Marmosa incana
43.9
46.1
169
NS
Marmosa cinerea
27.8
62.2
356
.001
Caluromvs Dhilander
16.3
73.8
49
.001
Nectomvs sauamioes
69.6
21.0
17
.05
RhiDidomvs mastacalis
28.8
61.1
5
NS
Akodon cursor
83.5
6.3
390
.001
Oecomvs trinitatis
31.2
58.8
19
NS
OrYzomvs caoito
62.9
27.0
19
.05
OrYzomvs subflavus
58.0
32.5
46
.01

27
Table 2-6. Results of Student’s t-tests between terrestrial and
arboreal behavior upon release of small mammals captured in all
habitats. NS= non significant. Species abbreviations are
explained in Table 2-4.
SPECIES
X
N
TERRESTRIAL
BEHAVIOR
% ARBOREAL
N BEHAVIOR
V
T
P <
DM
33
73.2
3
16.7
34
3.265
0.01
MN
150
80.7
4
9.1
152
4.934
0.001
MI
95
53.5
52
36.5
145
3.448
0.001
MC
11
10.8
294
78.9
303
7.743
0.001
CP
2
18.4
18
71.6
18
2.488
0.05
NS
12
90.0
0
0.0
10
10.882
0.001
RM
1
24.0
5
67.8
4
1.393
NS
AC
70
90.0
0
0.0
68
26.282
0.001
OT
9
64.8
2
25.3
9
1.763
NS
OC
13
90.0
0
0.0
11
11.326
0.001
OS
10
90.0
0
0.0
8
9.933
0.001

28
that this species is an insectivore-omnivore (Robinson and Redford,
1986) or frugivore-omnivore (Hunsaker, 1977).
Marmosa incana Lund 1840. This mouse opposum is endemic to the
Brazilian Atlantic Rainforest (Streilein, 1982). M. incana occurs in
both secondary and primary forest habitat. This species 1s small in
size (adults, average weight = 62 g) and strongly sexually dimorphic
in body size and color. Males tend to have larger ears and hind feet
(Table A-2) while females tend to have a more rose colored venter
and less pronounced face mask (P. Myers, pers. 1itt.). Females do not
have a true pouch. This species tends to use both the ground and
arborescent vegetation. I classify the spatial adaptation of this
species as scansorial (Table 2-3). There was no significant difference
in the proportion of terrestrial and arboreal captures (Table 2-5, p >
.90, df=169), however; individuals tended to remain on the ground
upon release (Table 2-6, p < 0.001, df=145). No data exist on the
feeding category of this species. Other species of Marmosa which
have similar body mass are classified as insectivore-omnivores.
Stomach content analysis (n=3) showed 100% insects from two orders,
Coleóptera and Orthoptera (Table 2-7). I classify this species as an
insect1vore-omnivore based on the relationship found between body
mass and dietary classification (Robinson and Redford, 1986).

29
Table 2-7. Stomach content analysis of small mammals captured in
Rio Doce State Forestry Park, Minas Gerais, Brazil (N= number of
stomachs analyzed).
SPECIES
FRUIT SEEDS GRASS INSECTS
Marmosa incana (n=3)
Akodon cursor (n=23)
Orvzomvs subflavus (n=1)
Orvzomvs nigripes (n=5)
Oxvmvcterus roberti (n=2)
Calomvs laucha (n=1)
Nectomvs sauamipes (n=2)
19%
19%
8%
5%
-
95%
34%
11%
21%
100%
50% - 50%
100%
52%
34%
100%

30
Marmosa cinérea Temminck (1824). M. cinérea has a disjunct
geographical distribution in South America; it occurs in northern
Venezuela through the Guianas, and it occurs in the Brazilian
Atlantic Rainforest extending into Paraguay (Streilien, 1982). M.
cinerea occurs in brushy and forested habitat, ranging from secondary
to primary. This species is a large bodied Marmosa (Table A-2,
average weight= 105g) and 1s highly sexually dimorphic based on
external body measurements. Males tended to have larger body, tail,
ear and foot (Table A-2). Females do not have a pouch. This
species is strongly arboreal and exploits the high forest stratum
(Charies-Domin1que, 1983; Miles et al., 1981). Miles et al. (1981)
found this species to be nocturnal and to construct open arboreal
nests rather than use cavities. M. cinerea tended to be caught a
greater proportion of the time in arboreal traps (Table 2-5, p <0.001,
df=356) and tended to exhibit arboreal rather than terrestrial behavior
upon release (Table 2-6, p < 0.001, df=303). This species is primarily
an Insectivore-omnivore (Robinson and Redford, 1986).
Marmosa microtarsus Wagner (1842). This species is restricted to the
Brazilian Atlantic Rainforest (Streilein, 1982). M. microtarsus differs
from its congener M. agí lis by possessing a pure colored white patch
of hairs on the throat and chin (Tate, 1933). There are insufficient
data in the literature to determine if this species is terrestrial,
scansorial, or arboreal. I only recorded one capture during the study,
in an intermediately disturbed habitat (Table 2-4). However, the
species has a long prehensile tail and short wide feet which suggest

31
an arboreal lifestyle. This species is probably an Insectivore-
omnlvore.
Caluromvs philander Linne (1758). This species has a disjunct
geographical distribution with populations in Venezuela, the Guianas
and northern Brazil and in southeastern Brazil (Streilein, 1982). The
woolly opossum is classified as a forest dwelling species (Nowak and
Paradiso, 1983). This species was present in the eucalypt, mosaic,
and primary forested habitats (Table 2-4). Based upon the external
body measurements, there was no sexual dimorphism in adults (Table
A-2). Females lack a true pouch. According to Charles-Dominique
(1983) and Miles et al. (1981) this species exploits the high forest
stratum and is nocturnal. My data showed that there was a higher
proportion of arboreal captures (Table 2-5, p< 0.001, df=49) and that
this species tended to climb more than remain on the ground upon
release (Table 2-6, p < 0.05, df=18). Fruit makes up a large portion
of this species’ diet (Charles-Dominique, 1983; Robinson and Redford,
1986).
Rodents - - Family Cricetidae
Oecomys (Oryzomys) trinitatus = (Q. concolor) Wagner (1845). This
genus is in need of revision and the subgenus Oecomys is currently
being revised (P. Myers, pers. comm.). This species was previously
called Orvzomvs concolor and was known, within Brazil, as an
Amazonian species (Alho, 1982). However, Nitikman and Mares (1987)
reported trapping this species in gallery forest in the Brazilian

32
cerrado. I captured this species in all native forested habitats (Table
2-4). The species is not sexually dimorphic (Table A-2).
Gyldenstolpe (1932) and Moojen (1952) stated that this species is
"more or less adapted for arboreal life." My data suggest that this
rat is scansorial; there were no significant differences in the
proportion of terrestrial and arboreal captures (Table 2-5, p < .10,
df=19) and no significant differences in terrestrial and arboreal
behavior upon release (Table 2-6, p < .20, df= 9). Most species of
the genus Orvzomvs are frugivore-granivores (Robinson and Redford,
1986).
Orvzomvs subflavus Wagner (1842). This species is distributed
throughout the Guianas, southeastern Brazil and eastern Paraguay
(Honackl et al., 1982; Alho, 1982). In Brazil, it occurs in the
cerrado, caatinga and Atlantic Rainforest (Alho, 1982). This species
was captured in wet meadow and heavily disturbed secondary habitat
(Table 2-4). There were no differences in body measurements
between sexes 1n adult Individuals (Table A-2). I classify this species
as terrestrial. This species tended to be captured more on the
ground than in the trees (Table 2-5, p < 0.01, df=46) and was never
observed to climb upon release (Table 2-6). Stomach content analysis
(n=1) showed 95% grass and 5% fruit (Table 2-7).
Oryzomys capito Olfers (1818) = 0. goeldi. laticeps and intermedius.
This species has a wide distribution throughout the Neotropics and
occurs in a variety of habitats ranging from agricultural fields

33
(Moojen, 1952) to humid forests (Alho, 1982). Orvzomvs capito was
primarily captured in humid forests ranging from intermediate levels
of disturbance to primary forests in the park (Table 2-4). There
were no significant differences in body measurements between sexes
for adults (Table A-2). My capture and release data are in
accordance with Alho’s (1982) classification for this species. 0.
capito tended to be caught more on the ground than 1n trees (Table
2-5, p < 0.05, df=19) and tended to remain on the ground upon
release (Table 2-6, p < 0.001, df=11).
Orvzomvs (01igorvzomvs) nigrioes (eliurus) Wagner (1845). This
small-bodied rodent (Table A-2) occurs in grassland, wet meadow and
secondary forest habitat in northern Argentina, eastern Paraguay,
southern Brazil and the Bolivian Beni (Honacki et al., 1982). In the
park, all captures were made in the wet meadow habitat (Table 2-4).
All captures were made on the ground (n=4), however; the Individuals
climbed readily in captivity (J. Stallings, pers. obs.). The results
from the stomach analysis (n=5) revealed a wide range of foodstuffs
(Table 2-7).
Abrawayaomvs ruschii Cunha and Cruz (1979). This species is only
known from the type locality in Espirito Santo, eastern Brazil. It is
endemic to the Brazilian Atlantic Rainforest. The single capture of
this species was recorded in an intermediately disturbed forest (Table
2-4). There is very little information available regarding the ecology

34
of this species and there are only three study skins found in
museums (A. Gardner, pers. comm.).
Rhipidomvs mastacalis Lund (1840). Climbing mice range south from
Margarita and Tobago Islands to Venezuela and Guianas to
northeastern and east central Brazil (Honacki et al., 1982). This
species was only captured in a relatively undisturbed primary forest
(Table 2-4). Sample size was too small to detect any differences
between terrestrial and arboreal captures and behavior upon release.
However, as the common name implies, this species climbs readily. I
captured two individuals 1n my house in the park, a commonly cited
“exotic" habitat for this species (Nowak and Paradiso, 1983).
Nectomvs sauamipes Brants (1827). The neotropical water rat occurs
in aquatic habitats either in grasslands and wet meadows or in
forests. This species distribution ranges from the Guianas to
Colombia to Peru and in Brazil, Paraguay and northeastern Argentina
(Honacki et al., 1982). This species tended to be caught more on the
ground (Table 2-5, p < 0.05, df=17) and exhibited a significant
tendency to remain on the ground upon release (Table 2-6, p < 0.001,
df=10). Stomach content analysis (n=2) showed 50% grass and stems
and 50% fruit (Table 2-7).
Akodon cursor Winge (1887). This species occurs in several habitat
types from southeastern and central Brazil to Uruguay, Paraguay and
northern Argentina. The wet meadow habitat in the park was the

35
primary habitat to capture this species (Table 2-4). A. cursor was
formerly Included in A. arviculoides (Honacki et al., 1982). This
species is sexually dimorphic in tail (p < 0.01) and body (p < 0.008)
length (Table A-2). A. cursor is strongly terrestrial (Table 2-5 and
Table 2-6). Analysis of stomach contents (n=23) revealed a high
proportion of insects, seeds and fruit (Table 2-7).
Ca lorn vs laucha Olfers (1818).. This species occurs 1n grassland and
wet meadows in southern Bolivia, southeastern Brazil, Paraguay,
central Argentina and Uruguay. I only captured this species in the
wet meadow habitat 1n the Park (Table 2-4). The results from one
stomach sample revealed 100% seeds (Table 2-7).
Oxymycterus roberti Thomas (1901). The burrowing mouse occurs in
a variety of habitats but is usually associated with moist substrate in
open or brushy habitats. I captured this species in wet meadow and
secondary habitats in the Park (Table 2-4). This species is endemic
to eastern Brazil. This semifossorial mouse is described as an
insectivore (Nowak and Paradisio, 1983). Stomach analysis (n=2)
revealed 100% insects (Table 2-7).
Family Caviidae
Cavia fulgida Wagler (1831). This species of cavy is endemic to the
open grasslands and wet meadows of the Atlantic Rainforest of
eastern Brazil (Honacki et al., 1982; Nowak and Paradiso, 1983). I
captured this species in grassland and wet meadow habitats in the

36
Park, however; it was not trapped in site RD/B. For this reason this
species does not appear in Table 2-4. Cavies are terrestrial and are
herbivore-grazers (Nowak and Paradiso, 1983).
Family Echimyidae
Eurvzvgomatomvs soinosus Fischer (1814). The single species of
guiara is endemic to southeastern Brazil, Parguay and northeastern
Argentina (Honacki et al., 1982). I captured this species in the wet
meadow habitat (Table 2-4). This species inhabits open grasslands
and wet meadows, is considered terrestrial or semifossorial (Alho,
1982), and is most probably a herbivore-grazer.
Trapping results
Tables 2-8, A-5, and A-6 present the capture results by species
for the three main habitat types: native forested, eucalypt with
native species subcanopy and wet meadow habitats, respectively. As
a group, marsupials represented 79.2% and 83.3% of first and total
captures, respectively, in native forested sites, and 67.7% and 82.9% in
eucalypt forest with native species subcanopy. Rodents represented
97.3% and 98.4% of the first and total captures, respectively, in the
wet meadow habitat.
In the native forested habitat, Marmosa cinerea represented over
40% of the marsupial captures (Table 2-8), while in the eucalypt
forest, this species represented more than 58% of the marsupial
captures (Table A-5). Akodon cursor was the major contributor to
the rodent captures in all three habitats. This species only

37
Table 2-8. Capture results from native forested plots in Rio Doce
State Forestry Park, Minas Gerais, Brazil. RECAP INDEX= total
captures/first captures, and represents the average number of times
that an individual of species X is captured. Numbers in
parentheses represents percent of contribution of capture per
species per taxonomic group. Species abbreviations are explained
in Table 2-4.
TOTAL FIRST RECAP
SPECIES
CAPTURES
% TOTAL
CAPTURES
% TOTAL
INDEX
MARSUPIALS
DM
35
4.5 ( 5.4)
32
7.8 ( 9.9)
1.1
MN
140
18.0 (21.6)
91
22.3 (28.1)
1.5
MI
154
19.8 (23.8)
90
22.1 (27.8)
1.7
MC
283
36.4 (43.7)
92
22.5 (28.4)
3.1
MM
1
0.1 ( 0.2)
1
0.2 ( 0.3)
1.0
CP
34
0.4 ( 5.3)
18
4.4 ( 5.5)
1.9
647
83.2(100.0)
324
79.4(100.0)
RODENTS
NS
15
1.9 (11.5)
9
2.2 (10.7)
1.7
RM
7
0.9 ( 5.4)
3
0.7 ( 3.6)
2.3
AC
52
6.7 (40.0)
27
6.6 (32.1)
1.9
OT
21
2.7 (13.8)
19
4.7 (22.6)
1.1
OC
18
2.3 (13.8)
15
3.7 (17.9)
1.2
OS
13
1.7 (10.0)
7
1.7 ( 8.3)
1.9
OR
3
0.4 ( 2.3)
3
0.7 ( 3.6)
1.0
AR
1
0.1 ( 0.81
1
0.2 ( 1.2)
1.0
130
16.8(100.0)
84
20.6(100.0)

38
Table 2-9. Trapping success of small mammals calculated by habitat
type in Rio Doce
State Forestry Park,
Minas
Gerais, Brazil.
HABITAT TYPE
NUMBER OF
TRAP NIGHTS
NUMBER OF
CAPTURES % SUCCESS
NATIVE FOREST
(EXCLUDING PLATFORMS)
30,960
710
2.3
NATIVE FOREST
PLATFORMS ONLY
1,050
66
6.3
WET MEADOW
1,980
373
18.8
EUCALYPT FOREST
W/NATIVE SPECIES
SUBCANOPY
6,000
158
2.6
EUCALYPT FOREST
W/NO SUBCANOPY
TOTALS
500
40,490
1
1,308
0.0

39
represented about 7% of the total captures in the native forested
habitat, but 40* of the rodent captures (Table 2-8). In the eucalypt
forest, A. cursor represented about 16* of the total captures and 93*
of the rodent captures (Table A-5). This rodent was the dominant
species captured in the wet meadow habitat, representing about 85*
of both the total and of the rodent captures.
In both the native and eucalypt forested habitats, marsupials in
general were recaptured at a high rate. Didelphis marsupial is and
Marmosa microtarsus both showed low recapture rates and reflect the
small sample size. Especially noteworthy was the relatively high
recapture rate of Marmosa cinerea (Tables 2-8 and A-5). No
individuals of other species, neither rodent nor marsupial, were
recaptured as frequently as individuals of this species. In the native
forested habitat, individuals of M. cinerea were recaptured on the
average 3.1 times, while in the eucalypt forest, individuals of this
species were recaptured on the average 7.7 times.
Akodon cursor was the only rodent that had a relatively high
number of captures and recapture rate (Tables A-5 and A-6). This
species had a recapture rate of 1.6 and 3.0 in the eucalypt and wet
meadow habitats, respectively.
Trapping success
Table 2-9 presents the trapping success by habitat type.
Trapping success was calculated for small mammals in the native
forested habitat. The platform trapping data was excluded. It must
be kept in mind that the sampling effort in each general habitat
category was different, however, comparisons of trapping success are

40
PERCENT SUCCESS
MONTHS
EUCALYPT -+- WET MEADOW NATIVE FOREST
Figure 2-4. Comparison of trapping success of small mammals in
three habitat types: eucalypt forest, wet meadow, and native
forest in the Rio Doce State Forestry Park, Minas Gerais, Brazil.

41
the result of the number of captures relative to the number of
trapping opportunities or nights. Overall, the wet meadow habitat
yielded the highest trapping success (18.8%) while the homogeneous
eucalypt habitat generated the lowest trapping success (0.02%).
Figure 2-4 compares the progression of the trapping successes of
the native forested habitat (without the platform data), the wet
meadow habitat and the eucalypt forest with subcanopy habitat.
Although these three habitats have unequal sampling effort and
trapping design, these habitats were sampled for the period of one
year and show important temporal trends. From the overall gross
comparison portrayed in Figure 2-4 and the percent success presented
in Table 2-9, in contrast to the wet meadow habitat, it appears that
the forested habitats, both native and exotic, were quite similar in
overall percent success and monthly trapping success. The wet
meadow trapping success fluctuated greatly throughout time, from a
high approaching 45% in March and April, to a crash lower than 10%
in May, October, and November.
I plotted the total captures, minimum known alive and first
captures through time for each of the three habitats. Figure 2-5
shows the capture curves for the native forested sites. Trapping
success was relatively low and stable from November through April,
with a noticeable increase in June, July and the early part of August.
This peak in trapping success dropped back to the levels observed
prior to the increase. Figures 2-6 and 2-7 compare the trapping
success of the eucalypt forest and the wet meadow habitat,
respectively. The highest number of captures at any one time in the

NUMBER OF CAPTURES
Figure 2-5. Small mammal capture curves for all native forested
sites in Rio Doce State Forestry Park, Hinas Gerais, Brazil.
Capture curves include total captures, minimum known alive (MKA),
and first captures.

43
RD/E EUCALYPT Sill
NUMBER OF CAPTURES
MONTHS
TOTAL CAPTURES MKA CAPTURES FIRST CAPTURES
Figure 2-6. Small mammal capture curves for eucalypt forest 1n
the Rio Doce State Forestry Park, Minas Gerais, Brazil. Capture
curves include total captures, minimum known alive (MKA), and
first captures.

44
NUMBER OF CAPTURES
MONTHS
TOTAL CAPTURES MKA CAPTURES FIRST CAPTURES
Figure 2-7. Small mammal capture curves for wet meadow habitat
in the Rio Doce State Forestry Park, Minas Gerais, Brazil.
Capture curves include total captures, minimum known alive (MKA),
and first captures.

