This article was downloaded by: [Dalhousie University]On: 18 April 2013, At: 05:39Publisher: Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House,37-41 Mortimer Street, London W1T 3JH, UK

Studies on Neotropical Fauna and EnvironmentPublication details, including instructions for authors and subscription information:http://www.tandfonline.com/loi/nnfe20

Influence of edge and fire-induced changes on spatialdistribution of small mammals in Brazilian Atlanticforest fragmentsAlexandra S Pires , Fernando AS Fernandez , Daniela de Freitas & Barbara R Felicianoa Department of Ecology, Institute of Biology, Federal University of Rio de Janeiro, BrazilVersion of record first published: 18 Oct 2011.

To cite this article: Alexandra S Pires , Fernando AS Fernandez , Daniela de Freitas & Barbara R Feliciano (2005): Influenceof edge and fire-induced changes on spatial distribution of small mammals in Brazilian Atlantic forest fragments, Studies onNeotropical Fauna and Environment, 40:1, 7-14

To link to this article: http://dx.doi.org/10.1080/01650520412331333747

PLEASE SCROLL DOWN FOR ARTICLE

Full terms and conditions of use: http://www.tandfonline.com/page/terms-and-conditions

This article may be used for research, teaching, and private study purposes. Any substantial or systematicreproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form toanyone is expressly forbidden.

The publisher does not give any warranty express or implied or make any representation that the contentswill be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses shouldbe independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims,proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly inconnection with or arising out of the use of this material.

ORIGINAL ARTICLE

Influence of edge and fire-induced changes on spatial distribution ofsmall mammals in Brazilian Atlantic forest fragments

ALEXANDRA S. PIRES, FERNANDO A. S. FERNANDEZ, DANIELA DE FREITAS, &

BARBARA R. FELICIANO

Department of Ecology, Institute of Biology, Federal University of Rio de Janeiro, Brazil

(Received 5 January 2000; accepted 8 September 2003)

AbstractThe spatial distribution of small mammals in relation to distance from forest edges, before and after fire, was studied inAtlantic Forest fragments in Brazil. None of nine species was captured exclusively on either edges or forest interior. Beforethe fire only the rodents Akodon cursor and Oecomys con-color were captured more often at the edge than randomly expected.After the fire only A. cursor remained associated with edges; it had increased in number and penetrated farther than beforeinto the forest fragments. The marsupial Micoureus demerarae became more restricted to the forest interior than randomlyexpected. These results suggest that small mammal species tolerant to habitat changes induced by edge effects and fire havebetter chances to survive in forest fragments.

Keywords: Atlantic coastal forest, edge effects, forest fragmentation, fire, rodents, marsupials.

Introduction

The rate of tropical deforestation exceeds 15 million

ha annually, resulting in extensive fragmentation of

forest landscapes (Whitmore, 1997). In the Atlantic

Forest along the Brazilian east coast, fragmentation

has already reached a very advanced stage, as the

forest has been dramatically reduced in the last

centuries, due to the expansion of agriculture, cattle

raising, mining, and human settlements (Dean,

1996). Today only about 5% of the original forested

area remains (Fonseca, 1985), mostly in small

fragments with areas of tens to hundreds of ha. As

the Atlantic Coastal Forest is one of the World’s

most species rich systems (Quintela, 1990; Myers et

al., 2000), it is a critical priority to carry out detailed

field studies on the responses of its species to

fragmentation.

A common feature of habitat fragmentation is a

sharp increase in the amount of induced habitat

edge. Consequently, plant and animal populations in

forest fragments are not only reduced and subdi-

vided, but also suffer the effects of increased

exposure to abiotic and biotic changes associated

with forest edges (Lovejoy et al., 1986; Wilcove et al.,

1986; Murcia, 1995; Kapos et al., 1997; Laurance,

1997). In comparison to interior forest, structural

changes found near edges often include reduced

canopy cover, higher abundance of lianas, higher

understorey foliage density, increased windthrow of

trees and broken limbs, and increased litter fall

(Laurance, 1991a, 1997; Malcolm, 1994; Didham &

Lawton, 1999). In the Neotropics, studies have

shown that such changes affect the distribution,

abundance, richness and diversity of many plants,

insects, birds and mammals (Lovejoy et al., 1986;

Laurance, 1991a, 1994; Brown & Hutchings, 1997;

Stevens & Husband, 1998). Small forest remnants

can have their entire areas affected as the effects of

edge expand to the center of the fragments. Then,

the resulting vegetation consists of little more than

edge-modified habitat, which does not support

organisms dependent upon conditions found only

in the interior forest.