45
Table 2-10.
T rapping
success by trap type for all species in all
habitat types.
, Trap
types are arranged according
to trapping
location: terrestrial or arboreal. Trap types are as follows: 1ST=
small terrestrial Sherman live trap; 3MT=
medium sized
terrestrial
live trap; 5LT= large
terrestrial live
trap; 2SA= small arboreal
Sherman live trap; 4MA
arboreal platform trap.
= medium sized
arboreal live
trap; 6PA=
NO.
NO.
PERCENT
TRAP TYPE
CAPTURES
TRAP NIGHTS
SUCCESS
TERRESTRIAL
1ST
370
4080
9.1
3MT
553
18000
3.1
5LT
42
5760
0.7
965
30480
3.2
ARBOREAL
2SA
48
2640
1.8
4MA
228
8640
2.6
6 PA
66
1050
6.3

46
wet meadow habitat was double the highest in the eucalypt forest.
However, the trends were similar. Both habitats showed two
pronounced peaks in their respective capture curves that corresponded
to the same months throughout the year. There was a peak in
February, March, and April followed by a crash, and another peak in
June, July, and August followed by another crash.
T.rap Types
Overall, trapping success was higher with terrestrial traps than
with arboreal ones (Table 2-10). Caution should be used in these
comparisons as the number of trap nights are unequal; almost three
times the number of terrestrial trap nights as the number of arboreal
ones. However, I feel that some degree of comparison can be drawn
from this analysis based upon the number of captures relative to the
number of trap nights.
Trapping success by trap type varied considerably (Table 2-10).
Small terrestrial Sherman live traps were the most successful (9.1%),
medium terrestrial traps represented the trap with the greatest
number of captures and trapping opportunities, and large terrestrial
live traps were relatively unproductive.
Arboreal trap type success varied. Small arboreal Shermans
were the least productive arboreal trap type with only 1.8% success.
The arboreal platform traps had a trapping success of 6.3%.
Table 2-11 reports the number of total captures of each species
and the number of captures per trap type. Odd numbered traps
represent terrestrial traps and even numbered ones represent arboreal

47
Table 2-11. Trap response by species across all habitat types.
Trap types are explained in Table 2-10. Species abbreviations are
explained in Table 2-4.
SPECIES
TRAP
TYPES
TOTAL
1ST
2SA
3MT
4MA
5LT
6AP
DM
42
0
0
24
2
15
1
MN
158
0
0
129
3
26
0
MI
171
13
14
106
38
0
0
MC
361
2
29
145
150
0
35
CP
51
0
0
9
13
0
29
MM
1
0
0
0
1
0
0
NS
19
4
0
13
1
1
0
RM
7
0
0
3
3
0
1
AC
392
312
2
78
0
0
0
OT
21
1
3
9
8
0
0
OC
21
8
0
11
2
0
0
OR
10
7
0
3
0
0
0
AR
1
0
0
0
1
0
0
OS
44
18
0
20
6
0
0
CL
2
2
0
0
0
0
0
ON
4
3
0
1
0
0
0
EG
2
0
0
2
0
0
0
TOTALS
1307
370
48
553
228
42
66
PERCENTAGES
28.3
3.7
42.3
17.4
3.2
5.0

48
traps. In general, the number of captures per species in terrestrial
and arboreal traps represented the spatial adaptation for each species.
For marsupials, Didelphis marsupial is and Metachi rus nudicaudatus
were captured principally in terrestrial medium live traps (3MT),
while Marmosa incana was captured in all trap types except for the
large terrestrial and arboreal platform traps. Marmosa cinerea was
captured in all trap types except the large terrestrial traps.
Caluromys philander was trapped principally in arboreal medium live
traps and arboreal platform traps. The sample size for rodents was
too small to allow for a clear trapping trend. Akodon cursor is the
clear exception. The small terrestrial Sherman live trap was most
effective for this species. This was mostly a consequence of the
approximately 80% of all Akodon captures made in the wet meadow
habitat (Table 2-10). Orvzomvs subflavus was trapped principally in
terrestrial small shermans and medium live traps.
Discussion
The trapping success realized in this study for neotropical humid
forests falls within the range of observed success rates (Table 2-12).
However, one striking difference between this and other neotropical
small mammal studies was the high number of marsupial captures
relative to rodent captures (Table 2-12). All reported studies show
rodent biases, and usually high captures of rodents relative to
marsupials. Only Emmons (1984) reported a marsupial to rodent
capture ratio approaching equality (Table 2-12). Fonseca (pers.
comm.) reported marsupial biased trapping results from a variety of

49
Table 2-12. Percent capture by taxonomic group, trapping success
and number of trap nights by habitat type for this study compared
to other Neotropical field studies. % M= percent of total
marsupial captures; % R= percent of total rodent captures; % T.S.=
percent trapping success; # T.N.= mumber of trap nights.
STUDY
SITE
% M
% R
% T.S.
# T.N.
NATIVE FOREST
THIS STUDY
BRAZIL
83.2
16.8
2.3
30,960
DIETZ ET AL., 1975
BRAZIL
9.3
90.7
—
—
CARVALHO, 1965
BRAZIL
0.3
99.7
3.6
10,080
EMMONS, 1984
PERU
48.0
52.0
6.9
2,987
EMMONS, 1984
PERU
—
—
7.0
4,390
EMMONS, 1984
BRAZIL
—
—
0.8
434
DIAS, 1982
BRAZIL
2.3
97.7
—
—
NITIKMAN & MARES,
1987
BRAZIL
30.7
69.3
6.0
12,170
LAEMMENT ET AL.,
1946
BRAZIL
31.0
69.0
10.0
30,000
AUGUST, 1984
VENEZUELA
25.0
75.0
0.9
30,269
DAVIS, 1945
BRAZIL
17.0
83.0
—
—
FLEMING, 1972
1974
PANAMA
19.0
81.0
16.0
24,732
O’CONNELL, 1979
VENEZUELA
12.0
88.0
—
—
WET MEADOW/SAVANNA/PANTANAL
THIS STUDY
BRAZIL
2.0
98.0
18.8
1,980
AUGUST, 1984
VENEZUELA
0.0
100.0
1.9
3,660
AUGUST, 1984
VENEZUELA
25.0
75.0
0.1
4,400
LACHER & ALHO,
IN PRESS
BRAZIL
0.0
100.0
4.2
3,582
BORCHERT & HANSON,
1983
BRAZIL
0.0
100.0
3.5
4,173
O’CONNELL, 1981
VENEZUELA
10.0
90.0
—
—
HOMOGENEOUS EUCALYPT
FOREST
DIETZ ET AL., 1975
BRAZIL
0.0
100.0
THIS STUDY
BRAZIL
0.0
100.0
0.2
500
EUCALYPT FOREST WITH
I NATIVE SUBCANOPY
THIS STUDY
BRAZIL
83.0
17.0
2.6
6,000

50
native forest sites in eastern Minas Gerais. Avila-Pires (1978)
captured 245 rodents and 40 marsupials from the R1o Doce Park.
Gastal (1982) reported that 5 species of marsupials were captured in
the Park but gave no comparative data for rodents. Dias (1982)
trapped more rodents than marsupials in the Rio Doce Valley in
Minas Gerais.
Hunsaker (1977) stated that marsupials require considerable
effort to trap. Perhaps one explanation for the observed high
marsupial/rodent capture ratio could be due to the habitat type found
in the Park. There is very little primary habitat in the Park relative
to secondary habitat (Table 2-1). Most of the forest habitat in the
Park has been altered by fire in the recent past (Lopes, 1982). The
primary forest plots that I sampled yielded the lowest species
richness and absolute captures of didelphid marsupials relative to the
other secondary forested habitats (J. Stallings, in prep.). Charles-
Dominique (1983) suggested that didelphid marsupials can reach high
local densities in areas of abundant food resources. He postulated
that these species are r-strategists and are adapted to the "unstable
environment of secondary forests.” In Panama, Didelphis marsupial is
tended to occur at higher densities in primary forest, while
Caluromvs and Philander were found at higher densities in secondary
forest than primary forest (Fleming, 1972).
Marsupials were recaptured with great frequency in this study,
especially Metachi rus nudicaudatus. Marmosa incana and Marmosa
cinerea. My recapture data on marsupials agree with data reported
by Fleming (1972, 1973), August (1984) and to some degree with that

51
found by O’Connell (1979). My data do not agree with Hunsaker
(1977) who stated that didelphid marsupials are difficult to recapture.
For example, individuals of Marmosa cinerea were captured on the
average 7.7 times in the eucalypt forest with native subcanopy
habitat. This high recapture rate could be explained by the fact that
this habitat in effect was surrounded by habitat insuitable for
arboreal species. In essence, this habitat was a forested island. On
one side there was a monoculture of Eucalyptus saligna with no
subcanopy, on another a marsh turned into a rice field, and the other
two sides of the forest were bounded by pasture. Arboreal species
should spend more time in arborescent vegetation than on the ground.
These species should be more hesitant to disperse than terrestrial
ones.
Another obvious difference between this study and other
inventories conducted in neotropical native forests was the absence of
echymid rodents. Species of the genus Proechimvs are the most
widespread taxa of the family Echymidae in the neotropics
(Hershkovitz, 1969). These forest species are terrestrial and usually
appear on species lists from forest inventories. In Panama,
Proechimys was a common forest capture (Glanz, 1982; Eisenberg and
Thorington, 1973). Handley (1976) reported Proechimvs as a common
species in Venezuela. Emmons (1984) and Terborgh et al. (1986)
reported captures of Proechimvs from forested sites in Peru and
Ecuador. There are several reports of Proechimvs captures in
Brazilian tropical moist forests. Laemmert et al. (1946), Emmons
(1984), Carvalho (1965), Miles et al. (1981) and Malcolm (pers. comm.)

52
reported Proechimvs in their inventories from the Brazilian Amazon.
In the Atlantic Forest, Davis (1945) and Fonseca (pers. comm.)
reported two species of Proechimvs from their studies in the states
of Rio de Janeiro and Minas Gerais, respectively. Dias (1982) trapped
one species of Proechimvs in 3 study areas in the Rio Doce Valley.
I did not capture one individual of Proechimvs from the Park in
approximately 35,000 trap nights from 1985-86, nor from an additional
30,000 trap nights from 1986-87 (J. Stallings, in prep.). One
explanation could be the presence of predators in the Park.
Eisenberg (1980) speculated that the abundance of rodents in some
neotropical sites and the paucity of rodents in other sites could be
the result of the presence or absence of top predators. Hershkovitz
(1969) stated that species of the genus Proechimvs "are the basic
source of protein for lowland predators in the Brazilian subregion."
The felid community in the Park is intact. All of the felids have
been observed by field workers in the recent past. I saw spoor from
jaguar, puma, ocelot, Geoffroy’s cat, and jaguarundi.
In the wet meadow habitat my findings were consistent with the
literature (Table 2-12). In every other study, rodent captures were
higher than marsupial captures. In this study, Akodon cursor was the
dominant species in terms of absolute numbers and captures.
O’Connell (1981) reported that Zvqodontomvs was the dominant rodent
in grass habitat in Venezuela and represented 85% of the total rodent
captures. One difference between this study and others was the
observed high trapping success. As can be observed from Table 2-4,

53
Akodon cursor made up 85% of the captures and was responsible for
the high trapping success.
The eucalypt forest with native subcanopy habitat yielded
surprising results. I did not expect to find many small mammals in
this habitat because of "plantation effects." However, I captured 7
species of small mammals, most of these being marsupials. In fact,
the marsupial/rodent capture ratio was similar to that observed in the
native forested habitat (Table 2-12). I could not find any studies in
the literature to compare to the results from this habitat. Perhaps
the study that sampled homogeneous eucalypt forest would be the
most appropriate (Dietz et al., 1975). Dietz et al. (1975) captured
two species of terrestrial rodents, Orvzomvs nigripes and Akodon
cursor, in the homogeneous plantations with grass/bamboo
undergrowth. The authors caught a total of 5 species, only one of
which was not strictly terrestrial, in the two native forested habitats.
The plantation habitats in this and my study are similar in that they
both were eucalypt plantations of similar age and that the terrestrial
substrate of both was covered by grass. The major difference was
the native species subcanopy. I captured a relatively high number of
terrestrial rodents and marsupials and a high number of arboreal
marsupials. I did not capture any arboreal rodents. The native
species subcanopy could be considered a secondary forest sere, if the
emergent eucalypt stratum is ignored. Following Charles-Dominique’s
(1983) hypothesis of increased didelphid marsupial abundances in
secondary habitat, it is not surprising that I captured marsupials at a
rate consistent with the native forested habitat.

54
The temporal capture results suggested that there were two
systems operating in the Park. The trapping data showed a
pronounced peak in the total number of captures, mka and first
captures for the native forested habitat during the cool, dry winter.
Davis (1945) reported a similar trend in the trapping results and
suggested that this trend was the result of more younger individuals
present 1n the trapping pool or because of a paucity of natural food
items during this time of year. My data do not support the
hypothesis that more younger individuals explain the pronounced
increase; rather 1t appears that food resource paucity results in the
increase (Chapter III).
The eucalypt and wet meadow habitats showed similar trapping
trends with a peak in the cool, dry winter, and a peak in late
summer. These two habitats might have yielded similar capture curves
because of the grass substrate. Akodon cursor was an important
component of both habitats and is an insectivore/omnivore that is
reported to use a high proportion of grass and grass seed in its diet
(Nowak and Paradiso, 1983). Harmosa incana and Metachi rus
nudicaudatus are insectivore/omnivores and frugivore/omnivores,
respectively, and perhaps track insect availability in grass substrate.
The grass species did not produce seeds until late May. Thus,
perhaps the peak observed 1n February, March and April can
be explained by the lack of food for both rodents and marsupials.
The second peak, which occurred in June, July, and August, could
also be explained in terms of a decrease in food availability. Insect
and fruit availability are usually low during the hibernal period in

55
seasonal neotropical forests (e.g., Janzen and Schonener, 1968). The
marsupial species rely heavily on these food resources. The results
of a preliminary stomach content analysis on Akodon. suggest that
insects are important items in this species diet (Stallings, in prep.).
Graminoids in this habitat were dry and seeds were not as readily
available as they were during April and May.
The trap type data analysis revealed that terrestrial small
Sherman Uve traps were very productive in the wet meadow habitat
but yielded relatively few captures in the forested habitats. The
same results were obtained for arboreal small Shermans in the
forested habitats. Large terrestrial Uve traps were unproductive 1n
the forested habitats. The most productive trap types were the
medium sized terrestrial and arboreal traps and the arboreal platform
traps. Some individuals of species that are considered terrestrial
were captured in arboreal traps. This can be explained by some low
arboreal traps that were connected to the ground by either a vine or
log.
The use of the arboreal platform traps did not allow me to trap
additional species that were not already trapped using the terrestrial
and low arboreal traps. However, I was able to increase the
frequency of capture of the highly arboreal marsupial Caluromvs.
Malcolm (per. comm.) obtained similar results with platform traps in
Manaus. Perhaps the first use of arboreal platforms for trapping
small mammals were in the studies described by Davis (1945) and
Laemmert et al. (1946). Unfortunately, I could not determine the
success of these traps nor the species captured from these studies.

56
In total, I logged 49 captures of this Caluromvs in both the eucalypt
with native species subcanopy and the native forested habitats. In
the latter habitat, I only trapped this species 5 times in the
terrestrial and low arboreal traps. I trapped 29 Caluromvs in the
native forested habitat by using the arboreal platforms. I would have
underestimated the presence of this species had I not used the
platform traps. Marmosa cinerea was trapped also with relative high
frequency in this trap type.
I was surprised to find such a high trapping frequency of
Caluromvs in the eucalypt forest with native species subcanopy.
Although this species can be quite common in native species forested
habitats, I found it unusual that this highly arboreal frugivore would
be inhabiting an exotic monoculture plantation. There obviously was
sufficient food resources available from the native species subcanopy.
The fact that this species was captured in terrestrial and low
arboreal traps, without using platforms, suggests that this species was
using the subcanopy.
The results of this small mammal inventory suggest that
marsupials play an important role in the community structure of small
mammals in one of the largest remaining native tracts of Atlantic
forest in Brazil. Wet meadow habitat in this region is speciose in
rodents, and perhaps dominated by one or two species. Eucalypt
forests with native species subcanopy can play an Important role in
the conservation of small mammal communities in a region greatly
altered by monocultura! plantations.

CHAPTER III
TEMPORAL VARIATION IN TRAPPING SUCCESS OF
DIDELPHID MARSUPIALS IN AN EASTERN BRAZILIAN PARK
Introduction
Tropical rainforests are classified as stable and evergreen, with
little seasonal change in the flora and fauna (Richards, 1952; Smith,
1974). However, tropical seasonal and semideciduous forests exhibit
seasonal changes 1n floral phenology (e.g., Augspurger, 1982; Foster,
1982; Garwood, 1982) and consequently variation in faunal populations
and activities (e.g., Howe, 1982; Worthington, 1982; Smythe et al.,
1982; Glanz et al., 1982). These changes have been related to
variation in the abiotic environment, such as changes in photoperiod,
precipitation, and temperature (Sinclair and Norton-Griffiths, 1979;
Leigh et al., 1982).
Tropical forests that exhibit a pronounced dry season seem to
have a peak of fruiting at the onset of the wet season immediately
following the dry season (Foster, 1980). While the reasons for this
peak in the fruiting period are not clear, the result is that fruit is
more abundant at certain times of the year. This Increase in the
availability of fruit drastically affects the animals that use these
resources for food.
Population fluctuations interpreted by trapping results of small
mammals must be carefully analyzed to determine if the population is
57

58
actually fluctuating or if animals are responding to trap bait during
periods when food is available (Hunsaker, 1977). Previous research
on didelphid marsupials in the neotropics (Davis, 1945; O’Connell,
1979; Fleming, 1972, 1973; August, 1984) have found that trapping
success of marsupials increased during the hibernal and prevernal
months. This increase in trapping success has been attributed to the
paucity of food resources during this period. Another interpretation
of the results could be linked to reproduction, such as greater
movements of males or a large number of immatures. These might be
related to the initiation or the cessation of reproductive activity.
One objective of this paper is to test the null hypothesis that
trapping success of marsupials does not vary with season in the
Atlantic Forest of eastern Brazil. Another objective is to test the
null hypothesis that any peak in trapping success was a consequence
of an increase in the population or reproductive activities.
Materials and Methods
Small mammals were trapped from October, 1985, through
September, 1986, in the Rio Doce State Forestry Park, Minas Gerais
state, Brazil. The Park lies between the coordinates 19 48’18" and 19
29’24" south latitude and 42 38’30" and 42 28’18" west longitude. The
climate of the Park is classified as tropical humid with a rainy season
between the months of November and February and drought during
the months of June, July, and August. Mean annual temperature is
approximately 22 C, with mean minimum monthly temperatures
fluctuating greatly throughout the year (Figure 3-1). Both the mean

TEMPERATURE (C )
SUMMER SEASON MONTHS WINTER SEASON
MEAN MAXIMUM TEMP —*— MEAN MINIMUM TEMP
DATA FROM 1954 - 1974
Figure 3-1. Temperature graph showing both minimum and maximum
mean temperatures per month in the Rio Doce State Forestry Park,
Hinas Gerais, Brazil. Winter and summer seasons are indicated
along the X axis.

60
maximum and minimum monthly temperatures reach their respective
lows during the drought season.
For the purposes of this paper, I divided the year into two
seasonal periods. This division is based on the monthly mean
minimum temperature. If the mean monthly minimum temperature was
above 17 C. the month was classified as summer, otherwise it was
considered the winter period. A comparison of my seasonal
classifications and those of Davis (1945) shows thermal and intuitive
consistencies (Figure 3-1). My winter season corresponds to the
period from April - September and the summer season from October -
March. My classification corresponds to those of Davis in the
following manner: my winter season includes the autumnal, hibernal
and prevernal seasons, while my summer season includes the vernal,
aestival and serótina! seasons.
The phytogeographical domain of the Park has been classified by
several authors. For example, according to Ab Saber (1977, in
Gilhaus, 1986), the Park occurs within the Tropical Atlantic Domain.
Rizzini’s (1963, in Gilhaus, 1986) classification defines the Park’s
domain as the Atlantic Province, Austro-Oriental sub-province,
Cordilheira sector. Gilhuis (1986) correctly pointed out that both
classification schemes locate the Park in a transitional zone bordering
the more humid Tropical Atlantic Domain and the drier Cerrados
Domain. I found tree species from both domains to be present in the
Rio Doce Park. Trees are more deciduous than would be expected in
Tropical Atlantic Domain forest. Most of the emergent tree species
and some others in the upper strata lose their leaves during the cool

61
and dry hibernal and prevernal seasons (Gilhuis, 1986; Stallings, pers.
obs.). The forest of the park is classified as Tropical Semideciduous
by Alonso (1977, in Gilhaus, 1986), Tropical Broadleaved by Azevedo
(1969, in Gilhaus, 1986) and Tropical Pluvial Seasonal by Lima (1966,
in Gilhaus, 1986).
Small mammals were live-trapped at 5 forested sites within the
Park boundary. Additional trapping was conducted in and around the
Park and the methodology is described in Chapter II. For the
purposes of testing the above mentioned hypotheses, I recorded the
following observations on each captured individual. The initial
capture of each individual was a first capture and a numbered ear tag
was placed on the left ear of all animals to identify individuals that
were captured previously. Total captures were first captures plus
subsequent captures of the same individual. The minimum known
alive (MKA) were the number of individuals captured on the first
occasion and those individuals recaptured only on one occasion during
each trapping period. Sex, age and reproductive information was also
recorded.
The probability of capturing a small mammal is affected by 3
factors: 1) temporal effects, 2) behavorial effects, and 3)
heterogeneity effects (White et al., 1982). The effects of
heterogeneity (e.g., number and placement of traps in an animal’s
home range or variation due to social dominance) vary among animals,
but capture probabilities for each animal remain constant per
occasion. In contrast, temporal (e.g., variation over time because of
environmental conditions or trapping effort) and behavior (e.g., trap

62
avoidance or fascination) effects can alter capture probabilities after
the initial capture of an Individual.
First capture data are more affected by heterogeneity and
temporal effects than behavioral effects (White et al., 1982). An
increase in the number of juveniles in first capture data through time
suggests that the population is increasing. However, if young
individuals do not account for an increase in first capture data, then
an alternative explanation 1s that the increase reflects individuals
present in the population that were not previously captured.
Total capture and minimum known alive data are influenced by
temporal, behavioral, and heterogeneity effects. Total capture data
are the result of all captures per occasion or through time, inclusive
of first captures, successive recaptures, and recapture of individuals
separated by time. These data should not be interpreted as a good
estimator of population abundance because they are strongly affected
by behavioral and temporal factors. An increase in recapture data
through time can be interpreted as a change in the capture
probabilities of marked individuals due to trap fascination, rather
than trap shyness which would reduce capture probabilities.
The minimum known alive data are the best population estimator
of the three. These data incorporate first captures and only the first
recapture of an individual captured in a previous occasion for each
trapping period. This estimator, in effect, uses the number of new
captures plus survivorship of animals captured previously to obtain an
estimate of abundance.