Several characteristics of edges can boost the

penetration, into forest fragments, of fires moving

across surrounding flammable habitats (like grass-

lands or pastures). These include abiotic factors of

Correspondence: F.A.S. Fernandez, Laboratorio de Ecologia e Conservacao de Populacoes, Departamento de Ecologia, Instituto de Biologia, Universidade

Federal do Rio de Janeiro, C.P. 68020, 21941-590 Rio de Janeiro – RJ, Brazil. E-mail: rodentia@biologia.ufrj.br

Studies on Neotropical Fauna and Environment, April 2005; 40(1): 7 – 14

ISSN 0165-0521 print/ISSN 1744-5140 online ª 2005 Taylor & Francis Group Ltd

DOI: 10.1080/01650520412331333747

Dow

nloa

ded

by [

Dal

hous

ie U

nive

rsity

] at

05:

39 1

8 A

pril

2013

edges like increased air temperatures, wind speeds

and solar radiation, and reduced air humidity, as well

as some features of the edge vegetation (e.g.

abundance of litter, fallen logs and flammable

understorey vegetation such as lianas and vines).

Favored by these factors, fire may penetrate easily or

at least damage the margins of unburnable forest at

the line of contact. Along the fire-damaged edge a

substantial amount of herbaceous vegetation will

then grow in the following season, thereby extending

the margin of the inflammable area (Janzen, 1986).

Recent studies in the Amazonian region have shown

that forest fires create positive feedbacks in future fire

susceptibility, fuel loading, and fire intensity (e.g.

Cochrane et al., 1999). Such fire effects are

particularly severe in fragmented areas, and therefore

can contribute to further fragmentation of tropical

forests (Nepstad et al., 1999).

Fire events change the structure, biomass and

species composition of the vegetation (Cochrane &

Schulze, 1999) and, in the long-term, favor coloniza-

tion by animal species characteristic of disturbed

habitats (Lovejoy et al., 1986; Laurance, 1991a;

Higgs & Fox, 1993). The crowding of animals within

the fragments, a pattern often observed in fragmen-

ted systems (Lovejoy et al., 1983, 1986), can also be

boosted by fire due to the influx of individuals from

the surrounding burned habitat. In both cases

species living in the fragments are often negatively

affected, at least in the short term, by competition

with invaders and refugees.

Non-flying mammals are regarded as the terrestrial

vertebrates most vulnerable to the effects of forest

fragmentation (Wilcox, 1980), but detailed informa-

tion on small mammals’ responses to habitat

fragmentation is still scarce in the Neotropics.

Regarding edge effects in particular, there are just a

few studies providing information on how they affect

Neotropical marsupials and rodents (e.g. Laurance,

1994; Heske, 1995; Stevens & Husband, 1998). The

present contribution, which is part of an ongoing long-

term study on the effects of fragmentation, had two

objectives: (1) to evaluate how the spatial distributions

of populations of small mammals were affected by the

distance from the forest edge in two Atlantic Forest

fragments in south-eastern Brazil; and (2) to deter-

mine if (and how) fire striking the fragments’ edges

affected the distribution of individuals.

Material and methods

Study area

The study was carried out in two Atlantic Forest

fragments which are part of a group of eight

fragments known as ‘Ilhas dos Barbados’, within

Poco das Antas Biological Reserve, Rio de Janeiro,

southeastern Brazil (22830’ – 22833’S, 42815’ –42819’W). The two fragments, named ‘A’ and ‘D’,

have areas of 7.1 and 8.8 ha, respectively. The

distance between them is 300 m. The transition from

the fragments to the vegetation of the matrix (open

area around the fragments) is abrupt, as the

fragments are situated on small mounds in the peaty

matrix soil, which is periodically flooded.

The climate of the region is warm tropical with

average annual temperatures above 248C. Average

annual precipitation reaches about 1700 mm and its

distribution is moderately seasonal, as nearly 30% of

the annual precipitation falls during the dry season

(M.C. Kierulff, pers. comm.).