63
I only used marsupial capture data to test the null hypotheses
that trapping success does not vary by season and that any expected
increase is due to reproduction. Marsupials comprised 83% of the
total small mammal captures and 80% of the first captures (Chapter
II). Marsupial species included in the analysis were Didelphis
marsupial is (azarae). Metachi rus nudicaudatus. Marmosa incana.
Marmosa cinerea. and Caluromvs philander. Fruit and insects are
important compontents of this group’s diet (Chapter II).
I used correlation analysis and simple regression to determine if
there was any association between trapping success and mean
minimum monthly temperature. I quantified trapping success of first
captures, minimum known alive and total captures for each season.
Chi-square tests for goodness of fit were then used to test for
significant deviation from expected capture frequencies between
seasons.
I looked at (1) an increase in the number of juveniles during
the winter season, (2) an increase in the number of males between
seasons, (3) a male biased sex-ratio during the winter season, and (4)
an increase in the number of lactating/pregnant females during the
winter season to test the null hypothesis of an expected increase due
to reproduction. For this analysis I used Chi-square tests goodness
of fit to test for significant deviation from the above expected
results.
Individuals were placed into either juvenile or adult subjective
age classes. Initially, the age class placement was based on the
overall morphological gestalt of each individual per species. I then

64
used the recorded body measurements (Chapter II) to determine the
threshold between the two age classes and I compared field
determined ages to ages derived from body measurements. Body mass
and hind foot measurements were relied on heavily to determine age
class.
Age class categories of the 5 species of marsupials are presented
in Table 3-1 and are taken from Chapter III. For Didelphis
marsuoialis. juvenile and adult age categories were based on the hind
foot size. This species can attain a body mass of 2 kg. The hind
foot would grow slower in comparison to smaller bodied species and
body mass would provide a better indicator of age. Animals with a
hind foot of 51 mm or smaller were classified as juveniles (average
weight 417 gm). Lengths greater than 51 mm (average weight 1049
gm) were classified as adults. I used body mass to determine the
threshold for Metachi rus nudicaudatus. Marmosa incana, and Marmosa
cincera. These species are relatively small and foot size reaches its
maximum at an early age. Body mass provided the better estimator
because mass increased as the individuals grew older. Metachlrus
nudicaudatus was placed into age categories based upon a threshold
weight of 90 gms. Age class placement was determined by a weight
of 35 gm and 50 gm for Marmosa incana and M. cinerea. respectively.
All Caluromys philander that were captured were considered adults.
The number of individuals of each sex were sorted for first,
MKA, and total capture data between and within seasons.
Comparisons were made for sex biased trapping success between

65
Table 3-1. Criteria used to place marsupials into either juvenile
or adult age categories. Individuals per species were classified
as juveniles if designated measurements were less than cut-of
measurements used in table. Species were not sexually dimorphic
based on body measurements listed.
SPECIES
BODY
MEASUREMENT USED
CUT-OFF
MEASUREMENT
DidelDhis marsuDialis
HIND FOOT
51 MM
Metachi rus nudicaudatus
MASS
90 G
Marmosa incana
MASS
35 G
Marmosa cinerea
MASS
50 G

66
seasons for all capture data. In addition, comparisons were made for
sex biased trapping success within seasons.
Female marsupials were examined to determine if they were
lactating and for the presence of young attached to the teat field.
Females with young attached were considered pregnant. Frequencies
of lactating/pregnant females were compared on a seasonal basis.
Results
There was a significant negative association observed between
mean monthly temperature and trapping success through time. This
association is significant only if we look at the number of total
captures (f=28.444, p < .0003) (Figure 3-2) or the minimum known
alive (f=9.939, p < .007) (Figure 3-3). First capture data did not
show a significant association (Figure 3-4).
The first capture data (0.05 0.001) and total captures (p< 0.001) of marsupials were greater than
expected in the winter season than in the summer season (Table 3-2).
Juveniles were observed throughout the year. Juveniles were
represented from four of the five species of marsupials (Table 3-3).
There was no significant difference in the minimum known alive or
first captures of juveniles between seasons. A significant difference
(p < .001) was observed in the total captures of juveniles between
seasons. More total captures of juveniles were observed in the
winter season than expected.
The number of juveniles captured per species between seasons
was fairly constant (Table 3-3) from minimum known alive and first

67
P BET
TOTAL
NUMBER OF CAPTURES
Figure 3-2. Relationship between temperature and total captures
of didelphid marsupials in the Rio Doce State Forestry Park,
Minas Gerais, Brazil.

NUMBER OF CAPTURES
i
Figure 3-3. Relationship between temperature and minimum known
alive captures (MKA) of didelphid marsupials in the Rio Doce
State Forestry Park, Minas Gerais, Brazil.

69
NUMBER OF CAPTURES
0 5 10 15 20
TEMPERATURE (C )
Figure 3-4. Relationship between temperature and first captures of
didelphid marsupials in the R1o Doce State Forestry Park, Minas
Gerais, Brazil.

70
Table 3-2. Comparison of the first captures, minimum known alive
(MKA) and total captures of five species of didelphid marsupials
between seasons in the Rio Doce State Forestry Park, Minas Gerais,
Brazil. * Significant at 0.05-0.01. ** Significant at 0.01—
0.001.
SEASONS
WINTER
SUMMER
CHI-SQUARE
FIRST CAPTURES
206
151
4.23*
MKA CAPTURES
276
157
16.35**
TOTAL CAPTURES
392
190
35.05**

71
Table 3-3. The number of first captures (FIRST), minimum known
alive (MKA) and total captures (TOTAL) of juvenile didelphid
marsupials per season. N.S= Non significant.
SUMMER SEASON WINTER SEASON
FIRST MKA TOTAL FIRST MKA TOTAL
SPECIES
D. marsupial is
M. nudicaudatus
M. incana
M. cinerea
TOTALS
FIRST CAPTURES X2
MKA CAPTURES X2
TOTAL CAPTURES X2
3
8
10
7
5
14
15
17
26
8
10
19
33
40
69
: 0.00,
N.S.
: 0.55,
N.S.
14.38,
P < 0.
001
6
6
7
5
5
21
14
18
35
8
21
85
33
50
148

72
capture data. The total capture data per species indicated that there
was a recapture bias during the winter season, although the
difference was not significant. Figure 3-5 shows that first captures
of adults, not juveniles, clearly represented the majority of captures
during May, June, and July.
The number of captures of each sex between seasons is
presented in Table 3-4. There were no significant differences
between the number of males captured between seasons or the number
of females captured between seasons using the first capture data.
Minimum known alive and total captures data showed significant
differences in the expected number of males or females between
seasons. More males and more females were captured during the
winter season than during the summer season, with a higher
proportion of males being captured during the winter season. For the
first capture data, minimum known alive and total capture data there
were no significant differences in the male/female sex ratio within
each season (Table 3-5).
More lactating females, whether carrying young or not, were
captured during the summer season than the winter season (Table
3-6). However, this difference, although approaching significance,
was not statistically significant (x2 =2.89; .10 < p <.05). One
striking observation was the obvious decrease in the occurrence of
lactating/pregnant females during the months of June, July and
August (Figure 3-6). This decrease is also observed when comparing
number of juveniles and lactating females per species through time
(Figures 3-7, 3-8, and 3-9).

>
73
RIpQ
inilO'Q)
OF ADULTS AND YOUNG BY MONTH
NUMBER OF CAPTURES
MONTHS
cm NO. YOUNG GO NO. ADULT
Figure 3-5. Comparison of first captures of adult and young
didelphid marsupials per month in the Rio Doce State Forestry Park,
Minas Gerais, Brazil.

74
Table 3-4. Between season comparisons of the number of first
captures (FIRST), minimum known alive (MKA) and total captures
(TOTAL) of didelphid marsupials per sex. * Significant at 0.05-
0.01. ** Significant at 0.01-0.001.
BETWEEN SEASON COMPARISON
SUMMER SEASON WINTER SEASON
X2
NUMBER OF MALES
FIRST CAPTURES
62
83
1.52
MKA CAPTURES
78
152
*
*
o
o>
f—
TOTAL CAPTURES
97
215
22.30**
NUMBER OF FEMALES
FIRST CAPTURES
66
70
0.05
MKA CAPTURES
79
124
4.98*
TOTAL CAPTURES
92
174
12.63**

75
Table 3-5. Within season comparison of the number of first
captures, minimum known alive captures (MKA) and total captures of
didelphid marsupials per sex. N.S.= Non significant. Significance
level set at p < 0.05.
SEX
X2
SUMMER SEASON
MALES
FEMALES
FIRST CAPTURES
62
66
0.06 N.S.
MKA CAPTURES
78
79
0.003 N.S.
TOTAL CAPTURES
97
92
0.06 N.S.
WINTER SEASON
MALES
FEMALES
FIRST CAPTURES
83
70
0.55 N.S.
MKA CAPTURES
152
124
1.42 N.S.
TOTAL CAPTURES
215
174
2.16 N.S.

76
Table 3-6. The frequency of lactating/pregnant didelphid
marsupials trapped per season in the Rio Doce State Forestry Park,
Minas Gerais, Brazil. N.S.= Non Significant.
SPECIES SEASONS
SUMMER WINTER
D. marsupial is 4 1
M. nudicaudatus 11 12
M. incana 9 0
M. cinerea 12 3
Caluromvs philander 1 3
TOTALS 37 19

CO h- CD lO ^ CO CM
77
NUMBER
NOV DEC JAN FEB MAR APR MAY JUN JUL AUG SEP OCT
MONTHS
EB Dldelphls Metachirua üMl M. incana
EiüH M. clnerea 1... I Caluromys
Figure 3-6. Comparison of lactating females of five species of
didelphid marsupials per month in the Rio Doce State Forestry Park,
Minas Gerais, Brazil.

78
NUMBER
M. nudicaudatus
Figure 3-7. Comparison of the number of captures of young and
lactating females of the didelphid marsupial Metachi rus
nudicaudatus in the Rio Doce State Forestry, Minas Gerais, Brazil.

79
NUMBER
M. incana
Figure 3-8. Comparison of the number of captures of young and
lactating females of the didelphid marsupial Marmosa incana in the
Rio Doce State Forestry Park, Minas Gerais, Brazil.

80
[QUENGY OF YOUNG
AND LACTATING FEMALES
NUMBER
MONTHS
CZ1 YOUNG IE3 LACTATING FEMALES
M. cinerea
Figure 3-9. Comparison of the number of captures of young and
lactating females of the didelphid marsupial Marmosa cinerea in the
Rio Doce State Forestry Park, Minas Gerais, Brazil.

81
Discussion
There is obviously a connection between animal population
numbers and the availability of food resources (Leigh et al; 1982).
An intuitive model is that population increases should track and peak
with maximum production of food resources. By timing reproduction
prior to the peak of available food resources, individuals maximize
their opportunities for the survival of their offspring.
There was a clear peak in the trapping success of marsupials
during the winter months in the Rio Doce State Forestry Park.
There was a significant negative association between the number of
total captures or minimum known alive each month and mean
minimum monthly temperature. However, the first capture data did
not support these previous findings. These data suggest that the
recapture of individuals plays an important role in the trapping
success numbers and that the numbers of new individuals did not
increase during the winter season.
August (1984) found an increase in the capture success of
Marmosa robinsoni during the dry season in Venezuela, while
O’Connell (1979) reported an increase in the minimum known alive for
Marmosa robinsoni. M. fuscata. and Didelphis marsupial is during the
same period in Venezuela. Fleming (1972) observed an increase in
the number of captures of three species of marsupials during the dry
season in Panama. Davis (1945) reported more captures and
recaptures of two species of didelphid marsupials, Didelphis

82
marsupialis and Caluromvs philander, during the dry, cold season in
Teresopolis, Brazil.
Research on neotropical marsupial mating behavior suggests that
typically males are more active prior to and during the reproductive
or mating season (Atramentowicz, 1982; Hunsaker, 1977). This may be
a consequence of the polygynous mating systems observed for these
small mammals (Eisenberg, 1981). There was no significant difference
between seasons in the number of first captures of male marsupials.
The same results were obtained for female first captures. There were
more male and female minimum known alive captures during the
winter season than the summer season. The same results were
obtained for the total captures of male and female marsupials during
the winter season, but at a higher level of significance. These data
again suggest that there is a recapture bias in the trapping data, and
that observed and significant differences in the number of captures
and individuals of each sex between seasons is a function of being
recaptured and not related directly to reproduction. However, the
minimum known alive and total capture data show male captures in
the winter season to be more significant than females captures during
the same period. These findings could be interpreted as increased
male activity over female activity at the onset of the reproductive
season.
A direct measure of population growth is the number of young
observed at any point in time. My results indicate that during the
winter season the number of young individuals captured was not
greater than in the summer season, but more young individuals were

83
recaptured during the this season than in the summer season.
O’Connell (1979) reported an increase in the number of individuals of
Didelphis marsupial is during the wet season in Venezuela. She
attributed this increase to the first captures of juveniles.
The last measure that was used to determine population growth
was that of the occurrence of lactating/pregnant females through
time. Statistically, there was no difference in the occurrence of
reproductively active females between seasons, although there was a
suggestion of more reproductively active females in the summer
season than in the winter season. O’Connell (1979) found that most
of the species she studied to be sexually inactive during the dry
season.
It is clear from the previous analyses that there was a
pronounced increase 1n the number of animals known to be alive and
the overall trapping success during the hibernal and prevernal seasons
(Figure 3-10). However, the data do not support the hypothesis that
this significant increase is due to more juveniles present in the
population, nor does it result from increased male activity or more
lactating/pregnant females. I conclude that population growth of
marsupials was constant if we consider only first capture data.
Clearly, the trapping success curve was being driven by adult
individuals; individuals that were being recaptured more during a
certain period in time than at another.
I did not measure the availability of fruit or insects during the
study. Phenological studies (CETEC, 1981) in the park from 1977-
1981 found that trees were in flower and fruit throughout the year,

NO. CAPTURES
200
150 -
100 -
OCT-NOV DEC-JAN FE8-MAR APR-MAY JUN-JUL AUG-SEP
MONTHS GROUPED ACCORDING TO SEASON
SEASONS GROUPED BY MONTHS (DAVIS, 1945)
Figure 3-10. Frequency of captures of didelphid marsupials by
season in the Rio Doce State Forestry Park, Minas Gerais, Brazil.

85
with a noticeable, but not statistically significant increase from
October through December (Figure 3-11). These studies also indicated
that the months of June, July and August represented the lowest
incidence of tree species with flowers and fruits. There was also a
low incidence of tree species with flower and fruits in January.
There are no quantitative data available on the seasonality of insect
density in the park. However, studies in other Neotropical areas
report a positive association between increased insect abundance and
precipitation and temperature (Janzen and Schonener, 1968; Wolda,
1978, in August, 1984; Davis, 1945). It is possible that such a trend
occurs in the park due to the high seasonality of rainfall and
temperature.
In other areas were neotropical didephids have been studied
(Fleming, 1972; O’Connell, 1979; August, 1984; Charies-Dominque,
1983), the dry season represented the period of the year when food
resources were scarce. In contrast, the three month period from
October - December in the park showed more species in fruit and
flower than any other three month period. These trends are
interesting when compared to the low and peak three month trapping
success periods. The highest trapping success coincides with the
lowest number of tree species in fruit and flower and likewise the
lowest period of trapping success coincides with the maximum period
of fruit and flower production.
However, it must be stated that trapping success remained
relatively constant for the period of December and January when the
number of tree species with fruit and flower were observed to be

86
ENCY OF FRUIT
AND FLOWERS
NO. SPECIES IN FRUIT
SEASONAL PERIODS IN MONTHS
Figure 3-11. Frequency of species of trees and shrubs in fruit and
flower per seasonal period in the Rio Doce State Forestry Park,
Minas Gerais, Brazil.

87
low. This two month period has the highest mean temperature and
precipitation of the year. If insect density is high at this time, then
perhaps marsupials are able to switch to insect prey or other foods
in response to the unavailability of fruit.
I would interpret my results of a significant increase in total
trap success as a response to the lack of fruit during the subtropical
winter and not due to (1) greater movement of males (i.e., more
captures of males than females during this period) nor (2) a
significant increase 1n the number of juveniles captured nor (3)
because of more reproductively active females during this period.
Marsupials fell into traps easier during this period because they were
searching for food. This interpretation is consistent with the
findings of Atramentowicz (1982), Charles-Dominique et al. (1981),
Charles-Dominique (1983), O’Connell (1979), Fleming (1972, 1973),
Davis (1945) and August (1984) and suggests that reproduction and
weaning of young marsupials occur at the end or after the dry season
when food resources are more abundant.

CHAPTER IV
SMALL MAMMAL ASSOCIATION AND MICROHABITAT
SELECTION IN AN EASTERN BRAZILIAN PARK
Introduction
There are several studies on small mammal habitat selection in
temperate zones (e.g., Dueser and Porter, 1986; Dueser and Shugart,
1978, 1979; Hallett, 1982; Rosenzweig and Winakur, 1969), but there
are relatively few small mammal habitat studies conducted in the
neotropics. August (1983, 1984) studied the relationship between
small mammals and habitat structure in Venezuela. Lacher and Alho
(in press) and Lacher et al. (1988) reported habitat selection by small
mammals in grassland habitats in southern Brazil. Nitikman and
Mares (1987) reported microhabitat preferences of small mammals in a
gallery forest in central Brazil. The purpose of this paper is to
examine the effect of habitat structure on habitat use by small
mammals in an eastern Brazilian forest, the Rio Doce State Forestry
Park.
Preliminary analysis of the habitat in this Park suggested that
there are distinct differences in the forest structure across several
forested habitats (Gilhaus, 1986). Forest fire, in the form of intense
crown fire, has played an important role, at least in the recent past,
in structuring the forest. This study tested the hypothesis that small
88

89
mammal abundances vary with habitat and that species select
microhabitats within each habitat.
Materials and Methods
This study was conducted in the Rio Doce State Forestry Park
(Rio Doce), Minas Gerais, Brazil (19 48’ 18“ and 19 29’ 24“ south
latitude and 42 38’ 30" and 42 28’ 18" west longitude). The Park
contains over 35,000 ha and elevation ranges from 230 to 515 m. The
mean annual precipitation for the Park was 1480 mm. from 1954 to
1974, however annual precipitation during the study amounted to 947
mm. (Figure 4-1). Mean monthly maximum and minimum temperatures
vary appreciably throughout the year (Figure 4-2).
Study Sites
There are several distinct forested and open/field habitats in the
Park (Gilhaus, 1986). Slope, soil quality and moisture, and elevation
all affect habitat type. However, forest fire has played the major
role affecting the forested habitat in this Park. The vegetation of
the Park is classified as tropical semi-deciduous and most of the
emergent tree species lose their leaves during the cool dry months.
In 1964 and 1967, major forest fires burned approximately 30% of the
forest (Lopes, 1982; Silva-Neto, 1984). Fire is important because leaf
Utter accumulates during the dry season.
During the course of this study, I surveyed small mammal
communities in 5 distinct forested sites which represent 3 habitat
types in the Park (Figure 4-3). The following is a brief description
of each study site.

90
WALTER AND LEITH
CLIMATIC DIAGRAM
MONTHLY RAINFALL (MM.) MEAN TEMPERATURE (0 )
MONTHS
AVERAGE ANNUAL RAJNFALL U0O mm
OAJA FROM 1664-1074
WALTER AND LEITH
CLIMATIC DIAGRAM
MONTHLY RAINFALL (MM) MEAN TEMPERATURE (O I
JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV OEO
MONTHS
ANNUAL AVERAGE RAINFALL 804 mm
DATA FROM mm
A.
5.
Figure 4-1. Walter and Leith climatic diagram characterizing
surplus precipitation and drought by month in the Rio Doce State
Forestry Park, Minas Gerais, Brazil.

TEMPERATURE (C )
MONTHS
MEAN MAXIMUM TEMP —'— MEAN MINIMUM TEMP
DATA FROM 1954 - 1974
Figure 4-2. Mean minimum and maximum monthly temperatures in the
Rio Doce State Forestry Park, Minas Gerais, Brazil.

92
RIO DOCE STATE
FORESTRY PARK
3 4 km
Figure 4-3. Map of the Rio Doce State Forestry Park, Minas
Gerais, Brazil. Circled numbers indicate trapping sites. The
five native forested trapping sites are indicated by number 2,
and numbers 4 through 7.

93
Low Secondary Habitat (RD/F). This forest type is equivalent to the
Medium Secondary Forest with Bamboos and Graminoids described by
Gllhaus (1986). This forest type is characterized by a mean forest
canopy height of less than 8 m, a relatively high number of tree and
shrub species, and high relative abundances of trees and shrubs, with
small diameters at breast height. The trapping site was burned
completely to the ground in 1967 and the resulting vegetative cover
is the result of succession since the fire.
Mosaic Forest Habitat (RD/M). This forest type corresponds to the
Medium to Tall Forest with Bamboos and Graminoids described by
Gilhaus (1986). This forest type has a variable canopy height, but
usually more than 12 m. This formation is intermediate between
primary and early secondary forest, with variable tree and shrub
density and height. This trapping site was burned in a mosaic
fashion in 1967. Some areas were burned to the ground whereas
other areas were not affected by the forest fire. A stream, which
drained a swamp located to the west of the site, cut through part of
this trapping area.
Mosaic Forest Habitat (RD/H). This forest type also corresponds to
the Medium to Tall Forest with Bamboos and Graminoids. The
trapping site was burned in a mosaic fashion in 1967. The canopy is
composed of both tall and short trees. Some open areas support a
dense herbaceous growth.

94
Primary Forest Habitat (RD/T). This forest type corresponds to the
Tall Primary Forest with Epiphytes (Gilhaus, 1986). This forest type
is characterized by a well stratified primary forest with a significant
amount of large diameter and tall trees. The trapping site occurs in
the vicinity of the Rio Turvo, but does not constitute part of a
gallery forest habitat. This forest has not been altered by forest fire
and the canopy reaches approximately 30 m. Arboreal epiphytes and
ground ferns are common.
Primary Forest Habitat (RD/C). This forest type also corresponds to
the Tall Primary Forest with Epiphytes. The trapping site occurs
near the Rio Doce, but 1t also does not form part of the gallery
forest. Forest site RD/C was not affected by the forest fires of 1964
or 1967 and supports a tall canopy. Arboreal epiphytes and ground
ferns are common.
Trapping Methods
I used three trapping lines, 300 m in length, in each trapping
site. There were 16 trapping stations per line, each separated by 20
m. At each post, I placed one squirrel-size wire live trap on the
ground. I placed arboreal and terrestrial Sherman live traps, as well
as arboreal squirrel-size wire live traps, systematically along the
trapping lines. Large wire live traps were placed on the ground at
every other station on the exterior lines. I used arboreal platforms
to sample the canopy dwelling community in trapping sites that
afforded suitable tall and developed canopies. Twelve arboreal

95
platforms with squirrel-size traps were placed in RD/C, RD/T, and
RD/H, while only 6 were placed in RD/M. Traps were open in each
trapping site for 5 consecutive nights per month. Trapping
commenced in November, 1985, and ended in October, 1987. Only the
results of the first 12 months are reported here.
Traps were baited with chunks of pineapple, dry oatmeal, and
cotton balls soaked with a mixture of sardines and cod-liver oil. The
following data were recorded upon the capture of a small mammal:
species, sex, ear tag number, reproductive condition, age, mass, body
length, tail length, hind foot and ear length. All animals were
released at the trapping station where captured and behavior upon
release was noted.
Only species that had at least 20 total captures or at least 15
first captures were used in the abundance, association and
microhabitat analysis. These species are Didelphis marsupial is.
Metachi rus nudicaudatus. Marmosa incana. Marmosa cinerea.
Caluromys philander. Akodon cursor. Oecomys trinitatis. and Orvzomvs
capito. The ecological role of each species is presented in Table 4-1.
Several authors (e.g., Hallett, 1982; Dueser and Porter, 1986;
Lacher and Alho, in press; Lacher et al.; in prep) have examined the
effects of habitat structure and competition on small mammal
communities. I will not examine the effect of competition and
habitat selection because of the variety of feeding guilds and
adaptations for the use of vertical space exhibited by the species
involved. These two considerations assume that food and spatial
resources are shared and limited. I did not measure the amount of

96
Table 4-1. Ecological role of eight species of small mammals
captured at Rio Doce State Forestry Park, Minas Gerais, Brazil.
GM = grasslands and wet meadows; B = brushy areas; S = secondary
forests; P = primary forests; T = terrestrial; S = scansorial; A =
arboreal; FG = frugivore/granivore; FO = frugivore/omnivore; 10 =
insectivore/omnivore. G.= mass in grams.
SPECIES
HABITAT
TYPE
USE OF
VERTICAL SPACE
DIET
MASS
MARSUPIALS
DidelDhis marsuoialis
B,S,P
S
FO
1000
Metachi rus nudicaudatus
S,P
T
10
258
Mamosa cinerea
B,S,P
A
10
104
Marmosa incana
B,S,P
S
10
63
Caluromvs philander
S,P
A
FO
188
RODENTS
Oecomvs trinitatis
S,P
S
FG
71
Oryzomvs capito
S,P
T
FG
59
Akodon cursor
GM,B,S
T
10
40
/

97
overlap and availability of food resources. Moreover, since I did
measure the forest structure, I will concentrate on the small mammal
associations and abundances and microhabitat selection, rather than
competition coefficients.
Habitat Methods
I used both quantitative and qualitative measurements to
describe the microhabitat at each trapping station in each site (Table
4-2). Each variable described the physical environment in the
immediate vicinity of the trapping station, and provided a measure of
the forest structure that influences the distribution and abundance of
small mammals (see Dueser and Shugart, 1978). Most of the habitat
measurements were recorded within a circle with a radius of 3.5 m
centered at the trapping station. Logs, ferns, palms, bamboo, snags
and stumps were measured within the 3.5 m radius circle as well as
outside of the circle in the immediate area of the trapping station.
All other measurements were made within the trapping circle.
All analyses were performed using programs in Statistical
Analysis System on a personal computer (SAS, Institute, Inc.; 1985).
Statistical significance was set at p < 0.05.
Results
Trapping Site Habitat
To distinguish habitat types, I conducted one way analyses of
variance for each of the 28 variables to identify habitat variables
that were significantly different between each habitat and a pooled

98
Table 4-2. Microhabitat variables (with corresponding mnemonic
abbreviations in parentheses) measured at each trapping station, in
all habitats in the Rio Doce
Brazil.
State Forestry Park, Minas Gerais,
HABITAT VARIABLE
DESCRIPTION
NUMBER OF TREE FAMILIES (NOFAM)
NUMBER OF TREE FAMILIES IN DBH
SAMPLE
NUMBER OF TREE SPECIES (NOSPP)
NUMBER OF TREE SEPCIES IN DBH
SAMPLE
NUMBER OF TREES (NOTREE)
NUMBER OF TREES IN DBH SAMPLE
MEAN DIAMETER AT BREAST HEIGHT
(XBDH)
MEAN DIAMETER (CM) OF ALL TREES
WITH AT LEAST A DBH OF 3.2 CM
WITHIN A 3.5 M RADIUS OF TRAP
STATION
MEAN TREE HEIGHT (XHT)
MEAN TREE HEIGHT (M) OF ALL
TREES IN DBH SAMPLE
CANOPY HEIGHT (CANHT)
CANOPY HEIGHT (M) WITHIN 3.5 M
RADIUS OF TRAP STATION
HERBACEOUS HEIGHT (HERBHT)
AVERAGE HEIGHT (M) OF
HERBACEOUS VEGETATION WITHIN
3.5 M RADIUS OF TRAP STATION
CANOPY CONNECTIVITY (CANCON)
DEGREE OF CONNECTIVITY OF
CANOPY TREES (SLIDING SCALE
0-3)
MIDSTORY CONNECTIVITY (MIDCON)
DEGREE OF CONNECTIVITY OF
SUBCANOPY TREES (SLIDING SCALE
0-3)
PERCENT CANOPY COVER (PERCAN)
PERCENTAGE OF CANOPY COVER
ABOVE TRAPPING STATION
PERCENT HERBACEOUS COVER
(PERHERB)
PERCENTAGE OF HERBACEOUS GROUND
COVER WITHIN TRAPPING CIRCLE
CANOPY VOLUME (CANVOL)
CANOPY VOLUME ABOVE TRAPPING
CIRCLE EXPRESSED IN SLIDING
SCALE (0-3)

99
Table 4-2 - continued
HABITAT VARIABLE
DESCRIPTION
MIDSTORY VOLUME (MIDVOL)
MIDSTORY VOLUME ABOVE TRAPPING
CIRCLE EXPRESSED IN SLIDING
SCALE (0-3)
HERBACEOUS VOLUME (HERBVOL)
HERBACEOUS VOLUME OF GROUND
COVER WITHIN TRAPPING CIRCLE
VINE DENSITY (VINE)
DENSITY OF VINES EXPRESSED IN
SLIDING SCALE (0-3) IN
IMMEDIATE VICINITY OF TRAPPING
STATION
EPIPHYTE DENSITY (EPIP)
DENSITY OF VASCULAR EPIPHYTES
IN SLIDING SCALE (0-3) IN
IMMEDIATE VICINITY OF TRAPPING
STATION
RELIEF (RELIEF)
VERTICAL RELIEF AT TRAPPING
STATION SLIDING SCALE (0-3)
SOIL DEPTH (SOILD)
AVERAGE SOIL DEPTH (CM) AT
TRAPPING STATION
SOIL CONDITION (SOILC)
SOIL MOISTURE AT EACH STATION
IN SLIDING SCALE (0-3)
NUMBER OF LOGS (NLOGS)
NUMBER OF LOGS WITHIN IMMEDIATE
VICINITY OF TRAPPING CIRCLE
HUMUS (HUMUS)
RELATIVE AMOUNT OF HUMUS
EXPRESSED IN SLIDING SCALE
(0-3)
FERN DENSITY (FERNS)
DENSITY OF TERRESTRIAL FERNS
IN IMMEDIATE VICINITY OF
TRAPPING CIRCLE IN SLIDING
SCALE (0-3)
PALM DENSITY (PALBRE)
DENSITY OF PALMS IN IMMEDIATE
VICINITY OF TRAPPING CIRCLE
IN SLIDING SCALE (0-3)
BAMBOO DENSITY (BAMBO)
DENSITY OF BAMBOO IN IMMEDIATE
VICINITY OF TRAPPING CIRCLE
IN SLIDING SCALE (0-3)

100
Table 4-2 - continued
HABITAT VARIABLE
DESCRIPTION
BANANEIRA DENSITY (BANA)
DENSITY OF BANANEIRAS IN
IMMEDIATE VICINITY OF TRAPPING
CIRCLE IN SLIDING SCALE (0-3)
SNAG DENSITY (SNAGS)
PRESENCE OR ABSENCE OF SNAGS
IN IMMEDIATE VICINITY OF
TRAPPING CIRCLE
STUMP DENSITY (STUMP)
PRESENCE OR ABSENCE OF STUMPS
IN IMMEDIATE VICINITY OF
TRAPPING CIRCLE
SUM OF BASAL AREA (SUMBAS)
SUM OF BASAL AREA (CM2) OF ALL
TREES IN DBH SAMPLE

101
sample for all other habitats. Each analysis was arranged as a two
sample comparison between all observations for a particular trapping
site versus all observations for the other 4 trapping sites combined.
The 5 trapping sites or habitat types were different in overall habitat
structure and physignomy (Table 4-3). There were 21 univariate
habitat variables found to be significantly different among trapping
sites. The number of significant variables per site ranged from 5 to
13 variables. The paucity of significant variables in trapping sites
RD/H and RD/M, the sites burned in a mosaic fashion, reflects the
generality of the habitat compared to the more homogeneous sites,
RD/C, RD/T and RD/F. Only one habitat variable, arboreal
epiphytes, was found to differ significantly across all habitats or
trapping sites and pooled samples.
Since many of the habitat variables are significantly correlated
with other variables, I used canonical discriminant analysis because it
accounts for correlations between variables and detects and quantifies
differences between sample groups from multivariate data. For each
trapping site, discriminant analyses were performed with the
significant habitat variables (e) identified by ANOVA in Table 4-3.
For each test, N=240 (total stations across 5 trapping sites), g=2 (a
trapping site versus a pooled sample), e_= 5, 8, 9, 11, and 13
significant habitat variables for trapping sites RD/M, RD/H, RD/T,
RD/C and RD/F, respectively. MANOVA F tests for overall
discrimination of habitat variables were performed between each
trapping site and pooled sample means. The results of the MANOVA
overall discrimination test were highly significant for each trapping

102
Table 4-3. Microhabitat variables and significance levels for
comparisons of each site to a pooled sample including all
observations combined for the other sites. (DF 1/254) * p < 0.01
** p < 0.001. Mnemonic abbreviations explained in Table 4-2.
Trapping site abbreviations explained in text.
MICROHABITAT VARIABLES
TRAPPING SITES
RD/F
RD/H RD/M RD/T
RD/C
NOFAM
**
**
NOSPP
*
*
XDBH
*
CANHT
**
*
**
**
HERBHT
**
CANCON
**
**
**
PERHERB
**
**
CANVOL
**
*
**
VINE
**
**
EPIP
**
**
**
*
**
RELIEF
tt
**
**
SOILC
**
**
**
FERNS
**
**
**
**
HERBVOL
**
BAM BO
**
**
**
PERCAN
*
BANA
*
*
NLOG
**
STUMP
**
NOTREE
**
PALBRE
**
TOTALS
13
8
5
9
11

103
Table 4-4. Correlations with the discriminant function and mean
vectors for the microhabitat variables which each site differed
significantly (p < 0.01) from a pooled sample consisting of all
observations combined for the other sites. Mnemonic microhabitat
variables explained in Table 4-2.
SAMPLE MEAN VALUE
MICROHABITAT VARIABLES (R) SITE POOLED
TRAPPING SITE RD/H
NOFAM
-0.410
4.04
5.03
NOTREE
-0.461
5.91
7.06
NOSPP
-0.385
4.50
5.55
CANHT
-0.720
14.94
18.19
EPIP
-0.518
0.18
0.83
PALBRE
+0.250
1.41
0.81
BAMBO
-0.155
0.04
0.47
BANA
+0.143
0.58
0.30
MANOVA OVERALL DISCRIMINATION TEST
F= 15.4067
DF= 8/231
P < 0.0001
TRAPPING SITE RD/F
NOFAM
-0.204
5.75
4.60
NOSPP
-0.156
6.19
5.14
XDBH
+0.200
8.55
11.00
CANHT
+0.594
11.02
19.17
HERBHT
+0.466
0.66
0.98
CANCON
+0.257
0.48
1.01
PERHERB
+0.961
21.83
43.31
CANVOL
+0.292
0.56
1.14
VINE
-0.529
2.79
1.51
EPIP
+0.537
0.00
0.89
RELIEF
-0.570
1.71
0.45
SOILC
+0.556
1.00
1.95
FERNS
+0.565
0.25
0.98
MANOVA OVERALL DISCRIMINATION TEST
F= 28.8941
DF= 13/226
P < 0.0001

104
Table 4-4 - continued
SAMPLE MEAN VALUE
MICROHABITAT VARIABLES (R1 SITE POOLED
TRAPPING SITE RD/C
CANHT
+0.429
21.71
16.54
CANCON
+0.255
1.35
0.79
PERHERB
+0.987
64.72
37.58
CANVOL
+0.251
1.48
0.91
HERBVOL
+0.651
1.92
1.34
VINE
-0.460
1.15
1.92
EPIP
+0.603
1.98
0.39
RELIEF
-0.513
0.04
0.87
SOILC
+0.440
2.00
1.71
FERNS
+0.638
1.83
0.58
BAMBO
-0.165
0.02
0.48
MANOVA OVERALL
DISCRIMINATION TEST
F= 25.6164
DF= 11/228
P < 0.0001
TRAPPING SITE
RD/T
CANHT
-0.699
21.56
16.54
CANCON
-0.106
0.52
0.99
PERCAN
-0.782
63.02
55.94
CANVOL
-0.219
0.75
1.09
EPIP
-0.326
1.08
0.61
RELIEF
+0.392
0.02
0.88
SOILC
-0.339
2.00
1.71
FERNS
-0.274
1.21
0.74
BANA
+0.090
0.15
0.41
MANOVA OVERALL DISCRIMINATION TEST
F= 12.2862
DF= 9/230
P < 0.0001

105
Table 4-4 - continued
SAMPLE MEAN VALUE
MICROHABITAT VARIABLES (R) SITE POOLED
TRAPPING SITE RD/M
EPIP
-0.213
0.29
0.81
NLOG
+0.447
1.42
0.81
FERNS
-0.663
0.04
1.03
BAMBO
+0.864
1.37
0.14
STUMPS
+0.217
0.15
0.02
MANOVA OVERALL DISCRIMINATION TEST
F= 61.6361
DF= 5/234
P < 0.0001

106
site and the pooled samples (Table 4-4). Trapping site RD/H was
characterized by having lower tree diversity and number of trees,
lower canopy, lower incidence of epiphytes and bamboo, but more
bananeiras than the pooled sample. Trapping site RD/M was
characterized by having lower incidence of epiphytes and ferns and
more logs, bamboo and stumps than the pooled sample. RD/F can be
described as rich in the number of tree families and species, with
short, small diameter trees, a short open canopy and herbaceous
stratum, few epiphytes and ferns, high vine density, and moderate
relief. RD/C and RD/T, the primary forest trapping sites, were
similar in forest structure. These sites had a high and well
developed canopy and herbaceous stratum. Vine and bamboo densities
were low. The terrain was relatively flat. Terrestrial fern and
arboreal epiphyte densities were high compared to the pooled samples.
Small Mammal Abundances and Spatial Overlap
I examined the interactions of the small mammal species at each
trapping station within each site and among the sites. I used the
log-likelihood ratio goodness of fit test to test the null hypothesis
that each small mammal species used trapping sites in proportion to
their abundances. Except for Metachi rus nudicaudatus and Mamosa
cinérea, none of the small mammal species used the habitat in
proportion to their abundances (Table 4-5, log-likelihood ratio
goodness of fit test [Sokal and Rohlf, 1981]). These data suggest
that Metachi rus and M. cinerea used the habitat in a random fashion.
I also tested the null hypothesis that among the species there was no

107
Table 4-5. Mean abundances (FIRST CAPTURES) of eight species of
small mammals per trapping line across five trapping sites.
Trapping sites abbreviations explained in text. Kruskal-Wallis
univariate nonparametric test for no site differences and manova
test with F approximations for no overall site differences with all
species considered simultaneously. * p < 0.05.
SPECIES
TRAPPING
SITES
RD/F
RD/H
RD/M
RD/T
RD/C
DidelDhis marsuDialis
0.66
0.66
0.33
1.33
4.33
Metachi rus nudicaudatus
5.33
7.33
4.33
6.00
7.00
Marmosa incana
4.00
14.66
4.00
5.33
2.00
Marmosa cinerea
6.66
10.33
5.66
5.33
3.33
Caluromvs Dhilander *
0.00
3.66
0.66
2.66
1.00
Akodon cursor *
1.00
7.00
0.66
0.00
0.33
Oecomvs trinitatis
1.33
2.33
1.00
1.00
0.66
Orvzomvs caoito *
0.00
1.33
3.00
0.66
0.00
MANOVA
WILK’S LAMBDA 4.6625
P < 0.0028

108
differential habitat utilization. I used the log-likelihood ratio test
for independence to determine if the species exhibited significant
differences in their use of the vegetation types or sites. The results
of the log-likelihood ratio test for independence suggest that all of
the species did exhibit significant differences in their use of the 5
trapping sites (G=51.717, df=20, p < 0.001).
I tested each pair of species for significant association at each
trapping station by using two-way contingency analysis of the
frequencies of presence and absence. Akodon and M. incana occurred
more frequently than expected by chance in RD/H, while Didelphis
was more abundant than expected by chance in RD/C. Caluromvs
exhibited higher than expected abundances in RD/H and RD/T.
Species capture overlaps per trapping station ranged from 2 to 6
species. There were no captures in 8.8% of the trapping stations
across all trapping sites. Approximately one third of all trapping
stations (35.4%) yielded single species captures. Two species captures
per site was also common (27.5%). Metachi rus. M. cinerea, and M.
incana were captured at 42%, 45% and 35% of all available trapping
stations, respectively. All other species were captured at less than
12% of all trapping stations. Metachi rus and Akodon were trapped at
the same station more frequently than expected and showed
significant association across all trapping sites (chi-square= 10.19,
df=1, p < 0.001). Metachi rus and Akodon (p < 0.025) and Metachi rus
and M. incana (p < 0.05) were trapped at the same trapping stations
in trapping site RD/H more frequently than expected under the null
hypothesis of independence.

109
I compared small mammal abundances by species per site using
Kruskal-WalHs test (Sokal and Rohlf, 1981). Small mammal abundance
was calculated with the first capture data per trapping line per
trapping site. Only those species with at least 15 first captures were
considered for this analysis. Mean number of individuals per species
per trapping line were considered independent samples and the overall
mean number of individuals per trapping site was compared across all
sites. Results of the Kruskal-Wal1 is test on abundances revealed
significant differences among trapping sites for Caluromvs philander.
Akodon cursor, and Orvzomvs capito (p < 0.05). Didelphis
marsupialis. Metachi rus nudicaudatus. Marmosa incana. M. cinerea and
Oecomvs trinitatis showed no significant differences in abundances
among trapping sites (Table 4-6). I used multivariate analysis of
variance (MANOVA) to test the null hypothesis that small mammal
abundances, considering all species simultaneously, did not vary among
sites. Multivariate analysis of variance, considering all of the
species abundances simulaneously, indicated significant variation in
abundance patterns among trapping sites (Table 4-6).
Small Mammal Microhabitat Selection
I used ANOVA and MANOVA for comparisons between
microhabitat observations for each small mammal species against a
pooled sample of observations for available habitat (August, 1984).
My null hypothesis was that there was no difference between small
mammal microhabitat and available habitat. Tests were made using all
habitat variables at trapping stations where the species in question

110
Table 4-6. Sampling effort and the number of individuals of small
mammals (species with > 15 first captures) in each of the five
trapping sites sampled in the Rio Doce State Forestry Park.
Trapping site abbreviations explained in text. Significant G5
scores indicate that species do not use the trapping sites in
random fashion (Log-Likelihood Ratio Goodness of Fit). Significant
G28 score suggests that the eight species exhibited significant
differences in trapping site use (Log-Likelihood Ratio Test for
Independence). * p < 0.025 ** p < 0.01
SPECIES
TRAPPING
SITE
NUMBER
TRAP NIGHTS DM
MN
MI
MC
CP
AC
OT
OC
RD/F
6000
2
16
12
20
0
3
4
0
RD/H
6780
2
23
44
31
11
21
7
4
RD/M
6300
1
13
12
17
2
2
3
9
RD/T
6300
4
18
16
16
8
0
3
2
RD/C
6780
13
21
6
10
3
1
2
0
TOTALS
32160
22
91
90
94
24
27
19
15
G5
**
*
*
*
G28 = 51.71
P < 0.001

111
was captured against all trapping stations irregardless if the species
was captured or not. These samples are not truly independent.
However, differences in microhabitat variables will be more
meaningful in this regard versus differences with capture station data
removed from the analyses. Differences in microhabitat variables
should be more sensitive with the capture station data removed, but,
real differences would be more pronounced with all stations included
in the analysis. I log transformed the habitat variable values because
preliminary analyses revealed that many of the habitat variables were
not normally distributed. Separate one way analyses of variance were
conducted for each habitat variable and each species. Multivariate
analyses of variance was used to test the null hypothesis with all
variables considered simultaneously.
Only Akodon showed significant differences in the univariate
analysis of habitat variables against the pooled habitat variables in
RD/H (Table 4-7). This means that for this species, the habitat
variable means of relief, herbvol, and perherb from all stations where
this species was captured differed significantly from these same
variables at all available trapping stations in this site. No other
species in any other trapping sites showed any significant differences
on any habitat variables. None of the multivariate analyses per
trapping site revealed any significant differences.
Six species showed significant univariate habitat variables when
compared with a pooled sample of observations for available habitat
across all trapping sites (Table 4-8). Metachi rus and 0. capito
tended to be captured at random and did not occur at any trapping

112
Table 4-7. Habitat profile of Akodon cursor in trapping site RD/H.
N= sample size, * p < 0.05, sign in parentheses indicates direction
of mean of that microhabitat variable. Mnemonic microhabitat
variables explained in Table 4-2.
MICROHABITAT
VARIBALES
F VALUE
PR > F
RELIEF
4.48
0.03
HERBVOL
4.54
0.03
PERHERB
5.53
0.02
MANOVA RESULTS
WILK’S LAMBDA
0.7186
EXACT F STATISTICS 0.4895
P > F
0.9725

113
Table 4-8. Univariate and multivariate analysis of variance for
comparisons between all significant microhabitat observations for
each species against a pooled sample of observations for available
habitat. * p < 0.05 ** p < 0.01 *** p < 0.001. Sign in
parentheses indicates direction of mean of that microhabitat
variable for each species. Species abbreviations given in Table 4-
6. Microhabitat mnemonics given in Table 4-2.
MICROHABITAT
VARIABLES
SPECIES
DM MN
MI MC
CP
AK OC
OT
NOFAM
*(-)
NOTREE
*(-)
NOSPP
**(-)
CANHT
*(+)
HERBHT
*(+)
CANCON
*(+)
PERHERB
***(+)
HERBVOL
*(+)
EPIP
*(+)
*(-)
RELIEF
*(+)
**(-)
SOILD
*(+)
SOILC
*(+)
*(+)
FERNS
*(+)
**(+)
***(+)
BANA
**( + )
STUMPS
*(+)
WILK’S
LAMBDA
P > F
0.91
0.81
0.97
0.99
0.93
0.87
0.95
0.96
0.85
0.29
0.78
0.16
0.88
0.93
0.90
0.73