The vegetation of the fragments is typical Atlantic

Coastal Forest, with trees about 20 m tall, and rich

in palms (mostly Astrocaryum aculeatissimum and

Attalea humilis). It was disturbed to a moderate

degree by selective logging in the past, but otherwise

has been protected from disturbance, except for fires,

since 1975. The most recent fire, the effects of which

were evaluated in the present study, occurred on 18

August 1997. It completely burned the matrix, which

is composed mostly of introduced grasses, and

severely damaged the edges of the forest remnants.

Captures of small mammals

Trapping lines were marked transversely across the

greatest length of each fragment, covering its whole

area. Lines were 50 m apart and trapping points were

20 m apart. This design resulted in 64 trapping

stations in fragment A and 78 in fragment D. At each

point a live-trap (either Tomahawk 17 6 17 648.5 cmor Sherman386 126 10 cm)was set on the

ground. At every second point an additional Sherman

trap was set on tree branches or vines at breast height

(about 1.5 m). From January 1997 to August 1997

tree traps were set in all capture points. As the two

types of ground traps were set on alternate points and

trap lines had different lengths (according to the

varying widths of each fragment), proportions of

Tomahawks and Shermans differed slightly among

distance classes. However, all species were readily

captured in both trap types, except forDidelphis aurita,

which was seldom captured in Shermans. Therefore,

we removed this species from any quantitative

analyses. Before the fire trapping sessions were carried

out every secondmonth fromMarch1995 to July 1997

in fragment A, and fromApril 1996 to August 1997 in

fragment D. The same trap design was also used after

the fire, from September 1997 to July 1998 in A, and

from October 1997 to August 1998 in D. Before the

fire the sampling effort was 6150 and 4088 trap-nights

in fragments A andD, respectively. After the fire there

were 3156 trap-nights in A and 4206 in D. Each

trapping session consisted of five consecutive nights of

8 A.S. Pires et al.

Dow

nloa

ded

by [

Dal

hous

ie U

nive

rsity

] at

05:

39 1

8 A

pril

2013

capture. All traps were baited with a mixture of oat,

banana, bacon and peanut butter placed on a manioc

slice. Traps were checked every morning and the

captured animals were marked individually with

numbered ear-tags.

To evaluate whether some species found in the

fragments were actually common in the surrounding

matrix and only used the fragments opportunisti-

cally, either before or after the fire, we placed eight

trapping lines between the two fragments studied.

These lines were run from May 1997 to August

1998. Each line had nine Sherman live-traps spaced

at 20 m intervals and placed on the ground; the

distance between trapping lines was also 20 m.

Additional Sherman traps, in a total of 167 trap-

nights, were placed at 1.5 m height in points that had

pioneer trees. The trapping effort in the matrix

amounted to 300 trap-nights before the fire and 2017

trap-nights after the fire.

Data analysis

Capture points were grouped into four distance

classes from the edge: 0 – 20 m, 20 – 40 m, 40 – 60 m

and 60 – 100 m (no capture point was farther than

100 m from the closest edge). The last class resulted

from pooling the 60 – 80 and 80 – 100 m classes,

because each of these had a small number of points in

each fragment and therefore the observed frequencies

of captures would be prone to sampling errors.

As the number of points in each distance class

varied, we used a Kolmogorov – Smirnov test for

grouped data (Zar, 1984) to evaluate if the distribu-

tions of captures of each species fit what would be

expected under the null hypothesis that captures

were randomly distributed among distance classes

according to trapping effort. This approach was

prefered to using ANOVA (with number of captures

of each species as variables, distance classes as

treatments and fragments as replicas) because

homocedasticity, an important assumption of ANO-

VA, was strongly violated for most species.

Kolmogorov – Smirnov analysis was carried out in

two ways: first, for each fragment separately; second,

pooling the data from both fragments. As the results

were identical for all species in both fragments (with

one exception mentioned below), for simplicity only

the results from the second analysis (fragments

pooled) are presented.

For species whose distributions differed from

random expectation in both periods, c2 tests were

also used to evaluate whether distribution found after

fire differed from that before fire. Additionally, for

each species with large enough sample size, overall

abundance (number of individuals captured) in the

fragments before and after fire was compared using

c2 tests with Yates’ correction for continuity (Zar,

1984).

Results

A total of 11 species were captured in the fragments

(Table I). The species captured most often, before

and after fire respectively, were the marsupial

Micoureus demerarae and the rodent Akodon cursor.

Three species were removed from any quantitative

analysis: Gracilinanus microtarsus and Rattus rattus,

that were captured only once, and Didelphis aurita,

because its trap bias posed problems for interpreting

its patterns.

No species was captured exclusively at edges or in

the interior, if both periods are taken into account

(Figure 1). Before the fire, only the distributions of

captures of Oecomys concolor differed between frag-

Table I. Number of captures and individuals () of each small mammal species, before and after fire, in two Atlantic Forest fragments in

southeastern Brazil.

Fragment A Fragment D

Before fire After fire Before fire After fire

MARSUPIALS

Caluromys philander 2(2) 11(5) 5(3) 15(4)

Didelphis aurita 1(1) 5(3) 6(3) 3(3)

Metachirus nudicaudatus 0 0 23(7) 17(7)

Philander frenata 1(1) 7(4) 2(1) 10(6)

Gracilinanus microtarsus 0 0 1(1) 0

Micoureus demerarae 184(39) 179(36) 114(22) 70(13)

RODENTS

Nectomys squamipes 0 6(3) 3(1) 1(1)

Oecomys concolor 94(42) 12(5) 23(16) 18(12)

Oligoryzomys nigripes 10(8) 18(18) 3(3) 7(6)

Akodon cursor 45(21) 233(103) 31(20) 122(65)

Rattus rattus 0 1(1) 0 0

Edge and fire effects on small mammals 9

Dow

nloa

ded

by [

Dal

hous

ie U

nive

rsity

] at

05:

39 1

8 A

pril

2013

ments, as the distribution deviated from the random

expectation in fragment A but not in fragment D

(Kolmogorov – Smirnov test, d=16.47, p5 0.05,

and 2.81, p4 0.50 respectively); this result was

possibly due to the smaller sample sizes in the latter

fragment. Among the remaining species, pooling the

fragments, only the distribution of captures of A.

cursor deviated from random expectation (Kolmo-

gorov – Smirnov test, d=32.2, p5 0.001). Oecomys

concolor in fragment A and A. cursor in the two

fragments were both captured more often near the

edge. This pattern was stronger for A. cursor which

had 68.4% of its captures in the first distance class.

After the fire only capture data for A. cursor andM.

demerarae deviated from random expectation (frag-

ments pooled, Kolmogorov – Smirnov for A. cursor

Figure 1. Trapping success of small mammals in relation to distance from edge in two fragments of Atlantic Forest in Rio de Janeiro state,

southeastern Brazil, before fire (March 1995 to July 1997) and after fire (September 1997 to August 1998). Bars representing the two periods

studied are as follows: ( before fire; _ after fire. The number of captures of each species in each period, respectively, are in parentheses.

10 A.S. Pires et al.

Dow

nloa

ded

by [

Dal

hous

ie U

nive

rsity

] at

05:

39 1

8 A

pril

2013

d=132.5; for M. demerarae d=51.90; in both cases

p5 0.001). Although A. cursor was always more

abundant near the edge, the proportion of captures

in the interior increased after fire in both fragments

(w2 = 8.065, df = 3, p=0.047). Micoureus demerarae

was captured less often near the edge after the fire,

causing the observed departure from random ex-

pectation. For O. concolor, the preference for edges

which was observed in fragment A before the fire was

not evident after the fire.

Before the fire the mammal community in the

adjoining matrix was dominated by the rodent A.

cursor, which comprised 80.0% of all captures (Table

II). The only marsupial caught in the matrix before

the fire was M. demerarae, which was caught twice.

After the fire the most abundant species were the

rodents A. cursor and Bolomys lasiurus, which together

made up 88.4% of captures. After the fire, a further

four species were captured in the matrix, including

the marsupials D. aurita and Philander frenata (Table

II).

Among the species common in the matrix, only

the rodent A. cursor was found in the fragments

before and after the fire. The other species captured

in both habitats were the rodent Oligoryzomys

nigripes, which had few captures in both places, and

the marsupials Didelphis aurita, Philander frenata and

Micoureus demerarae, which were captured most often

in the fragments.