114
stations characterized by any single variable. Didelphis occurred at
trapping stations with more arboreal epiphytes and terrestrial ferns (p
< 0.05). M. incana occurred at areas with fewer arboreal epiphytes
(p < 0.05). M. cinerea tended to be captured at trapping stations
that were in hilly, rather than flat, terrain (p < 0.05). Oecomvs was
captured more frequently at stations which had stumps (p < 0.05).
Akodon and Caluromvs were the only species that were captured at
stations characterized by several habitat variables. Akodon occurred
in areas with low tree diversity and number and relatively flat
terrain. This species was trapped at areas that had a well developed
herbaceous stratum, high bananeira and ground fern density, and
moist to wet soils. Caluromvs was captured at trapping stations with
high and well connected canopies, moist to wet soils and dense
ground ferns.
The results of the multivariate analysis, which tested the null
hypothesis that habitat variables from the capture profile of the
species are equivalent to all habitat variables of available habitat,
showed no significant differences between the species microhabitat
and available habitat.
Discussion
The observed differences in small mammal abundance patterns
across all trapping sites was primarily due to the variation in
abundances of Caluromvs. Akodon and Orvzomvs capito. All but two
of the species, Metachi rus and Marmosa cinerea. used the habitat in a
non-random fashion and the log-1 ike1ihood test for independence

115
suggested that the frequency of occurrence of a species in a trapping
site is dependent on the trapping site or habitat type. These results
suggest that the small mammal species are occurring at certain
trapping sites over others and that relative abundances vary per site.
Upon preliminary examination of these data, I would expect that
there should be significant microhabitat competition by the small
mammals.
Willig et al. (1986) report on. the use of univariate and
multivariate approaches in the assessment of morphometric variation
in natural populations, and conclude that the multivariate approach is
a more informative statistical test. By extension, multivariate tests
should also be the more appropriate test to evaluate overall group
differences in this study since these tests account for the
correlations among the microhabitat variables. Two species,
Caluromvs and Akodon. were captured at stations characterized by
several habitat variables. The remaining species either were captured
at stations that did not differ from the pooled sarnie, or that differed
on only one habitat variable. Caluromvs and Akodon appear to be
habitat specialists, while the remaining species are habitat generalists.
No species appeared to have a microhabitat preference when all of
the habitat variables were considered simultaneously. These results
suggest that all the species are habitat generalists. These results are
not consistent with my expectations.
The univariate method, although less informative than the
multivariate method, has some validity in determining the microhabitat
preferences of Caluromvs and Akodon and the lack of microhabitat

116
preferences of the other species in this study. From this chapter’s
analyis, we have seen that relative abundances of Akodon were higher
than expected in trapping site RD/H and that in this habitat this
species tended to be trapped more frequently at stations with little
relative relief and high herbaceous volume and percent cover. Across
all trapping sites, this species tended to occur in stations with low
species diversity of trees, well developed herbaceous stratum and
little vertical relief. Akodon cursor is strictly terrestrial and occurs
more frequently in grass/open habitats rather than forested habitats
(Chapter I). However, it is considered a pioneer species and
colonizes gaps in an efficient manner. Thus, although Akodon
appears to be a habitat specialist in the forest, it is not restricted to
the forest and because of its reliance on dense herbaceous ground
cover, is actually a grass/open habitat specialist.
Caluromvs is a highly arboreal forest species (Chapter I).
Across all trapping sites, this species occurred more frequently than
expected at stations where canopy height and connectivity were
great, with deep and moist soil and high terrestrial fern densities.
Caluromvs was captured more frequently than expected in trapping
sites RD/H and RD/T, both of which have areas with the above
mentioned microhabitat features.
Metachi rus. M. incana and M. cinerea were the most abundant of
the species captured. They were captured across all habitat types or
trapping sites. Individual abundances did not vary among trapping
sites and these species appeared to use the available habitat in a

117
random fashion. Together these 3 species were captured at
approximately 40* of all available trapping stations.
If most of the species do not appear to be selecting
microhabitats within a given area, how do they partition the feeding
and spatial resources? The 8 species used in this analysis frequent
three forest strata. Hetachirus. 0. cacito and Akodon are terrestrial
and occur on the forest floor. Didelphis. M. incana and Oecomvs are
scansorial and occupy the herbaceous and subcanopy strata. M.
cinerea and Caluromvs are highly arboreal and tend to occupy the
upper canopy stratum, although they do come to the ground.
Species that co-exist in each habitat stratum should differ
substantially in body mass or feeding guild (MacArthur, 1972) and
temporal activity (Pianka, 1983). One requirement for co-existing
species follows the "square of 2" rule (Hutchinson, 1959) which states
that the larger of two co-existing species should be approximately
twice the mass of the smaller. Larger species should be capable to
displace smaller species from feeding sites (Clutton-Brock and Harvey,
1983). Species differing by this much should have significant
differences in food resources, food particle size and resting space
requirements. Inherent within this requirement for co-existence is
the assumption that species occurring in the same habitat (stratum)
should differ in feeding strategies in order to avoid or reduce
interspecific competition. Temporal variation in activity patterns and
social structure differences also could lead to niche co-existence.
Small mammal body mass varied by spatial segregation (Chapter
I). Metachi rus has a mean mass of 258 g, while 0. capito and

118
Akodon. 60 and 40 g, respectfully, are much smaller. Metachi rus and
Akodon were trapped at the same station more frequently than
expected, while 0. capito and Akodon showed no such significant
association. There was also variation in the mean mass of the
scansorial species. Didelphis was the heaviest species used in the
analysis with a mean mass of 1000 g. Oecomvs and M. incana have
similar mean mass, 70 and 63 g, respectfully. There was no
significant association between any pair of these scansorial species.
For the two arboreal species, Caluromvs is approximately twice the
mass of M. cinerea.
Trophic category also varied by spatial segregation (Chapter I).
Metachi rus and Akodon are terrestrial insectivore / omnivores and 0.
capito is a terrestrial frugivore / granivore. Didelphis is a scansorial
frugivore / omnivore, M. incana a scansorial insectivore / omnivore
and Oecomys a scansorial frugivore / granivore. Caluromvs is an
arboreal frugivore / omnivore and M. cinerea is an arboreal
insectivore / omnivore.
Temporal activity overlapped for most species, as most of the
species are nocturnal. Didelphid marsupials are nocturnal (Nowak and
Paradiso, 1983; Miles et al., 1981; Charles-Dominique, 1983). Akodon
probably exhibits episodic activity bouts throughout the 24 hour
period (J. Eisenberg, pers. litt.), while Oecomvs and 0. capito
probably are nocturnal. Didelphid marsupials are usually solitary and
defend no exclusive area or territory, with little agonistic behavior
exhibited among conspecifics (Charles-Dominique, 1983).

119
There appears to be clear divisions in body size, spatial
segregation and dietary classifications of the small mammals in spite
of a general lack of microhabitat selection. These patterns are in
somewhat accord with the observations of Miles et al. (1981) on the
importance of the stratification of tropical forests. Within each
stratum (terrestrial, herbaceous and subcanopy, and canopy), there are
distinct trophic strategies. This stratification can be perceived as
"parallel microhabitats" with distinct trophic categories in each
stratum or parallel microhabitat. The clear divisions in the mass and
dietary and spatial adaptations of the species in question reduce
competition for space and food resources. These observations were
also made for didelphid marsupials by Eisenberg and Wilson (1981).
An interesting phenomenon observed in this study was the
ubiquitous distribution and relative high density of three species of
marsupials, Metachi rus. M. cinerea and M. incana. This observation is
in accord with Hanski (1982, cited in Weins, 1985) who noted a
bimodal frequency distribution of species among locations, where
“core species" occur at high densities at most sites and "satellite
species" occur at low frequencies. These three didelphid species
have distinct body masses and are spatially segregated. However
these species are not readily separated by diet; all three species are
basically omnivorous. Perhaps their success in absolute numbers
within each spatial category can be explained in terms of being a
generalist, rather than a specialist.
If a species is abundant, it usually is more capable of using the
production of resources over less abundant species (MacArthur, 1972).

120
These opposums are broadly adapted to a variety of food resources
within their distinct spatial categories. Omnivores are able to switch
to the commonest suitable resource and keep all resources rare.
However, these species can not completely exploit all possible
resources and other food competitors, that have stricter diets, can
gain access to these resources. In this fashion, stenophagic species
can co-exist with euryphagic species, albeit at lower densities.
Contemporary ecological dogma states that in order for a species
to be successful in tropical forests, it needs a well defined ecological
niche because of the high degree of species richness. However,
eurytopic species may be favored by available resources. A proximate
explanation for high density species could be that the abundance is
proportionate to the available habitat.
Charles-Dominique (1983) pointed out that in sites with abundant
food resources, didelphid opposums can reach high densities because
of their high reproductive capacity, rapid growth rate, and lack of
territoriality. The habitats examined in this study ranged from early
succession, to mid-stage succession to primary forests. Secondary
forests generally are more productive than primary forests (Foster,
1980) because the former invests more energy into production of fruit
and seeds, leaves and woody material. Climax adapted plant species
tend to invest more energy into longevity (Charles-Dominique, 1983).
The didelphid marsupials have evolved mechanisms enabling the
various species to exploit different spatial strata and a wide range of
food resources. These mechanisms have enabled this group to
dominate each trophic category and use the habitat in a general

121
fashion. Forest rodents also have evolved mechanisms to exploit the
spatial strata, but are bound by a more specialized diet. These
differences in the trophic categories of the two groups helps tp
explain how some species of the didelphid marsupials are the
numerically dominant group in some forested habitats.
The numerically abundant, as well as some of the less abundant,
species in this park did not tend to select microhabitats. One
interpretation of these results is that the habitat structure in the
park does not impose selective pressure for microhabitat specialists.
This could be because of the vast disturbances that have occurred, in
the form of intense crown fires, in recent and perhaps evolutionary
past history. However, perhaps the true habitat specialists are those
that were trapped at low frequencies and subsequently were not
included in the analysis. Species such as Abrawavomvs ruschii are
rare and very little is known of their natural history. These species
may not necessarily have low densities, rather, they may be difficult
to trap due to habitat peculiarities or bait preferences. But for the
species analyzed in this study, it appears that the spatial adaptation,
dietary classification and body size play important roles in the
manner in which these small mammals partition the available
resources.

CHAPTER V
FOREST FIRE AS A DETERMINANT OF
SMALL MAMMAL DIVERSITY IN A BRAZILIAN FOREST
Introduction
The hypotheses that have been developed to account for the
high level of species diversity observed in tropical forests have
traditionally been based on “climax" species equilibrium theory
(MacArthur and Levins, 1967; Ashton, 1969; MacArthur, 1969). The
view that environmental stability promotes species diversity derives in
part from the observation that diversity decreases with latitude and
the assumption that tropical environments are more stable because
they do not experience the seasonal fluctuations characteristic of
more temperate areas. This lack of disturbance allows for increased
niche diversification, higher speciation, and lower extinction rates
(Ashton, 1969; Stebbins, 1974). High diversity is therefore maintained
in climax conditions. If disturbed, the system returns to an earlier
successional stage with lower species diversity.
A current view is that environmental instability promotes species
diversity. "Climax" species diversity is low, but climax is seldom
reached in natural systems because frequent disturbances hinder the
process of competitive exclusion. High diversity is the result of
constantly changing conditions that promote a community of
competing species. Recent theory (Connell, 1978) suggested that high
122

123
species diversity is only maintained at some intermediate level of the
frequency, extent, and intensity of the disturbance. Connell (1978)
suggested that intermediate disturbances caused by hurricanes
increased species diversity of sessile organisms in a tropical reef.
Garwood et al. (1979) described landslides caused by earthquakes as a
source of disturbance to tropical forests. Epiphyte loads causing
treefalls (Strong, 1977) and tree falls due to wind (Connell, 1978)
have also been suggested as mechanisms to promote species diversity
in non equilibrium theory.
It is not widely accepted that fire 1s a major determinant of
tropical diversity. However, fire has long been associated with
tropical savanna formation or maintenance (Boughley, 1963;
Daubenmire, 1968; Scott, 1977) and forest fire is not uncommon in
seasonally dry tropical forests of Australia (Stocker and Mott, 1981;
Recher and Christensen, 1981; Vickery, 1984). Recent evidence
suggests that fire in the lowland humid tropical forest may not be as
uncommon as previously believed (Sanford et al., 1985; Stocker and
Mott, 1981; Uhl and Buschbacher, 1985) and may cause widespread,
rather that local disturbance. Such was the case of the 1983 fire in
Borneo, which burned three million hectares of evergreen tropical
rainforest (Leighton, 1984). Fire was cited by officials as a major
threat to the vegetation in tropical and subtropical parks (Machiis
and Tichnell, 1985). Core samples from the minimum critical size
experiment near Manaus in the Amazon basin show evidence of
extensive burning of this humid forest (R. Biergaard, pers. comm.).
Sanford et al. (1985) suggest that tropical forest fire be considered a

124
moderate disturbance factor in the upper Rio Negro region in the
north central Amazon Basin. Stocker and Mott (1981) stated that
humid forests in Australia burn, although not as frequently as drier
habitats.
Humid forests are succeptible to forest fire especially toward
the end of an exceptionally severe dry period (Phillips, 1974).
Leighton (1984) suggested that the 1982-83 El Nino Southern
Oscillation event caused intense drought in the humid old world
tropical forests of Borneo and consequently fire was able to destroy
millions of hectares of tropical forest. Forest fire has been
associated with shaping forest communities since the Mesozoic
(Harris, 1958) and it appears that fire might play a major role in the
formation and structure of tropical forests. The mechanism by which
fire disturbance affects the species diversity of mobile organisms
might include a) alteration of the structural complexity of the area
affecting alpha or within habitat diversity, and b) the creation of a
habitat mosaic comprised of early and late successional areas thereby
affecting habitat heterogeneity or beta (between habitat) diversity.
August (1983) defined complexity for small mammals as the
development of vertical strata within a habitat. MacArthur and
MacArthur (1961) used foliage height diversity as a measure of
structural complexity. Habitat structural complexity has been
correlated with bird species diversity (MacArthur, 1957, 1964, 1965;
MacArthur and MacArthur, 1961; MacArthur et al., 1966; Cody, 1968;
Recher, 1969; Karr and Roth, 1971). Rozenzweig and Winakur (1969)
reached similar conclusions for rodents, and Pianka (1967) for lizards.

125
Widespread fire can increase horizontal homogeneity within a
forest, whereas localized burning can create a mosaic structure
resulting in a heterogeneous vegetation (Wiens, 1985). Karr and
Freemark (1985) suggested that the impact of localized disturbances
in producing habitat mosaics plays an important role in determining
community attributes. Creation of a habitat mosaic would allow early
successional plant species to colonize the disturbed areas (Foster,
1980). Disturbed areas, depending upon the areal extent of the
disturbance, should therefore be characterized by spatial
heterogeneity and include areas containing species from different
successional stages. Plant species diversity should increase with this
form of disturbance, but effects of animal diversity are unclear.
Levin (1974) concluded that increased habitat heterogeneity increased
species diversity. August (1983) found no correlation between
patchiness or habitat heterogeneity and small mammal species
diversity, while others reported positive correlations for birds
(MacArthur et al., 1966; Roth, 1976).
Clearly, disturbance plays a crucial role in the structure of
natural communities (Sousa, 1984) and the areal extent of the
disturbance is dependent on several factors. Intense crown fire can
completely destroy a forested habitat or it can burn the forest in a
mosaic fashion. It seems clear that on the local scale, patchiness
would be maximized at some intermediate level of disturbance.
The objective of this study was to test the intermediate
disturbance hypothesis (Connell, 1978) using small mammals as
indicator species. This hypothesis predicts that disturbances of

126
intermediate scale promote higher species diversity of tropical trees
and corals. The data for tropical trees comes mostly from
successional studies (Connell, 1978) where the intermediate levels of
frequency, time after disturbance, and size of the disturbance yielded
higher species diversity. In this study, I predicted that small mammal
species diversity would be maximized in forested areas that are
characterized by disturbances of intermediate intensity. This
prediction assumes that patchiness of the environment is maximized in
intermediate disturbances.
Materials and Methods
I sampled 5 forested habitats in a large (35,000 ha) Brazilian
state forestry park in the Altantic Rain Forest. This study was
conducted in the Rio Doce State Forestry Park (PFERD), Minas
Gerais, Brazil (19 38’S, 42 33’ W). PFERD is one of the largest
(35,000 ha.) protected areas in the highly endangered Atlantic
Rainforest and is managed by the State Forestry Institute. The Park
is covered by continuous forest with approximately 40 ox bow lakes
interspersed throughout the sanctuary. I considered that any
differences in the characteristics of the small mammal communities
are due to habitat differences between the sampling areas.
The vegetation of the Park is classified as tropical
semideciduous (Gilhaus, 1986). The climate of the Park is classified
as tropical humid, with approximately 1500 mm of annual precipitation
(Chapter I). Most of the annual precipitation occurs during the
summer months and the winter months are characterized by drought.

127
Sampling Areas
The Park has been well protected from hunting and fishing,
however, forest fire has been a problem since the Park’s creation in
1944. The forest in the Park is characterized by frequent burning,
which frequently totally removes the forest cover in some areas.
Regeneration is rapid, and areas totally burned 20 years ago now
support a developed forest 10 to 15 m tall. All of the Park has
probably been burned in historical times. The habitat within the
Park has been altered at least twice by intense crown fires since
1960. In 1963 and 1967, large fires burned substantial portions of the
Park (Lopes, 1982). The areas most affected by the intense crown
fires occurred on slopes of hills or on hilltops, rather than lowland
forests (J. Stallings, pers. obs.). Areas not burned in either 1963 and
1967 support a tall (25-35 m), complex, and relatively homogeneous
forest. Forest stands on hilltops which have not been burned for
several decades support tall forests with dense epiphyte loads (J.
Stallings, pers. obs.; J. Ladeira, per. comm.). Forest fire therefore
maintains a mosaic of different forest or habitat types.
The frequent fire disturbances in the Park have resulted in
forest at different stages of regrowth, age and heterogeneity. Casual
observations suggested that the burned areas harbored a richer, yet
different, flora than previously observed in same forests before the
fire (Lopes, 1982). The diverse burned areas supported an increase in
certain wildlife species, such as tapirs (Tapirus terrestris). titi
monkeys (Cal 1icebus personatus) and cracids (Penelope obscura).

128
Five areas were selected for study (Table 5-1). These areas
represented 3 of the 5 forested habitats that have been described for
the Park (Gilhaus, 1986; Chapter I). In each area I cut 3 300 m
parallel lines separated by 100 m through the forest. Each line
included 16 trapping stations separated from one another by 20 m.
Each area had equal number of traps and all traps were placed within
a 3.5 m radius circle at each trapping station. Traps consisted of
large terrestrial live traps, and Sherman and medium sized live traps
positioned at varying heights in standing vegetation, as well as on
the ground. Traps were baited with dry oatmeal, pineapple, and
cotton balls soaked in cod’s liver oil and sardines. Traps were
opened for 5 consecutive days each month for 12 consecutive months.
Trapping procedures are presented in more detail in Chapter I.
Results of a previous discriminant analysis (Chapter IV)
indicated that the areas chosen for established trapping lines differed
in forest structure. Sampling area RD/F was burned totally to the
ground in 1967 and is characterized by high tree and shrub species
diversity, high understory vine density, low canopy, low canopy
volume and connectivity, small diameter trees, and underdeveloped
herbaceous stratum. This forest was impenetrable due to dense shrub
and vines and was the epitome of a secondary succession forest as
described by Ewell (1980) and Budowskl (1963, 1965).
Sampling areas RD/M and RD/H were both burned intensively,
but heterogeneously, in the 1967 fire. This resulted in a mosaic of
different habitats in both areas, containing both primary and
secondary forest stands. Area RD/M had a small stream cutting

129
Table 5-1. Study areas sampled in the Rio Doce State Forestry
Park, Minas Gerais, Brazil. Habitat types are compared to the
areas examined in this study and are taken from Gilhaus (1986).
THIS HABITAT DEGREE OF DISTURBANCE
STUDY TYPE BY FOREST FIRE (1967)
RD/C PRIMARY FOREST
WITH EPIPHTYES
NO EVIDENCE OF FIRE
RD/T PRIMARY FOREST
WITH EPIPHYTES
UNDERSTORY ALTERED BY
1967 GROUND FIRE
RD/M
MEDIUM TO TALL FOREST
WITH BAMBOOS AND GRAMINOIDS
SUBSTANTIAL CROWN FIRE
AFFECTING SOME LARGE
TREES IN 1967
RD/H
MEDIUM TO TALL FOREST
WITH BAMBOOS AND GRAMINOIDS
SUBSTANTIAL CROWN FIRE
AFFECTING SOME LARGE
TREES IN 1967
RD/F
MEDIUM SECONDARY FOREST BURNED TO GROUND
WITH BAMBOOS AND GRAMINOIDS IN 1967

130
across two of the 3 lines. Overall, these two habitats were similar in
structure when compared with the other 3 areas. RD/M and RD/H
had fewer epiphytes, fewer large and buttressed trees and a lower
canopy than the two primary forest areas. Area RD/F had lower
loadings in these categories than RD/H and RD/M.
Sampling areas RD/C and RD/T were considered primary forest
habitats. RD/C appeared to have been untouched by intense crown
fires since at least the creation of the park in 1944. However, area
RD/T was burned in 1967 and portions of the subcanopy were
destroyed. Both areas supported numerous large and buttressed trees,
high epiphyte densities, well developed canopy stratum, and high
terrestrial fern densities.
Estimate of Small Mammal Diversity
I used the Shannon-Wiener diversity index (H*) to estimate small
mammal diversity. H’ scores were generated using first capture data.
The progression of H’ scores was plotted for each month per area by
using the cumulative data from the first trapping session (month 1)
to the last trapping session (month 12). Using ANOVA, I tested the
null hypothesis that the cumulative H’ scores from the 12th trapping
session did not differ among sampling areas. Variance and standard
deviation of the expected value of H’ was calculated using the
method of Poole (1974). Individual t-tests were used to test the null
hypothesis that pairs of H’ scores were not different (Poole, 1974).
Cumulative species richness and diversity curves were plotted each
month to determine if the respective curves would asymptote. Curves

131
that fluctuate widely through time and show no sign of flattening out
would not be useful data to test the H’ scores (Pielou, 1975).
Estimates of Patchiness
In order to test the prediction that species diversity is
maximized in a forest characterized by spatial heterogeneity produced
by disturbance, it is necessary to quantify patchiness. In an earlier
analysis, I predicted that the forest structure would differ
significantly in areas that that were burned on a different areal
scale. I used mean of habitat variables measured at each trapping
station in each area to test this prediction. Variables used in this
analysis were mean tree height, coefficient of variation of the sum of
the basal area of trees, the number of trees, mean diameter of breast
height (DBH), and tree species diversity. Tree species diversity was
calculated using the Shannon-Weiner index. I expected that mean
tree height should be low in area RD/F, relatively moderate in RD/M
and RD/H, and higher in RD/T and RD/C. Mean DBH should be
highest in RD/C and RD/H and lowest in RD/F. Trees in RD/H and
RD/M should have moderate diameters. I expected more trees in
RD/F, moderate abundances in RD/M and RD/H, and fewer trees in
RD/C and RD/T. As indicated in Chapter IV and summarized in
Table 5-2, none of these indices generated results consistent with the
prediction, except for the coefficient of variation of the sum of the
basal area. I also expected that tree species diversity would be
higher in the mosaic habitats, however, these habitats had the lowest
diversity indices.

132
Table 5-2. Habitat variables used to test the prediction that
habitat structure differed in areas disturbed on different scales
in the Rio Doce State Forestry Park, Minas Gerais, Brazil. Only
the coefficient of the sum of the basal area (CV BASAL) provided
results consistent with the prediciton. Areas are listed in
decreasing order of areal disturbance.
VARIABLES AREAS
RD/F
RD/H
RD/M
RD/T
RD/C
MEAN TREE HEIGHT
6.13
6.49
7.06
6.73
7.15
CV BASAL AREA
76
248
209
138
167
NUMBER OF TREES
399
284
344
394
324
DIAMETER OF BREAST HEIGHT
8.5
10.9
10.2
11.7
11.2
H* (TREE SPECIES)
3.4
3.1
3.2
2.9
3.3

133
Only the coefficient of the sum of basal area (CV basal) was
related to extent of disturbance. CV basal scores were higher in
RD/H and RD/M mosaic sites and lower in the heavily disturbed
(RD/F) and relatively undisturbed (RD/T and RD/C) sites. This
relationship is intuitive because regenerating forests that were burned
completely to the ground should be composed of small diameter trees,
with few large diameter trees. The CV would be low. Primary
forests and primary forests that were only burned in the understory,
should be composed on large and small diameter trees, and should
have a high CV. Regenerating forests that were burned in an
intermediate or mosaic fashion, should have a very high CV because
many of the large trees being destroyed.
Another measure was the relative abundance of vascular
epiphytes and the variation of vascular epiphytes. I used the mean
epiphyte density in the burned sampling areas to quantify the scale
of disturbance caused by fire. Areas burned to the ground within the
last 20 years should not have any epiphytes. Areas burned in a
mosaic fashion should have features of secondary and primary
habitats, and should support some epiphytes in relatively undisturbed
stands of habitat. Areas not burned by crown fires should support
high densities of epiphytes. At each trapping station, the relative
density of epiphytes was recored in a subjective manner using a
sliding scale (0-3). A score of 0 indicated no vascular epiphytes in
the immediate vicinity of the trapping station, a score of 1 indicated
a modest amount of epiphytes, a score of 2 indicated a relatively

134
high amount of epiphytes and a score of 3 indicated a heavy load of
epiphytes.
Patchiness brought about by fire disturbance was defined by
using the coefficient of variation of the mean density of epiphytes
per burned area. The coefficient of variation (CV) a good indicator
of the degree of patchiness in each area because this measure
compares the variation independent of the magnitude of the means
(Sokal and Rohlf, 1981). A low CV indicates that there is very little
variation around the mean density of epiphytes and suggests that the
habitat is relatively homogeneous either because disturbance had been
extreme or because it had been slight. A high CV indicates that
habitat is spatially heterogeneous presumably because the disturbance
had been of an intermediate intensity producing a mosaic of habitat
variations. Forested habitats with a high CV are characterized by
stands of both early secondary and climax adapted species.
One of Connell’s (1978) predictions proposed that diversity is low
in areas of slight and extreme disturbances, and is highest when
disturbances are intermediate in intensity. This prediction suggests a
curvilinear relationship between the two variables (Figure 5-1). One
of the mechanisms by which this pattern is produced could be that
patchiness is maximized at some intermediate Intensity. Another
mechanism could be that species diversity should increase with an
increase in patchiness. These predictions will be tested using mean
epiphtye density as an estimator of intensity of disturbance and CV
of epiphyte abundance and CV of sum of basal area as estimators of
the patchiness of the disturbance. If diversity is higher in areas

135
EIATIONSHIP
H’ AND DJSTURI
SPECIES DIVERSITY (H’)
CURVE TAKEN FROM CONNELL (1978)
Figure 5-1. Curvilinear relationship between species diversity
and disturbance as suggested by Connell’s intermediate
disturbance hypothesis. Species diversity should be maximized at
some intermediate level of disturbance.

136
disturbed by intermediate intensities, then there should be a
significant positive correlation between the variables. I tested these
predictions using simple linear regression.
Methodological Caveats
I believe that the Shannon-Weiner H’ index is the best estimator
of species diversity of small mammals. Other authors have used
several estimators, some of which include species richness, densities,
biomass and Pielou’s evenness index. I am in accord with Pielou
(1975) and Poole (1974) that the H’ is the correct estimator to be
used because I am fairly confident that the total number of species
that could be trapped by my methodology are represented by the
trapping results. Cumulative species curves remained constant after
approximately eight months in all but one case. Some of the species
that were captured are very rare indeed. A case in point is the
rodent Abrawayomvs ruschii which has only been reported in the
literature since the description of type specimen in 1979 (Chapter I).
However, relative trapping frequencies of Caluromvs. a highly arboreal
species, can be significantly increased by using arboreal traps placed
high in the canopy. I did not use the high arboreal trap results to
calculate H’ in the areas that had these traps. The arboreal trapping
effort was not equal among the areas and would have made the
interpretation of the results difficult. In addition to the live traps, I
also sampled the small mammal populations with snap traps which
often tend to be more effective than live traps (J. Stallings, pers.
obs.). Snap trap results revealed the same number of species.

137
Sampling areas were relatively homogeneous with respect to
moisture availability except for the stream that cut across mosaic
habitat RD/M. Two species associated with aquatic habitats,
Oxvmvcterus roberti and Nectomys sauamioes were trapped in this
habitat and their presence and proportions may have played a role in
the high diversity index calculated for this area. However, these two
species were also trapped in the other mosaic sampling area where no
streams occurred.
Does the sample size and trapping design allow the prediction to
be tested? Some researchers believe that 40 traps per line in
tropical forests are insufficient to sample small mammals because of
notoriously low live trapping results. My trapping success of 2.5%
falls within the observed range of trapping successes reported for
Neotropical forest studies, and my trapping effort of 31,000 trap
nights in one year ranks among the highest (Chapter I).
Vascular epiphytes depend on the forest structure (Richards,
1952). In the study area, vascular epiphytes were represented by
arboreal cacti, brome!iads, ferns, orchids, and palms. Epiphytes do
not recolonize an area for approximately 20 years following a
disturbance (Budowski, 1963). Drudgeon (1923, cited in Johansson,
1974) also reports that in the course of epiphytic succession, ferns
and vascular epiphytes did not appear in the secondary forests for
approximately 20 years. Others have supported the view of no to
few vascular epiphytes occurring in early to secondary serai stages
(Budowski, 1963; Johansson, 1974; Whitmore, 1975; Yeaton and
Gladstone, 1982). The reason for the long lag time is that the forest

138
must produce a suitable microenvironment for epiphytes (Richards,
1952; Johansson, 1974). Vascular epiphytes depend on relatively
undisturbed forest stands because of several factors, some of which
are high humidity and suitable substrate on climax adapted species
(e.g., rough bark for gaining an attachment site).
Because epiphytes are dependent on forest structure, they are
good indicators of the degree of disturbance in forests disturbed by
fire, even though casual observations may not detect physical patches
or gaps due to previous forest fire. There should be no vascular
epiphytes in regenerating forests burned to the ground over a large
area within twenty years of the disturbance. A large burned area
represent a large scale disturbance. Areas burned in a mosaic fashion
should support epiphytes in areas not affected by the fire, while
burned portions of the forest should be devoid of epiphytes. These
forests represent areal disturbances of an intermediate scale. Primary
forest habitats that have only the subcanopy disturbed by fire should
have high epiphyte densities because they represent disturbed on a
fine scale. Natural gaps caused by treefalls should account for the
majority of the variation of epiphyte densities observed in primary
forests not affected by fire. These forests also are disturbed on a
fine scale.
Results
Species Diversity
Species richness of small mammals varied among the sampling
areas (Table 5-3). The heavily disturbed area, RD/F, and the two
primary forest areas, RD/T and RD/C, each had seven species of

139
Table 5-3. Species richness, species diversity, and total number
of individuals of small mammals captured in each area in Rio Doce
State Forestry Park, Minas Gerais, Brazil.
AREA
SPECIES
RICHNESS
SPECIES
DIVERSITY
NUMBER OF
INDIVIDUALS
RD/F
7
1.45
64
RD/H
12
1.80
137
RD/M
10
1.78
63
RD/T
7
1.40
56
RD/C
7
1.30
49

140
small mammals. The two intermediately disturbed areas, RD/H and
RD/M, had 13 and 10 species, respectively.
Figures 5-2 through 5-6 illustrate the community structure of
small mammals in each area. In all cases, didelphid marsupials
dominated the communities in terms of absolute numbers, and 3
species, Metachi rus nudicaudatus. Marmosa incana and Marmosa
cinerea. were ubiquitous and abundant. Cumulative species curves
tended to flatten out after the eighth month of the study (Figure 5-
7). H’ scores, generated in a cumulative fashion for each trapping
session, also stabilized after approximately 8 or 9 months (Figure 5-
8).
H’ scores varied significantly among sampling areas (ANOVA, F=
3.94, p < 0.05). Sampling area RD/F had a significantly lower H’
score than area RD/H (t = 6.93, p < 0.001), forest mosaics RD/H and
RD/M did not differ significantly (t=0.44, p > 0.90), forest mosaic
RD/M and primary forest RD/T differed significantly (t= 3.51,
p < 0.001) and primary forest areas RD/T and RD/C did not differ
( t=0.92, p > 0.20).
Patchiness and Intensity of Disturbance
Table 5-4 quantifies epiphyte abundance and degree of
disturbance in the five habitats. Mean epiphyte densities ranged from
a low 0.00 in RD/F, to high densities of 1.08 and 1.98 in primary
forest sites RD/T and RD/C, respectively. The coefficients of
variation of epiphytes, which estimates degree of disturbance, ranged
from 0.0 to 278.9 percent. Area RD/F, which was burned in a coarse
scale, had no epiphytes and correspondingly no variation. The areas

141
SMALL MAMMAL COMMUNITY
SPECIES
NUMBER OF INDIVIDUALS
Figure 5-2. Community structure of small mammals 1n primary forest
site RD/T in the Rio Doce State Forestry Park, Minas Gerais,
Brazil.

142
SPECIES
NUMBER OF INDIVIDUALS
Figure 5-3. Community structure of small mammals in primary forest
site RD/C in the Rio Doce State Forestry Park, Minas Gerais,
Brazi1.

143
SPECIES
M.incana
M. nudicaudatus
M. cinerea
A. cursor
0. trinitatis
0. capito
N.squamipes
C. philander
D. marsupialis
O.roberti
0. subfiavus
M. microtarsus
A. ruschii
0 10 20 30 40 50
NUMBER OF INDIVIDUALS
Figure 5-4. Community structure of small mammals in mosaic forest
site RD/H in the Rio Doce State Forestry Park, Minas Gerais,
Brazil.

144
¡N FOREST SITE RD/M
SPECIES
NUMBER OF INDIVIDUALS
Figure 5-5. Community structure of small mammals in mosaic site
RD/M in the Rio Doce State Forestry Park, Minas Gerais, Brazil.

145
SPECIES
NUMBER OF INDIVIDUALS
Figure 5-6. Community structure of small mammals in secondary
forest site RD/F in the Rio Doce State Forestry Park, Minas Gerais,
Brazil.

146
OUMUIATIV
CURVES
NUMBER OF SPECIES
MONTHS
RD/h RD/C -*-RD/F ~a~ RD/M RD/T
Figure 5-7. Cumulative species curves for all forested sites in
the Rio Doce State Forestry Park, Minas Gerais, Brazil.

147
H* SCORES
MONTHS
RD/F RD/H RD/M RD/T RD/C
Figure 5-8. H’ scores for small mammals generated in cumulative
fashion in all forested sites in the Rio Doce State Forestry
Park, Minas Gerais, Brazil.

148
Table 5-4. Quantification of patchiness caused by fire disturbance
in forested sampling areas in Rio Doce State Forestry Park, Minas
Gerais, Brazil. Patchiness is derived from vascular epiphyte
relative densities mean sum of basal area in the sampling areas.
Patchiness reflects the amount of variation of epiphyte densities
and sum of basal area within each area and is estimated using the
coefficient of variation. CV EPIPHYTES = coefficient of variation
of mean epiphyte densities. CV SUM BASAL AREA = coefficient of
variation of the mean sum basal area of trees.
SAMPLING
AREA
INTENSITY
PATCHINESS
MEAN EPIPHYTE
DENSITIES
CV
EPIPHYTES
CV SUM
BASAL AREA
RD/F
0.0
0.0
76
RD/H
0.2
278.9
248
RD/M
0.3
255.2
209
RD/T
1.1
73.1
138
RD/C
1.9
43.9
167

149
burned in mosaic fashions or an intermediate scale, RD/H and RD/M,
had similar and relatively high coefficients of variation. The primary
forest areas, RD/T and RD/C, also showed variation in the density of
epiphytes. RD/T and RD/C were both disturbed on a fine scale, by
forest fire and tree falls, respectfully.
I did not find a significant linear relationship between
patchiness and intensity of disturbance, using either CV epiphyte or
CV sum of basal area as estimators of patchiness (Figure 5-9).
Rather, these results suggest a curvilinear relationship.
Small Mammal Diversity and Disturbance
Species diversity (H’) of small mammals was significantly
correlated with degree or scale of disturbance (r= 0.93, Ho: p=0, p <
0.02). I found a linear relationship between species diversity of small
mammals and disturbance (Figure 5-10). However, the relationship
between species diversity of small mammals and CV of sum basal area
was not as significant (r= 0.71, Ho: p=0, p> 0.20).
Primates and Disturbance
The prediction also holds for other taxa. I examined the data
obtained from census counts of larger vertebrates observed while
walking the trapping transects (Chapter I). Table A-3 lists the
species that I observed from 60 km. of repeat censuses per area. I
chose to use the primate data to test the prediction because primates
are relatively easy to see in comparison to the other taxa listed in
Table A-3 and are dependent on forest structure since they are
arboreal. In Table 5-5, I present primate abundance (number of
individuals observed per Km.) and diversity (number of species

150
NATION!
PATCHINESI
M
Ini
DiSTURBANG
PATCHINESS
INTENSITY OF DISTURBANCE
CV EPIPHYTES —CV SUM BASAL AREA
Figure 5-9. Relationship between patchiness of habitat and
intensity of disturbance for all forested sites in the Rio Doce
State Forestry Park, Minas Gerais, Brazil.

151
RELATIONSHIP BETWEEN
H' & PATCHINESS
RELATIONSHIP BETWEEN
H’ & PATCHINESS
H*
H*
â– 
â– 
â–  â–  , , .
.
PATCHINESS
PATCHINESS
CV EP1P
CV BASAL APEA
Figure 5-10. Relationship between H’ of small mammals and
patchiness of habitat using both CV EPIP and CV BASAL AREA as
indicators of patchiness in the Rio Doce State Forestry Park,
Minas Gerais, Brazil.

152
Table 5-5. Estimates of abundance and diversity of primates in
five sampling areas in the Rio Doce State Forestry Park, Minas
Gerais, Brazil. Site abbreviations are explained in text. Primate
diversity is the number of species (# SPP) and abundance is the
number of individuals observed per linear Km. (#/KM.) in 60 Km. of
repeat censuses in each sampling area. Disturbance is the relative
amount of distrubance caused by fire, this variable is discussed in
text. Primate species are the following: A= Callicebus personatus.
B= Cebus apella. C= Brachyteles arachnoides. D= Alouatta fusca. E=
Callithrix geoffrovi. F= Cal 1ithrix aurita.
DISTURBANCE
SITE
# SPP
#/KM.
SPP.
HEAVY
RD/F
1
0.16
A
INTERMEDIATE
RD/H
4
1.29
A,B,C,F
INTERMEDIATE
RD/M
4
1.00
A,B,D,E
PRIMARY
RD/T
2
0.41
B, F
PRIMARY
RD/C
2
0.43
B, F

153
observed during study) in each of the areas. These data suggest that
at least for primates, there is a linear relationship between species
richness and abundance of primates and the CV of epiphyte
abundance (Figure 5-11). More primates and primate species occurred
in areas that had a high coefficient of variation of epiphytes.
It is not surprising that Callicebus was found to occur in the
highly and intermediately disturbed habitats. Cal 1icebus personatus
inhabits a wide range of habitat type and perhaps is the most
adaptable species of the genus (Kinzey, 1981). Outside of the park,
this species was frequently heard calling from the isolated and
degraded hill top forests that represent the majority of the native
habitat left in the region. Cebus monkeys tend to have wide
tolerance levels and occur in all forested habitat types (Freese and
Oppenheimer, 1981). Brachvteles was formerly considered a climax
forest adapted species, but Fonseca (1983) found that this species can
occur in very disturbed habitat. Alouatta tends to be found near
bodies of water (Eisenberg, pers. comm.). Sampling area RD/H occurs
near a large lake in the park and might help to explain its presence
in this habitat. Members of the genus Cal 1ithrix occur in various
habitat types (A. Rylands, pers. comm.). C. geoffroyi. native to the
Brazilian cerrado, was introduced into the park in the late 1970’s
(Stallings, in prep.). C. aurita is the marmoset species that is native
to the park. It was only observed to occur in the mosaic site RD/M
(at low densities, Stallings, in prep.) and in the primary forest
habitats, RD/C and RD/T.

154
RELATIONSHIP BETWEEN
PRIMATE ABUNDANCE & PATCHINESS-
RELATIONSHIP BETWEEN
SPECIES RICHNESS 4 PATCHINESS*
PRIMATE ABUNDANCE
NO. PRIMATE 3PEOIE3
'DERIVED FROM OV OF EPIPHYTE ABUNDANCE
•DERIVED PROM CM OF EPIPHYTE ABUNDANCE
A.
Figure 5-11. Relationship between patchiness of habitat and A)
primate abundance and B) primate species richness in the Rio Doce
State Forestry Park, Minas Gerais, Brazil.