Only O. concolor and A. cursor exhibited significant

overall differences in abundance in the fragments

before and after the fire. After the fire there was a

decrease in O. concolor (c2 with Yates’ correc-

tion=27.075, df = 1, p5 0.0001), and an increase

in A. cursor (c2 with Yates’ correction =296.67,

df = 1, p5 0.0001). The increase in A. cursor did

not occur immediately after the fire but only after a

delay of some months (Figure 2).

Discussion

Species’ responses to edge and fire effects

Working at two Atlantic Forest fragments of 145 and

547 ha, Stevens and Husband (1998) found that

some small mammal species were not captured at the

edges of the fragments. In our study, however, no

species occurred exclusively either at edges or in the

interior of fragments. As the fragments studied are

much smaller, the effect of the forest edges is

relatively large, leaving only a rather small inner

area. This is perhaps too small to maintain viable

populations of species requiring habitats with char-

acteristics of continuous forest. Two non-exclusive

hypotheses could explain the patterns we found.

First, species that need conditions present only

inside the forest could have already disappeared

form the fragments, or second, those species are still

present in the fragments but may be using sub-

optimal areas. The specific responses of each small

mammal to the habitat changes produced by edge

and fire effects are discussed below.

Akodon cursor

This rodent was one of the dominant species in the

matrix. Paglia et al. (1995) observed that it was more

common in areas of anthropic fields and scrub than

in an Atlantic Forest fragment, whereas Gentile and

Fernandez (1999) found that its abundance was

related to herbaceous density near the ground and

increased litter density. The latter are often found on

forest edges (Lovejoy et al., 1986; Laurance, 1991a;

Malcolm, 1994), a pattern that could explain the

high abundance of A. cursor along the edges in our

Table II. Number of captures and individuals () of each small

mammal species, before and after fire, in the matrix between two

Atlantic Forest fragments in southeastern Brazil.

Before fire (300 trap-

nights)

After fire (2017 trap-

nights)

MARSUPIALS

Didelphis aurita 0 1(1)

Philander frenata 0 5(2)

Micoureus demerarae 2(1) 0

RODENTS

Oligoryzomys nigripes 20(13) 17(11)

Akodon cursor 125(33) 242(92)

Bolomys lasiurus 9(3) 292(65)

Mus musculus 0 46(36)

Rattus rattus 0 1(1)

Figure 2. Number of individuals of Akodon cursor captured at

Atlantic Forest fragments in Rio de Janeiro state, southeastern

Brazil, before and after fire. (A) Fragment A, (B) Fragment D.

Edge and fire effects on small mammals 11

Dow

nloa

ded

by [

Dal

hous

ie U

nive

rsity

] at

05:

39 1

8 A

pril

2013

study. Akodon cursor feeds mostly on invertebrates

(Fonseca & Kierulff, 1989; Stallings, 1989; Carvalho

et al., 1999) thus edges could be an especially

suitable habitat for it, as invertebrate abundance

usually increases towards the borders of the forest

(Didham, 1997). The increased abundance of A.

cursor in the fragments after the fire cannot be

explained by an influx of individuals from the burned

matrix (a crowding effect sensu Lovejoy et al., 1983,

1986), as numbers did not increase immediately after

the fire. Instead the increase was probably due to

physiognomic and ecological changes brought by the

penetration of edge effects following the fire, making

the interior of the fragments more suitable for this

species. Nonetheless, as rodent densities can vary

greatly within or between years in the same habitat

even without fires, in the absence of control plots we

cannot be sure whether the observed variation in A.

cursor densities was due to fire effects.

Micoureus demerarae

This marsupial apparently used both the edge and

the interior of the forest fragments. A similar pattern

was found by Stevens & Husband (1998) for the

same species (as M. cinereus). Such results indicate

that the spatial distribution of M. demerarae is not

restricted to the interior of the forests.

This may be due to their preference for dense vines

(Emmons & Feer, 1997) commonly growing at the

forest borders. Moreover, movements of adult males

of this species were detected between the fragments

studied (Pires & Fernandez, 1999; Pires et al., 2002),

showing that M. demerarae occasionally crosses the

matrix. However, the increased proportion of cap-

tures in the interior of fragments suggests that edges

became unsuitable for this marsupial after the fire, as

trees and vines were mostly burned.

Oecomys concolor

These rodents showed some preference for edges

before the fire but this pattern was not found after the

fire when the species became scarcer. As O. concolor

is mostly arboreal, being most numerous in dense

vines (Emmons & Feer, 1997), its smaller number of

captures after the fire could be due to the damage of

lianas. However, as rodent densities may vary

considerably in the absence of fires, there could have

been other causes for the decrease.

Caluromys philander

The higher abundance in the interior of fragments

probably reflects this marsupial’s preference for the

middle and upper levels of the forest, especially the

canopy (Julien-Laferriere, 1995; Passamani, 1995;

Leite et al., 1996; Grelle, 2003). As edges have

sparser canopies (Malcolm, 1994; Laurance, 1997),

they should be unsuitable for C. philander. The

increased number of captures after the fire (in traps

on the ground or at breast height) may reflect an

increase in trappability, as a shortage of resources

following this event could have forced the animals to

move down to the ground searching for food.

Metachirus nudicaudatus, Philander frenata and

Oligoryzomys nigripes

Our study did not reveal clear habitat preferences for

these species. This is in contrast to the results of

Stevens and Husband (1998) who never capturedM.

nudicaudatus within 80 m of edges. In contrast,

individuals of M. nudicaudatus often move among

forest fragments (Pires et al., 2002), showing that

they are not confined to continuous forests. Philander

frenata and O. nigripes are both found in primary and

secondary forests as well as in a variety of open

vegetations (Cerqueira et al., 1990; Paglia et al.,

1995; Emmons & Feer, 1997) and this versatility

may explain their lack of preference for either edges

or interior forest.

Nectomys squamipes

This semi-aquatic rat did not seem to enter farther

than 60 m into the fragments. As this species is often

found in flooded grasslands (Emmons & Feer,

1997), parts of the matrix may provide a suitable

habitat, from where it invades the forest borders.

Concluding remarks

After each fire, edges advance further into the

forest fragments (Janzen, 1986), setting the balance

at a new point a little farther from the structure of

a continuous forest. As each mammal species has

its own habitat needs, this slow process of habitat

degradation is reflected by the species that are able

to survive and to maintain viable populations in

the fragments. Thus it seems that M. demerarae

that is well adapted to secondary forests, and A.

cursor adapted to forest edge conditions are less

susceptible to the changing conditions inside the

forest or may even profit from the increased edge

effects.

Our findings for A. cursor, as well as the presence

in fragments of several marsupials which cross the

matrix (M. demerarae, D. aurita, P. frenata and M.

nudicaudatus; see Pires et al., 2002), both corroborate

Laurance’s (1991b, 1994) findings that those forest

mammals that are most tolerant to the matrix

habitats will survive better in small fragments. It

remains an open question whether populations of

12 A.S. Pires et al.

Dow

nloa

ded

by [

Dal

hous

ie U

nive

rsity

] at

05:

39 1

8 A

pril

2013

animals which live mostly in the canopy, such as C.

philander and Gracilinanus microtarsus, will be able to

persist in small fragments in face of ongoing

degradation, which tends to produce a lower and

more open canopy. Long-term field studies are

clearly crucial to trace the process of environmental

changes in Neotropical forest fragments, and to

understand their effects on small mammal commu-

nities.

Acknowledgments

We thank IBAMA – Brazilian Institute of Environ-

ment and Renewable Natural Resources – for

allowing us to work at Poco das Antas and providing

many facilities there. We also thank the many

colleagues who helped in field work. J.L. Camargo

provided the fragments’ areas. William F. Laurance

provided several excelent criticisms and suggestions

which were very useful for improving the ms. Anne

Zillikens and an anonymous referee made very useful

comments. The work was funded by Fundacao O

Boticario de Protecao a Natureza, The MacArthur

Foundation, FAPERJ, FUJB and PROBIO (PRO-

NABIO, with the support of BIRD/GEF, MMA and

CNPq). Personal grants were given by CEPG-UFRJ,

FAPERJ, CAPES and CNPq.

References

Brown Jr. KS, Hutchings RW. 1997. Disturbance, fragmentation,

and the dynamics of diversity in Amazonian forest butterflies.

In: Laurance WF, Bierregaard RO, editors, Tropical Forest

Remnants – Ecology, Management, and Conservation of

Fragmented Communities. Chicago and London, University

of Chicago Press, p 91 – 110.