155
These data propose that intermediate disturbances brought about
by fire tend to increase the diversity of the primate species that
occur in the park. Species that depend on primary habitat (e.g.,
Callithrix aurita petronius) might be displaced by introduced species
(e.g., C. geoffrovi) in the event of a habitat disturbance.
Discussion
The Atlantic forest of Brazil has a rich and endemic small
mammal fauna (Table A-I). I suggest that fire affects species
diversity of this fauna in this region. Fire has been proposed to
shape forest communities since the Mesozoic (Harris, 1958) and
tropical forests that have pronounced annual dry seasons are prone to
fires, whether of natural or anthropogenic origin. The El Nino
Southern Oscillation has been documented to cause dramatic
environmental effcts since the early 1700’s. Interestingly, the fire
that occurred in the Rio Doce Park in the dry 1963-64 period
overlapped with a strong 1963 El Nino.
Periodic disturbances brought about by fire could help to shape
tropical forests. Some authors (e.g., Hartshorn, 1980) believe that
the evolution of tropical succession from pioneer stages to late
succession or primary habitat takes approximately 300 to 500 years.
It is not difficult to perceive that during this period of time an
oscillation could occur in the usually humid climatic pattern and since
the frequency of lighting storms increases with a decrease in latitude,
fire could have played a major role in shaping forest communities.
The high level of small mammal species richness could be the result

156
of a mosaic of different aged forest communities with species evolved
to fill each of the available niches.
From a historical perspective, fire has been a major factor in
shaping tropical forest communities. Slash and burn agriculturists
and even nomadic peoples in tropical forested areas have had an
impact on forest communities by creating a mosaic pattern of
different aged forest seres. These disturbances have kept, and
continue to keep, these forests in a state of non-equilibrium. From
an evolutionary perspective, fire could have helped to shape these
forest communities without human participation. Forests in this
region are semi-deciduous and leaf litter is deep at the end of the
annual drought season. Lightning storms are frequent at the onset of
the rainy season. It is not difficult to imagine lightning as the spark
needed to set off a major forest fire in stands that have deep litter.
Small mammals, although mobile organisms, are probably unable
to flee the destruction brought about by intense wildfire (Cook, 1959)
compared to larger and more vagile mammals. In forests that are
burned by intense crown fires, small mammals are forced to
recolonize burned areas from adjacent undisturbed habitat.
Recolonization time is dependent upon the areal extent of the fire
and the progession of the vegetational succession, as well as
competitive interactions between the species. In forests that are
burned in an intermediate fashion, but in an intense manner,
remaining local mosaics of pre-existing habitat could be local pools of
founder stock for small mammal recolonization. These forest mosaics
would be prime habitat for pioneering species that occur in the

157
undisturbed habitat or nearby early succession seres unaffected by
fire. It is intuitive that mosaic habitats would offer a wide array of
available habitat for species with differing needs.
Small mammals are largely unaffected by fire regimes, except
during, and immediately after, the burn. Rather, their abundance and
distributional changes associated with fire should be related to the
vegetational changes brought about by the fire. There are several
studies on the effect of fire on small mammal communities in
Australia (for a thorough treatise, see Fox and McKay, 1981). The
majority of these studies focus on the recolonization time and species
replacement pattern of small mammals in succession seres following
forest fire. These studies indicate that early colonizing or pioneer
species usually increase their numbers rapidly during the initial phase
of succession, overlap with later colonizing species in secondary
succession seres, and decrease in number as secondary succession
moves to early primary forest. Catling and Newsome (1981) propose
that species diversity should be highest in the intermediate fire-
prone forest in Australia.
Species diversity of mammals increased during the first 25 years
then decreased as the forest canopy closed following forest fires in
Yellowstone National Park (Taylor, 1972). Cook (1959) observed a
change in small mammal communities from grassland to brushland
species as succession evolved following a fire.
Several authors have examined the effect of succession after
disturbance not caused by fire on small mammals with varying results.
Isabirye-Basuta and Kasenene (1987) found a higher species diversity

158
of small mammals in a selectively logged forest than primary forest in
Uganda. Huntly and Inouye (1987) found no correlation between
species diversity of small mammals and time since last disturbance in
old field habitats. Barry (1984) reported a higher species diversity in
primary forest versus a forest mosaic of secondary and primary forest
in Australia.
This study focused on the species diversity of small mammals in
fire shaped forested habitats in the Atlantic forest of Brazil. Species
diversity of small mammals in grassland habitats probably is lower
than that in forested habitats (Anthony et al., 1981; Stallings, in
prep.; August, 1984). Vegetational structure in grassland habitat is
not as complex as forested habitat and these habitats are usually
dominated by one or two species. It appears that species diversity in
grasslands is more affected by density dependent factors rather than
vegetative structure. My data suggest that density independent
factors (variation in epiphyte abundance and sum of basal area) may
affect species diversity of small mammals rather than density
dependent factors (Chapter IV). The two variables that comprise the
species diversity index (H’) are the number of species and the number
of individuals of each species. There were five species of small
mammals that occurred in all habitat types, and three that did not
differ in abundance. Most notably, three didelphid marsupials,
Metachi rus. Mamosa incana and M. cinerea. were observed to have a
ubiquitous distribution and occur in high densities in all habitats. In
a previous analysis (Chapter IV), I proposed that these species are
able to dominate the foreste communities because of their generalized

159
diets. The marsupial Didelphis and the cricetine rodent 0. trinitatus
also occurred in all habitats but at varying densities. Several species
were represented by low capture rates, such as the rodents
Rhipidomvs. Oxvmvcterus. and Abrawavomvs and the marsupial
Marmosa microtarsus. These species augmented the species list in the
two mosaic habitats and in spite of their low proportions, increased
the H’ values.
As shown in this study, species diversity of small mammals
increased with increasing patchiness. The increase in species
diversity was primarily due to the addition of more terrestrial rodents
in patchy areas. Secondary and primary forested habitats, which
were not patchy, had depauperate faunas compared to the mosaic
habitats. Besides the usual 3 didelpid marsupials, the secondary
habitat favored terrestrial rodents, while the primary habitats favored
the more arboreal rodents and marsupials.
It is intuitive that there should be a linear relationship between
species diversity and intensity of disturbance. Obviously, the more
complete the disturbance the less patchy the resources. The data
presented in this study support Connell’s intermediate disturbance
hypothesis for small mammals in a tropical forest.

CHAPTER VI
CONCLUSIONS AND SYNTHESIS
The salient results of this study emphasize the high level of
species diversity and endemism of small mammals in the Atlantic
Forest of eastern Brazil. Davis (1945) and Fonseca (pers. comm.) also
reported similar findings in Rio de Janeirio and Minas Gerais states,
respectfully. These findings support the notion of a high level of
endemism suggested for other taxa in the Atlantic Forest (e.g.,
primates, Mittermeier et al., 1982; Kinzey, 1982; trees, Mori et al.,
1981; reptiles, Muller, 1973; birds, Haffer, 1974).
One striking difference between this study and other small
mammal studies conducted in the neotropics is the high number of
marsupial captures relative to rodent captures. This finding is
consistent with Simpson’s (1980) observation that South America
should also be considered the "land of the marsupials" as is Australia.
Species richness of rodents was clearly numerically superior to that
of marsupials, however, marsupial captures accounted for
approximately 80% of total captures across all forested habitats
sampled. The grass or wet/meadow habitat was the only community
that was dominated by more species and more individuals of rodents
relative to marsupials.
160

161
Several studies have demonstrated the importance of
microhabitat, use of space, and competition in structuring small
mammal communities in Neotropical forests and grasslands (Nitikman
and Mares, 1987; Lacher and Alho, in press; August, 1983, 1984). The
findings in this study did nojt support the notion that the small
mammals were selecting microhabitat from the available habitat. Use
of vertical space, dietary classification, and body size appear to play
important roles in the manner in which these small mammals partition
the available resources.
The findings of this study suggest that environmental instability
promotes species diversity of small mammals in an area subject to
frequent forest fire. Areas burned in a heterogeneous fashion
supported a higher species diversity than in areas burned in a
homogeneous fashion or in areas of primary forest. These results are
in accord with the suggestions of Connell (1978) and do not support
the notion that primary forests support a higher species diversity.
None of the species captured during the study appeared to be
obligate or "climax" adapted species. Primate species richness and
abundance also yielded similar results. This is not to say that no
species depend on primary forest for their survival. It is probably
true that a wide array of species from different taxonomic groups are
dependent on primary forest stands.
The vast amount of secondary growth in the eastern Atlantic
forest probably is an important variable in determining the species
composition and abundances of small mammals. Charles-Dominique
(1983) hypothesized that second growth is optimal habitat for

162
dldelphid marsupials because this forest type provides a steadier
supply of food resources. At high densities, didelphid marsupials can
out compete forest rodents for food because marsupials are basically
omnivorous. These hypotheses help to explain the pattern of species
diversity in the region and the relative high abundance of marsupials.
Deforestation brought about by human activities in the recent past
account for the majority of disturbance to once forested habitats.
However, forest fire could have also played a role in helping to
maintain forest mosaics of secondary and primary stands. This
hypothesis does not require human participation. The subtropical
forests of this region are prone to forest fire because of a
pronounced dry season. Most trees are deciduous and semi-deciduous
and leaf litter is deep at the end of the drought period. Lightning
storms occur at high frequency at the onset of the rainy season and
forest fires could be caused by lightning strikes.
The results of this study provide a theoretical base for
developing conservation strategies in this highly threatened
ecosystem. Currently, much attention is directed to the fate of the
world’s tropical rainforests and maintaining the biotic diversity that
occurs within these forests. In areas of advancing agricultural and
industrial development, many parks and reserves have become islands
surrounded by mono-cultural plantations or large expanses of
urbanization. It is clear that these reserves are the future stock of
our wild species, but in many instances, their maintenance will
require active, rather than passive, management.

163
The Rio Doce State Forestry Park is an ideal model of a
tropical park that is managed by passive techniques. The Park has
experienced intense crown fires in the past because of passive
management. Large amounts of accumulated litter have been
responsible for the intense fires. Current fire policy in Brazil, and
in many other Latin American countries, is fire suppression and
prevention rather than adopting a policy that fire is natural and
using fire as a management tool. Control burns are currently used in
the United States National Parks and Forests as a tool to remove
excess and potential fuel, to set back successional evolution, and to
maintain the forests in different aged stands. This policy has
maintained forest litter at manageable levels and has increased the
species diversity of the flora and fauna.
I do not advocate the use of fire in all systems. Some forests
may not respond well to the use of fire and may alter the forest
composition for centuries. Species have evolved to fill available
niches in primary forests and these species depend on this habitat for
their survival. By destroying the primary forest in an intense crown
fire, these species may be lost forever because of the long
regeneration time of climax adapted forest species. Seeds stored in
the ground and all trees will most probably be destroyed in forests
that are burned to the ground in intense fires. These fires probably
hinder the pre-existing trees to coppice. These areas can only be
recolonized from outside sources and the resulting "climax'' forest will
be greatly different in composition from the previous forest. Fires of
an intermediate nature are probably of most use in setting succession

164
in tropical forests. The result of these practices allow pioneer, mid
successional and climax adapted species to co-exist in protected
areas, while maintaining and maximizing species diversity.

APPENDIX

166
Table A-1. Preliminary check list of the non-volant mammal
species that probably occur in the Atlantic Forest region of
Brazil per state. *= endemic species, += endemic genus, ?=
occurrence uncertain. References available from author upon
request. State codes: A= Bahia, B= Minas Gerais, C= Espirito
Santo, D= Rio De Janeiro, E= Sao Paulo.
ORDER/SPECIES
STATES
A
BCD
E
MARSUPIAUA
Caluromvs philander
X
X
MonodelDhis americana*
X
X
X
X
X
M. dimidiata
X
X
X
X
M. domestica
X
X
X
X
X
M. henseli
X
M. iherinqi*
X
X
M. scaloDS*
X
X
M. sorex*
X
X
X
X
M. theresa*
X
M. touan
X
X
X
M. unistriata*
X
Marmosa anil is*
X
X
M. cinerea
X
X
X
X
X
M. incana*
X
X
X
X
M. microtarsus*
X
X
X
M. murina
X
X
M. scaDulata*
X
M. velutina
X
Philander oDossum
X
X
X
X
X
Metichirus nudicaudatus
X
X
X
X
Lutreolina crassicaudata
X
X
DidelDhis marsuDialis
X
X
X
X
X
D. albiventris
X
PRIMATES
Callithrix iacchus*
X
C. qeoffrovi*
X
X
C. aurita*
X
X
X
C. flaviceDS*
X
X
C. kuhlii*
X
LeontODithecus+ rosalia*
X
L. chrYSODvqus*
X
L. chrvsomelas*
X
Callicebus Dersonatus*
X
X
X
X
X
Cebus apella
X
X
X
X
X

ORDER/SPECIES
STATES
ABODE
Alouatta fusca*
Brachvteles+ arachnoides*
EDENTATA
Mvrmecophaqa tridactvla
Tamandúa tetradactvla
Bradvpus variegatus
B. torauatus*
Priodontes maximus
Cabassous tatouav
C. unicinctus
Euphractus sexcinctus
Dasvpus nóvemeinctus
D. septemcinctus
CARNIVORA
Dusicvon gvmnocercus
D. thous
Speothos venaticus
Procvon cancrivorus
Nasua nasua
Potos flavus
Galictis vittata
G. cu.ia
Conepatus chinga
C. semistriatus
Eira barbara
Lutra longicaudis
Pteronura brasi1iensis
Felis pardal is
F. tigrina
F. concolor
F. yagouaroundi
F. wiedii
Pantera onca
PERISSODACTYLA
Tapi rus terrestris
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 XX
X X X X X
X X X X
XX 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 X X X
X X X X X
X X X X
X

168
Table A-1 - continued
ORDER/SPECIES STATES
A B C D E
ARTIODACTYLA
Tavassu ta.iacu X
I. pécari X
Ozotoceros bezoarticus X
Mazama americana X
M. gouazoubira X
M. rufina?
LAGOMORPHA
Svlvilagus brasi1iensis X
RODENTIA
Sciurus (aestuans) alphonsei* X
S. (aestuans) garbei* X
S. (aestuans) ingrami* X
Orvzomvs nigripes (=e1iurus) X
0. nitidus X
0. ratticeps
0. subflavus
Oecomvs cinnamomeus* (=trinitat1s?) X
Nectomvs sauamipes X
Rhipidomvs maculipes* X
R. mastacalIs*
Thomasomvs dorsal is*
I. pictipes?
Phaenomvs+ ferrugineus* X
Rhagomvs+ rufescens*
Bolornvs lasiurus X
Abrawavaomvs+ ruschii*
Akodon serrensis
A. nigrita X
A. arviculoides X
Bibimvs labiosus*
Oxymycterus angularis*
0. hisoidus* X
0. auestor*
0. roberti
0. roste 1latus* X
0. rutiIans X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
XXX
XXX
XXX
XXX
XXX
X
XXX
X
X
XXX
X X
XXX
X X
XXX
X
XXX
X
X
X X
X
X
X X
XXX
XXX
X X
X

169
Table A-1 - continued
ORDER/SPECIES
STATES
ABODE
Calomvs callosus
C. laucha
C. leucodactvlus?
Blarinomvs+ breviceps*
Wiedomvs pvrrhorhinos
Holochilus brasi1iensis
H. phvsodes*
H. russatus*
H. sciureus
H. wagneri*
Microxus iherngi?
Prcechimvs albispinus*
P. dimidiatus*
P. iheringi*
P. mvosurus*
P. setosus*
Eurvzvgomatomvs spinosus
Clvomvs+ latíceps^
Trichomvs apereoides
Isotrix pictus*
Echimvs blainvi11 el *
E. nigrispinus*
Kannabateomvs amblyonvx
Galea spixi1
Cavia aperea
C. fulgida*
Hvdrochaeris hvdrochaeris
Dasvprocta aguti
JD. prvmnolopha
Agouti paca
Chaetomvs subspinosus
Coendou prehensilis
X
X
XX 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
XX X
X
XX 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
TOTALS:
GENERA.
131 SPECIES; 50
ENDEMIC SPECIES; 63 GENERA; 7 ENDEMIC

170
|3
Table A-2. Standard body (MM) and mass (G) measurements for
species of small mammals captured in Rio Doce State Forestry Park,
Minas Gerais, Brazil. Measurements are separated for both sex and
age. N=sample size, MIN.=minimum measurement, MAX.=maximum
measurement, MEAN + STD. DEV.=arithmetic mean and one standard
deviation. Significant differences in measurements between sexes
for each species are designated by * (p < 0.05) and ** (p <
0.001).
DidelDhis marsuDialis
ADULTS, HIND FOOT > 51 MM.
MALES N MIN.
MAX.
MEAN + !
3TD. DEV.
MASS
9
578.00
1300.00
938.89
+
295.32
BODY
9
292.00
415.00
354.78
+
39.98
TAIL
9
312.00
393.00
355.33
+
28.41
EAR
9
47.00
55.00
50.67
+
2.78
FOOT
9
51.00
63.00
57.89
+
4.20
FEMALES
MASS
8
568.00
1855.00
1158.62
+
369.98
BODY
8
335.00
400.00
372.87
+
24.82
TAIL
8
345.00
400.00
377.25
+
18.01
EAR
8
46.00
58.00
51.37
+
3.93
FOOT
8
51.00
64.00
57.87
+
4.97
JUVENILES,
HIND
FOOT < 51
MM.
MALES
MASS
7
233.00
606.00
386.71
+
157.02
BODY
7
192.00
310.00
256.86
+
45.16
TAIL
7
236.00
333.00
277.14
+
38.33
EAR
7
41.00
51.00
45.29
+
3.20
FOOT
7
40.00
51.00
45.29
+
4.54
FEMALES
MASS
1
447.00
447.00
447.00
BODY
1
300.00
300.00
300.00
TAIL
1
380.00
380.00
380.00
EAR
1
50.00
50.00
50.00
FOOT
1
49.00
49.00
49.00

171
Table A-2 - continued
Metachi rus nudicaudatus
ADULTS, MASS > 90 G.
MALES
N
MIN.
MAX.
MEAN + STD. DEV
MASS*
35
102.00
480.00
281.51 +
117.00
BODY
35
170.00
300.00
233.89 +
36.76
TAIL
35
227.00
373.00
307.66 +
41.67
EAR
35
28.00
40.00
35.40 +
2.66
FOOT**
35
35.00
52.00
43.71 +
3.86
FEMALES
MASS
51
91.00
345.00
235.88 +
68.93
BODY
50
150.00
265.00
222.93 +
28.56
TAIL
51
178.00
363.00
297.96 +
43.19
EAR
50
31.00
43.00
35.72 +
2.62
FOOT
JUVENILES,
51
MASS < 90 G.
34.00
47.00
41.35 ±
2.96
MALES
MASS
6
37.00
81.00
63.83 +
19.34
BODY
6
115.00
160.00
142.00 +
17.44
TAIL
6
164.00
223.00
194.67 +
24.34
EAR
6
23.00
32.00
28.67 +
3.27
FOOT
6
27.00
34.00
31.33 +
2.73
FEMALES
MASS
5
60.00
88.00
76.60 +
10.90
BODY
5
140.00
170.00
157.80 +
10.92
TAIL
5
171.00
231.00
209.60 +
24.27
EAR
5
28.00
35.00
31.20 +
2.77
FOOT
5
30.00
34.00
32.60 +
1.52

172
Table A-
-2 - continued
Marmosa
incana
ADULTS,
MASS > 35 G.
MALES
N
MIN.
MAX.
MEAN +
STD.
DEV.
MASS
46
35.00
130.00
66.04
+
27.74
BODY
46
95.00
192.00
138.00
+
21.96
TAIL
45
162.00
296.00
195.09
+
21.52
EAR»*
46
24.00
32.00
27.52
+
1.96
FOOT**
46
18.00
26.00
21.52
+
1.72
FEMALES
MASS
22
44.00
73.00
58.64
+
8.32
BODY
22
110.00
162.00
137.50
+
11.90
TAIL
22
163.00
199.00
183.41
+
8.62
EAR
22
23.00
29.00
25.86
+
1.49
FOOT
22
16.00
24.00
19.82
+
1.79
JUVENILES,
MASS <
35 G.
MALES
MASS
13
14.00
34.00
29.69 +
5.85
BODY
13
82.00
120.00
105.46 +
11.18
TAIL
12
111.00
169.00
155.17 +
16.13
EAR
13
19.00
27.00
23.92 +
2.25
FOOT
13
16.00
22.00
19.62 ±
1.56
FEMALES
MASS
20
16.00
34.00
25.70 +
5.38
BODY
20
81.00
120.00
103.40 +
10.94
TAIL
20
115.00
176.00
151.50 +
17.34
EAR
20
17.00
29.00
23.85 +
2.68
FOOT
20
16.00
21.00
18.35 +
1.27

173
Table A-2 - continued
Marmosa cinerea
ADULTS,
MASS > 50 G.
MALES
N
MIN.
MAX.
MEAN + STD. DEV
MASS
36
56.00
194.00
109.94 + 28.75
BODY
36
146.00
210.00
176.89 ± 16.08
TAIL*
35
200.00
291.00
259.63 ± 20.42
EAR*
36
24.00
35.00
30.81 + 2.49
FOOT*
36
24.00
31.00
28.19 ± 1.62
FEMALES
MASS
28
53.00
230.00
99.07 ± 41.40
BODY
28
125.00
205.00
165.68 + 23.09
TAIL
28
192.00
293.00
248.68 + 26.41
EAR
28
24.00
34.00
29.14 + 2.77
FOOT
28
22.00
35.00
26.64 + 2.57
JUVENILES,
MASS
< 50 G.
MALES
MASS
3
20.00
48.00
31.00 + 14.93
BODY
3
100.00
140.00
114.67 + 22.03
TAIL
3
155.00
196.00
170.33 + 22.37
EAR
3
22.00
26.00
24.00 + 2.03
FOOT
3
21.00
26.00
22.67 + 2.89
FEMALES
MASS
13
19.00
49.00
32.77 + 10.94
BODY
13
95.00
150.00
114.38 + 16.97
TAIL
13
149.00
203.00
176.85 + 17.69
EAR
13
18.00
28.00
23.23 + 2.52
FOOT
13
21.00
27.00
23.08 + 2.18

174
Table A-2 - continued
Marmosa microtarsus
ADULTS
MALES
N
MIN.
MAX.
MEAN + STD. DEV.
MASS
1
31.00
31.00
31.00
BODY
1
106.00
106.00
106.00
TAIL
1
148.00
148.00
148.00
EAR
1
14.00
14.00
14.00
Foot
1
17.00
17.00
17.00
Caluromvs
Dhilander
ADULTS
MALES
N
MIN.
MAX.
MEAN + STD. DEV
MASS
18
123.00
261.00
189.83 + 35.72
BODY
18
180.00
245.00
215.33 + 17.08
TAIL
18
225.00
322.00
295.72 ± 22.15
EAR
18
31.00
38.00
33.67 + 1.94
FOOT
18
32.00
41.00
35.61 + 2.00
FEMALES
MASS
12
115.00
286.00
186.75 + 44.67
BODY
12
200.00
240.00
216.75 ± 12.27
TAIL
12
283.00
313.00
299.83 ± 9.68
EAR
12
30.00
37.00
33.42 ± 2.02
FOOT
12
30.00
42.00
35.00 + 2.92

175
Table A-2
- continued
Nectomvs
sauamiDes
ADULTS, MASS > 60
G.
MALES
N
MIN.
MAX.
MEAN +
STD. DEV
MASS
8
’75.00
235.00
146.12
+
64.97
BODY
8
138.00
203.00
173.00
+
24.63
TAIL
8
160.00
302.00
199.75
+
47.06
EAR
8
20.00
24.00
22.00
+
1.51
FOOT
8
42.00
53.00
47.87
+
3.52
FEMALES
MASS
4
98.00
217.00
160.50
+
57.97
BODY
4
165.00
200.00
183.00
+
17.11
TAIL
4
188.00
231.00
206.75
+
20.35
EAR
4
21.00
24.00
23.00
+
1.41
FOOT
4
46.00
50.00
48.00
+
1.83
JUVENILES
, MASS <
60 G.
MALES
MASS
1
31.00
31.00
31.00
BODY
1
95.00
95.00
95.00
TAIL
1
108.00
108.00
108.00
EAR
1
18.00
18.00
18.00
FOOT
1
35.00
35.00
35.00
RhiDidomvs mastacalis
ADULTS
MALES
N
MIN.
MAX.
MEAN + STD.
DEV.
MASS
1
80.00
80.00
80.00
BODY
1
130.00
130.00
130.00
TAIL
1
156.00
156.00
156.00
EAR
1
22.00
22.00
22.00
FOOT
1
27.00
27.00
27.00
JUVENILES
FEMALES
MASS
1
39.00
39.00
39.00
BODY
1
104.00
104.00
104.00
TAIL
1
142.00
142.00
142.00
EAR
1
18.00
18.00
18.00
FOOT
1
28.00
28.00
28.00