Carvalho FMV, Pinheiro PS, Fernandez FAS, Nessimian JL.

1999. Diet of small mammals in Atlantic Forest fragments in

southeastern Brazil. Rev Bras Zooc 1:91 – 101.

Cerqueira R, Fernandez FAS, Quintela MFS. 1990. Mamıferos da

restinga de Barra de Marica, Rio de Janeiro. Pap Avul Zool

37:141 – 157.

Cochrane MA, Alencar A, Schulze MD, Souza Jr CM, Nepstad

DC, Lefebvre P, Davidson EA. 1999. Positive feedbacks in the

fire dynamic of closed canopy tropical forests. Science

284:1832 – 1835.

Cochrane MA, Schulze MD. 1999. Fire as a recurrent event in

tropical forests of the eastern Amazon: effects on forest

structure, biomass, and species composition. Biotropica

31:2 – 16.

Dean W. 1996: A Ferro e Fogo. A Historia e a Devastacao da

Mata Atlantica Brasileira. Sao Paulo, Companhia das Letras.

Didham RK. 1997. The influence of edge effects and forest

fragmentation on leaf litter invertebrates in Central Amazonia.

In: Laurance WF, Bierregaard RO, editors, Tropical Forest

Remnants – Ecology, Management, and Conservation of

Fragmented Communities. Chicago and London, University

of Chicago Press, p 55 – 70.

Didham RK, Lawton JH. 1999. Edge structure determines the

magnitude of changes in microclimate and vegetation structure

in tropical forest fragments. Biotropica 31:17 – 30.

Emmons LH, Feer F. 1997. Neotropical Rainforest Mammals: A

Field Guide. Chicago & London, The University of Chicago

Press.

Fonseca GAB. 1985. The vanishing Brazilian Atlantic Forest. Biol

Conserv 34:17 – 34.

Fonseca GAB, Kierulff MCM. 1989. Biology and natural history

of Brazilian Atlantic Forest small mammals. Bull Fla State Mus

Biol Sci 34:99 – 152.

Gentile R, Fernandez FAS. 1999. Influence of habitat structure on

a streamside small mammal community in a Brazilian rural

area. Mammalia 63:29 – 40.

Grelle CEV. 2003. Forest structure and vertical statification of

small mammal populations in a secondary forest, southeastern

Brazil. Stud Neotrop Fauna Environm 38:81 – 85.

Heske EJ. 1995. Mammalian abundances on forest-farm edges

versus forest interiors in southern Illinois: is there an edge

effect? J Mammal 76:562 – 568.

Higgs P, Fox BJ. 1993. Interspecific competition: a mechanism for

rodent succession after fire in wet heathland. Aust J Ecol

18:193 – 201.

Janzen DH. 1986. The eternal external threat. In: Soule ME,

editor. Conservation Biology: The Science of Scarcity and

Diversity. Sunderland, Sinauer Associates, p 286 – 303.

Julien-Laferriere D. 1995. Use of space by the woolly opossum

Caluromys philander (Marsupialia, Didelphidae) in French

Guiana. Can J Zool 73:1280 – 1289.

Kapos V, Wandelli E, Camargo JL, Ganade G. 1997. Edge-related

changes in environment and plant responses due to forest

fragmentation in Central Amazonia. In: Laurance WF,

Bierregaard RO, editors. Tropical Forest Remnants – Ecology,

Management, and Conservation of Fragmented Communities.

Chicago and London, University of Chicago Press, pp 71 – 84.

Laurance WF. 1991a. Edge effects in tropical forest fragments:

application of a model for the design of nature reserves. Biol

Conserv 57:205 – 219.

Laurance WF. 1991b. Ecological correlates of extinction prone-

ness in Australian Tropical Rain Forest mammals. Conserv Biol

5:79 – 89.

Laurance WF. 1994. Rainforest fragmentation and the structure of

small mammal communities in tropical Queensland. Biol

Conserv 69:23 – 32.

Laurance WF. 1997. Hyper-disturbed parks: edge effects and the

ecology of isolated rainforest reserves in Tropical Australia. In:

Laurance WF, Bierregaard RO, editors. Tropical Forest

Remnants – Ecology, Management, and Conservation of

Fragmented Communities. Chicago and London, University

of Chicago Press, p 71 – 84.

Leite YLR, Costa LP, Stallings JR. 1996. Diet and vertical space

use of three sympatric opossums in a Brazilian Atlantic Forest

reserve. J Trop Ecol 12:435 – 440.

Lovejoy TE, Bierregaard Jr. RO, Rankin JM, Schubart HOR.

1983. Ecological dynamics of tropical forest fragments. In:

Whitmore TC, Chadwick AC, Sutton SL, editors. Tropical

Rain Forest: Ecology and Management. Oxford, Blackwell,

p 377 – 384.

Lovejoy TE, Bierregaard Jr. RO, Rylands AB, Quintela CE,

Harper LH, Brown Jr. KS, Powell AH, Powell GVN, Schubart

HOR, Hays MB. 1986. Edge and other effects of isolation on

Amazon Forest fragments. In: Soule ME, editor. Conservation

Biology: The Science of Scarcity and Diversity. Sunderland,

Sinauer Associates, p 257 – 285.

Malcolm JR. 1994. Edge effects in central Amazonian forest

fragments. Ecology 75:2438 – 2445.

Murcia C. 1995. Edge effects in fragmented forests: implications

for conservation. Trends Ecol Evol 10:58 – 62.

Edge and fire effects on small mammals 13

Dow

nloa

ded

by [

Dal

hous

ie U

nive

rsity

] at

05:

39 1

8 A

pril

2013

Myers N, Mittermeier RA, Fonseca GAB, Kent J. 2000.

Biodiversity hotspots for conservation priorities. Nature

403:853 – 858.

Nepstad DC, Verıssimo A, Alencar A, Nobre C, Lima E, Lefebvre

P, Schlesinger P, Potter C, Moutinho P, Mendonza E,

Cochrane M, Brooks V. 1999. Large-scale impoverishment of

Amazonian forests by logging and fire. Nature 398:505 – 508.

Paglia AP, De Marco Jr. P, Costa FM, Pereira RF, Lessa G. 1995.

Heterogeneidade estrutural e diversidade de pequenos mamı-

feros em um fragmento de mata secundaria de Minas Gerais,

Brasil. Rev Bras Zool 12:67 – 79.

Passamani M. 1995. Vertical stratification of small mammals in

Atlantic Hill forest. Mammalia 59:276 – 279.

Pires AS, Fernandez FAS. 1999. Use of space by the marsupial

Micoureus demerarae in small Atlantic Forest fragments in

southeastern Brazil. J Trop Ecol 15:279 – 290.

Pires AS, Lira PK, Fernandez FAS, Schittini GM, Oliveira LC.

2002. Frequency of movements of small mammals among

Atlantic Coastal Forest fragments in Brazil. Biol Conserv

108:229 – 237.

Quintela CE. 1990. An S.O.S. for Brazil’s beleaguered Atlantic

Forest. Nat Conserv Mag 40:14 – 19.

Stallings JR. 1989. Small mammals inventories in an eastern

Brazilian park. Bull Fla State Mus Biol Sci 34:153 – 200.

Stevens SM, Husband TP. 1998. The influence of edge on small

mammals: evidence from Brazilian Atlantic forest fragments.

Biol Conserv 85:1 – 8.

Whitmore TC. 1997. Tropical forest disturbance, disappearance,

and species loss. In: Laurance WF, Bierregaard RO, editors.

Tropical forest remnants – ecology, management, and con-

servation of fragmented communities. Chicago, University of

Chicago Press, p 3 – 12.

Wilcove DS, McLellan CH, Dobson AP. 1986. Habitat fragmen-

tation in the temperate zone. In: Soule ME, editor.

Conservation Biology: The Science of Scarcity and Diversity.

Sunderland, Sinauer Associates, p 237 – 256.

Wilcox B. 1980. Insular ecology and conservation. In: Soule ME,

Wilcox BA, editors. Conservation Biology. Sunderland,

Sinauer Associates, p 95 – 117.

Zar JH. 1984. Biostatistical Analysis. Englewood Cliffs, New

Jersey, Prentice-Hall.

14 A.S. Pires et al.

Dow

nloa

ded

by [

Dal

hous

ie U

nive

rsity

] at

05:

39 1

8 A

pril

2013