176
Table A-
-2 - continued
Akodon i
cursor
ADULTS,
MASS >
25 G.
MALES
N
MIN.
MASS
73
25.00
BODY*»
73
83.00
TAIL*
65
75.00
EAR
71
14.00
FOOT
72
21.00
FEMALES
MASS
38
26.00
BODY
38
76.00
TAIL
38
76.00
EAR
37
16.00
FOOT
38
23.00
JUVENILES, MASS
< 25 G.
MALES
N
MIN.
MASS
25
7.00
BODY
26
51.00
TAIL
25
47.00
EAR
26
10.00
FOOT
26
21.00
FEMALES
MASS
8
11.00
BODY
8
66.00
TAIL
8
64.00
EAR
8
11.00
FOOT
8
22.00
MAX. MEAN + STD. DEV.
66.00
41.18
+
10.55
123.00
104.82
+
9.24
108.00
93.49
+
7.65
22.00
18.62
+
1.46
30.00
26.06
+
1.38
66.00
38.53
+
10.04
120.00
99.53
+
10.99
103.00
89.58
+
7.02
20.00
18.30
+
1.02
30.00
26.00
+
1.47
MAX. MEAN + STD. DEV.
24.00
18.60
+
4.53
102.00
81.85
+
10.26
89.00
72.28
+
9.66
18.00
16.12
+
1.75
27.00
24.23
+
1.70
24.00
19.62
+
4.21
95.00
84.25
+
9.45
85.00
77.00
+
6.52
19.00
16.00
+
2.45
27.00
24.50
+
1.69

177
Table A-
-2 - continued
Orvzomvs caDito
ADULTS,
MASS > 35
G.
MALES
N
MIN.
MAX.
MEAN +
STD. DEV
MASS
5
38.00
72.00
59.60
+
13.81
BODY
5
118.00
135.00
127.60
+
8.14
TAIL
5
124.00
145.00
132.00
+
8.69
EAR
5
16.00
24.00
21.00
+
3.16
FOOT
5
23.00
35.00
30.80
+
4.55
FEMALES
MASS
10
50.00
65.00
59.10
+
5.22
BODY
10
110.00
147.00
124.40
+
10.86
TAIL
10
115.00
135.00
125.40
+
6.15
EAR
9
21.00
22.00
21.44
+
0.53
FOOT
10
30.00
34.00
31.80
+
1.32
JUVENILES
MALES
MASS
2
21.00
22.00
21.50
+
0.71
BODY
2
85.00
90.00
87.50
+
3.54
TAIL
2
92.00
93.00
92.50
+
0.71
EAR
2
18.00
20.00
19.00
+
1.41
FOOT
2
27.00
28.00
27.50
+
0.71

178
Table A-2 - continued
Oecomvs trinitatis
ADULTS
MALES
N
MIN.
MASS
10
42.00
BODY
10
114.00
TAIL
10
133.00
EAR
9
16.00
FOOT
10
13.00
FEMALES
MASS
5
61.00
BODY
5
124.00
TAIL
5
129.00
EAR
4
18.00
FOOT
5
28.00
MAX. MEAN + STD. DEV.
100.00
65.50
+
16.95
135.00
124.60
+
7.53
155.00
147.40
+
7.99
25.00
18.67
+
2.78
39.00
29.50
+
8.66
95.00
76.20
+
12.87
143.00
131.80
+
7.79
236.00
166.60
+
42.07
21.00
19.00
+
1.41
33.00
30.60
+
2.07
Orvzomvs
ADULTS
MALES
niqriDes
N MIN.
MAX.
MEAN +
STD. DEV
MASS
2
25.00
28.00
26.50
+
2.12
BODY
2
83.00
100.00
91.50
+
12.02
TAIL
2
122.00
132.00
127.00
+
7.07
EAR
2
16.00
17.00
16.50
+
0.71
FOOT
2
22.00
26.00
24.00
+
2.83
FEMALES
MASS
2
7.00
22.00
14.50
+
10.61
BODY
2
47.00
85.00
66.00
+
26.87
TAIL
2
83.00
115.00
99.00
+
22.63
EAR
2
15.00
15.00
15.00
+
0.00
FOOT
2
20.00
24.00
22.00
+
2.83

179
Table A-
-2 -
continued
Orvzomys subflavus
ADULTS,
MASS
> 40 G.
MALES
N
MIN.
MAX.
MEAN + STD. DEV.
MASS
10
54.00
101.00
77.90 + 17.64
BODY
9
118.00
170.00
144.00 + 16.15
TAIL
9
147.00
193.00
172.67 + 16.40
EAR
9
22.00
26.00
23.78 ± 1.30
FOOT
9
34.00
38.00
35.33 + 1.22
FEMALES
MASS
12
58.00
135.00
88.58 + 22.22
BODY
11
114.00
170.00
146.00 ± 19.28
TAIL
11
140.00
200.00
178.09 + 17.48
EAR
11
20.00
27.00
23.82 + 2.32
FOOT
11
27.00
37.00
34.45 ± 2.77
JUVENILES, MASS < 40 G.
FEMALES
N
MIN.
MAX.
MEAN + STD. DEV.
MASS
5
6.00
35.00
23.00 + 10.51
BODY
6
50.00
110.00
90.83 ± 20.88
TAIL
5
92.00
131.00
117.00 + 14.95
EAR
6
13.00
22.00
19.67 + 3.61
FOOT
5
22.00
32.00
28.60 + 3.85
Abrawavaomvs
ruschii
ADULTS
MALES
N
MIN.
MAX.
MEAN + STD. DEV.
MASS
1
63.00
63.00
63.00
BODY
1
128.00
128.00
128.00
TAIL
1
146.00
146.00
146.00
EAR
1
20.00
20.00
20.00
FOOT
1
31.00
31.00
31.00

180
Table A-
-2 -
continued
Calomvs
laucha
ADULTS
FEMALES
N
MIN.
MAX.
MEAN +
STD. DEV.
MASS
1
20.00
20.00
20
.00
BODY
0
—
—
—
TAIL
1
74.00
74.00
74
.00
EAR
1
17.00
17.00
17
.00
FOOT
1
19.00
19.00
19
.00
Oxymycterus
roberti
ADULTS
MALES
N
MIN.
MAX.
MEAN +
STD. DEV.
MASS
5
45.00
120.00
73.80
+
29.19
BODY
5
94.00
150.00
125.80
+
20.20
TAIL
3
105.00
125.00
118.00
+
11.27
EAR
5
19.00
25.00
21.40
+
2.19
FOOT
5
33.00
39.00
35.00
+
2.35
FEMALES
MASS
4
82.00
110.00
93.00
+
12.62
BODY
4
145.00
245.00
173.00
+
48.19
TAIL
2
121.00
122.00
121.50
+
0.71
EAR
4
22.00
23.00
22.25
+
0.50
FOOT
4
30.00
36.00
32.50
+
2.65
Eurvzqomatomys
SDinosus
ADULTS
MALES
MASS
2
165.00
210.00
187.50
+
31.82
BODY
2
185.00
188.00
186.50
+
1.14
TAIL
2
61.00
65.00
63.00
+
2.83
EAR
2
17.00
18.00
17.50
+
0.71
FOOT
2
35.00
35.00
35.00
+
0.00

181
Table A-2 -
- continued
Cavia fulqida
ADULTS
MALES N
MIN.
MAX.
MEAN + STD. DEV.
MASS 1
285.00
285.00
285.00
BODY 1
223.00
223.00
223.00
TAIL 1
—
—
—
EAR 1
22.00
22.00
22.00
FOOT 1
46.00
46.00
46.00
FEMALES
MASS 1
280.00
280.00
280.00
BODY 1
234.00
234.00
234.00
TAIL 1
17.00
17.00
17.00
EAR 1
21.00
21.00
21.00
FOOT 1
47.00
47.00
47.00
DasYDrocta
azarae
ADULTS
MALES
MASS 1
2560.00
2560.00
2560.00
BODY 1
450.00
450.00
450.00
TAIL 1
28.00
28.00
28.00
EAR 1
45.00
45.00
45.00
FOOT 1
130.00
130.00
130.00
FEMALES
MASS 1
2056.00
2056.00
2056.00
BODY 1
460.00
460.00
460.00
TAIL 1
19.00
19.00
19.00
EAR 1
38.00
38.00
38.00
FOOT 1
120.00
120.00
120.00
JUVENILES
MALES
MASS 2
449.00
589.00
519.00 + 98.99
BODY 2
210.00
250.00
230.00 + 28.28
TAIL 2
14.00
16.00
15.00 + 1.41
EAR 2
35.00
36.00
35.50 + 0.71
FOOT 2
80.00
96.00
88.00 + 11.31

182
Table A-3. Species observed during one year of census walks in
five sampling areas in the Rio Doce State Forestry Park. Each
census walk measured 1050 M. in length and each sampling area had
60 Km. of repeat censuses. A= mean number of encounters per
month, B= mean number of animals per month, C= mean group size, D=
percentage of total encounters, and E= percentage of total
individuals. Site abbreviations explained in text.
SITE/
SPECIES
A
B
C
D
E
RD/F
Callicebus Dersonatus
0.50
0.83
1.66
21.4
22.2
Sciurus aestuans
0.08
0.08
1.00
3.6
2.2
Nasua nasua
0.08
0.08
1.00
3.6
2.2
Taoirus terrestris
0.08
0.08
1.00
3.6
2.2
PeneloDe obscura
1.25
2.41
2.07
57.2
64.4
RD/H
Callicebus Dersonatus
0.50
1.00
2.00
12.0
9.0
Cebus aDella
0.75
2.91
3.88
18.0
26.0
Alouatta fusca
0.80
0.25
3.00
2.0
2.2
Callithrix hvbridus
0.58
2.33
4.00
14.0
21.0
Sciurus aestuans
0.58
1.33
2.28
14.0
12.0
Taoirus terrestris
0.08
0.08
1.00
2.0
0.7
DasvDrocta azarae
0.42
0.50
1.20
10.0
4.4
PeneloDe obscura
0.25
0.75
3.00
6.0
6.6
Nothura sd.
0.08
0.83
10.00
2.0
7.4
CrvDturellus sd.
0.83
1.25
1.50
2.0
11.1
RD/M
Callicebus Dersonatus
0.75
2.20
2.90
21.4
24.8
Cebus aDella
0.25
1.33
5.33
7.1
15.2
Callithrix a. Detronius
0.08
0.08
1.00
2.4
1.0
Brachyteles arachnoides
0.17
1.5
9.0
4.7
17.4
Sciurus aestuans
0.25
0.25
1.00
7.1
2.8
Nasua nasua
0.08
0.08
1.00
2.4
1.0
DasvDrocta azarae
0.25
0.25
1.00
7.1
2.8
PeneloDe obscura
1.16
2.50
2.14
33.3
28.6
Nothura sd.
0.08
0.16
2.00
2.4
2.0
CrvDturellus sd.
0.16
0.16
1.00
4.7
2.0
Tinamus solitarius
0.16
0.16
1.00
4.7
2.0
OdontoDhorus sd.
0.08
0.08
1.00
2.4
1.0

183
Table A-3 - continued
SITE/SPECIES
A
RD/T
Cebus apella
0.33
Callithrix a. Detronius
0.08
Nasua nasua
0.16
Dasvprocta azarae
0.66
EuDhractus sexcinctus
0.08
Tapirus terrestris
0.08
Penelope obscura
0.33
Nothura sd.
0.25
CrvDturellus sd.
0.16
RD/C
Cebus aDella
0.50
Callithrix a. Detronius
0.08
Sciurus aestuans
0.08
DasvDrocta azarae
0.25
Mazama sd.
0.08
PeneloDe obscura
0.75
Nothura sd.
0.08
CrvDturellus sd.
0.66
Tinamus solitarius
0.08
B C D E
1.50
4.50
15.4
27.7
0.42
5.00
4.0
7.7
0.33
2.00
7.7
6.2
0.66
1.00
30.7
12.3
0.08
1.00
4.0
1.5
0.08
1.00
4.0
1.5
0.58
1.75
15.4
10.7
1.58
6.33
11.5
29.2
0.16
1.00
7.7
3.0
2.00
4.00
19.4
39.3
0.16
2.00
3.2
3.3
0.08
1.00
3.2
1.6
0.25
1.00
9.7
4.9
0.08
1.00
3.2
1.6
1.58
2.11
29.0
31.1
0.08
1.00
3.2
1.6
0.75
1.13
25.8
14.8
0.08
1.00
3.2
1.6

184
Table A-4. Collection of bats from the Rio Doce State Forestry
Park, Minas Gerais, Brazil. * Species captured from Rio Casca,
approximately 50 Km. east of the park. + Captured in forest, a =
captured in man made structures, b = captured along water edge, c
= common, r = rare.
FAMILY/SPECIES
Embaílonuridae
Rhvnchonvcteris naso*br
Nocti1ionidae
Noctilio leporinusbc
Phyllostomatidae
Carol lia persoicilata+abc
Anoura caudiferae
Glossophaga soricinaac
Artibeus 1iteratus+c
Artibeus iamaicensis+r
Chrotooterus aurita+r
Sturnira 1 i 1ium+c
Micronvcteris sp.+r
Phyllostomus hastatus*r

185
Table A-5. Capture results from eucalypt forest with native
forest subcanopy. RECAP INDEX= total captures/first captures and
represents the average number of times that an individual of
species X is captured. Numbers in parentheses represent percent
of contribution of capture per species per taxonomic group.
Species abbreviations are explained in Table 2-4.
TOTAL FIRST RECAP
SPECIES
CAPTURES
% TOTAL
CAPTURES
% TOTAL
INDEX
MARSUPIALS
DM
7
4.4 ( 5.3)
3
5.6 ( 8.3)
2.3
MN
18
11.4 (13.7)
9
16.7 (25.0)
2.0
MI
14
8.7 (10.7)
8
14.8 (22.2)
1.8
MC
77
48.7 (58.8)
10
18.5 (27.8)
7.7
CP
15
9.6 (11.5)
6
11.1 (16.7)
2.5
131
82.8(100.0)
36
66.7(100.0)
RODENTS
AC
25
15.8 (92.6)
16
29.6 (88.9)
1.6
OC
2
1.4 ( 7.4)
2
3.7 (11.11
1.0
27
17.2(100.0)
18
33.3(100.0)

186
Table 2-10. Capture results from wet meadow site in Rio Doce
State Forestry Park, Minas Gerais, Brazil. RECAP INDEX= total
captures/first captures and represents the average number of times
an individual of species X was captured. Numbers in parentheses
represent percent contribution of captures per species per
taxonomic group. Species abbreviations are explained in Table 2-
4.
TOTAL
SPECIES
CAPTURES
% TOTAL
MARSUPIALS
MI
3
0.8 (50.0)
MC
1
0.3 (16.7)
CP
2
0.5 (33.3)
6
1.6(100.0)
RODENTS
NS
5
1.3 ( 1.4)
AC
315
84.5 (85.8)
OC
1
0.3 ( 0.3)
OS
31
8.3 ( 8.4)
ON
4
1.0 ( 1.1)
CL
4
1.0 ( 1.1)
OR
4
1.0 ( 1.1)
ES
2
0.5 ( 0.5)
373
98.4(100.0)
FIRST RECAP
CAPTURES
% TOTAL
INDEX
2
1.3 (50.0)
1.5
1
0.7 (25.0)
1.0
L
0.7 (25.0)
2.0
4
2.7(100.0)
4
2.7
( 2.7)
1.3
105
70.0
(71.9)
3.0
1
0.7
( 0.7)
1.0
22
14.7
(15.1)
1.4
4
2.7
( 2.7)
1.0
4
2.7
( 2.7)
1.0
4
2.7
( 2.7)
1.0
2
1.3
( 1.4)
1.0
150
97.3(100.0)

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BIOGRAPHICAL SKETCH
Jody R. Stallings was born in Marion, Illinois, on January 29,
1954. He received his Bachelor of Science degree in wildlife biology
from Murray State University, Murray, Kentucky, in 1977. In 1978,
Mr. Stallings accepted the position of Wildlife Specialist with the
Smithsonian Institution/Peace Corps Environmental Program and was
stationed in Paraguay. He served in that capacity until early 1982.
In 1982, Mr. Stallings enrolled as a graduate student in the Center
for Latin American Studies at the University of Florida. He received
his Master of Arts degree in 1984. In 1984, Mr. Stallings began his
doctoral studies in the Department of Wildlife and Range Sciences at
the same university.
200

I certify that I have read this study and that in my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, in scope and quality, as a
dissertation for the degree of Doctor of Philosopher^
John Gr. Robinson, Chairman
Assoc/ate Professor of Forest
/Resources and Conservation
I certify that I have read this study and that 1n my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, in scope and quality, as a
dissertation for the degree of Doctor of Philosophy.
JoTm^g. Elsenberg
Katharine Ordway Professor
Ecosystem Conservation
I certify that I have read this study and that in my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, 1n scope and quality, as a
dissertation for the degree of Doctor of Philosophy.
L'O .K I
Mai
Wayne R.\ Marlon
Associate Professor of Forest
Resources and Conservation

I certify that I have read this study and that in my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, in scope and quality, as a
dissertation for the degree of Doctor of Philosophy.
Assistant Professor of Forest
Resources and Conservation
I certify that I have read this study and that in my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, in scope and quality, as a
dissertation for the degree of Doctor of Philosophy.
X- ^ v f -
Nigel J. H. Smith
Professor of Geography
This dissertation was submitted to the Graduate Faculty of
the School of Forest Resources and Conservation in the College
of Agriculture and to the Graduate School and was accepted as
partial fulfillment of the requirements for the degree of
Doctor of Philosophy.
August, 1988
Director, Forest
Conservation
Resources
Dean, Graduate School




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7DEOH $ FRQWLQXHG 0HWDFKL UXV QXGLFDXGDWXV $'8/76 0$66 0$/(6 1 0,1 0$; 0($1 67' '(9 0$66r %2'< 7$,/ ($5 )227rr )(0$/(6 0$66 %2'< 7$,/ ($5 )227 -89(1,/(6 0$66 s 0$/(6 0$66 %2'< 7$,/ ($5 )227 )(0$/(6 0$66 %2'< 7$,/ ($5 )227

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7DEOH $ FRQWLQXHG 0DUPRVD LQFDQD $'8/76 0$66 0$/(6 1 0,1 0$; 0($1 67' '(9 0$66 %2'< 7$,/ ($5}r )227rr )(0$/(6 0$66 %2'< 7$,/ ($5 )227 -89(1,/(6 0$66 0$/(6 0$66 %2'< 7$,/ ($5 )227 s )(0$/(6 0$66 %2'< 7$,/ ($5 )227

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7DEOH $ FRQWLQXHG 0DUPRVD FLQHUHD $'8/76 0$66 0$/(6 1 0,1 0$; 0($1 67' '(9 0$66 %2'< s 7$,/r s ($5r )227r s )(0$/(6 0$66 s %2'< 7$,/ ($5 )227 -89(1,/(6 0$66 0$/(6 0$66 %2'< 7$,/ ($5 )227 )(0$/(6 0$66 %2'< 7$,/ ($5 )227

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7DEOH $ FRQWLQXHG 0DUPRVD PLFURWDUVXV $'8/76 0$/(6 1 0,1 0$; 0($1 67' '(9 0$66 %2'< 7$,/ ($5 )RRW &DOXURPYV 'KLODQGHU $'8/76 0$/(6 1 0,1 0$; 0($1 67' '(9 0$66 %2'< 7$,/ s ($5 s )227 )(0$/(6 0$66 %2'< s 7$,/ s ($5 s )227

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7DEOH $ FRQWLQXHG 1HFWRPYV VDXDPL'HV $'8/76 0$66 0$/(6 1 0,1 0$; 0($1 67' '(9 0$66 f %2'< 7$,/ ($5 )227 )(0$/(6 0$66 %2'< 7$,/ ($5 )227 -89(1,/(6 0$66 0$/(6 0$66 %2'< 7$,/ ($5 )227 5KL'LGRPYV PDVWDFDOLV $'8/76 0$/(6 1 0,1 0$; 0($1 67' '(9 0$66 %2'< 7$,/ ($5 )227 -89(1,/(6 )(0$/(6 0$66 %2'< 7$,/ ($5 )227

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7DEOH $ FRQWLQXHG $NRGRQ L FXUVRU $'8/76 0$66 0$/(6 1 0,1 0$66 %2'
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7DEOH $ FRQWLQXHG 2UY]RPYV FD'LWR $'8/76 0$66 0$/(6 1 0,1 0$; 0($1 67' '(9 0$66 %2'< 7$,/ ($5 )227 )(0$/(6 0$66 %2'< 7$,/ ($5 )227 -89(1,/(6 0$/(6 0$66 %2'< 7$,/ ($5 )227

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7DEOH $ FRQWLQXHG 2HFRPYV WULQLWDWLV $'8/76 0$/(6 1 0,1 0$; 0($1 67' '(9 0$66 %2'< s 7$,/ ($5 )227 )(0$/(6 0$66 %2'< 7$,/ ($5 )227 2UY]RPYV $'8/76 0$/(6 QLTUL'HV 1 0,1 0$; 0($1 67' '(9 0$66 %2'< 7$,/ ($5 )227 )(0$/(6 0$66 %2'< 7$,/ ($5 )227

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7DEOH $ FRQWLQXHG 2UY]RP\V VXEIODYXV $'8/76 0$66 0$/(6 1 0,1 0$; 0($1 67' '(9 0$66 %2'< 7$,/ ($5 s )227 )(0$/(6 0$66 %2'< s 7$,/ ($5 )227 s -89(1,/(6 0$66 )(0$/(6 1 0,1 0$; 0($1 67' '(9 0$66 %2'< s 7$,/ ($5 )227 $EUDZDYDRPYV UXVFKLL $'8/76 0$/(6 1 0,1 0$; 0($1 67' '(9 0$66 %2'< 7$,/ ($5 )227

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7DEOH $ FRQWLQXHG &DORPYV ODXFKD $'8/76 )(0$/(6 1 0,1 0$; 0($1 67' '(9 0$66 %2'< f§ f§ f§ 7$,/ ($5 )227 2[\P\FWHUXV UREHUWL $'8/76 0$/(6 1 0,1 0$; 0($1 67' '(9 0$66 %2'< 7$,/ ($5 )227 )(0$/(6 0$66 %2'< 7$,/ ($5 )227 (XUY]TRPDWRP\V 6'LQRVXV $'8/76 0$/(6 0$66 %2'< 7$,/ ($5 )227

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