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Original Article 

Biosci. J., Uberlândia, v. 29, n. 3, p. 751-759, May/June 2013 

THE MANED WOLF IN THE ECOTONE BETWEEN FOREST AND 
GRASSLANDS AT THE LIMITS OF ITS DISTRIBUTION IN A 

SUBTROPICAL ENVIRONMENT 
 

O LOBO-GUARÁ NO ECÓTONO ENTRE FLORESTA E CAMPO NOS LIMITES DE 
SUA DISTRIBUIÇÃO EM UM AMBIENTE SUBTROPICAL 

 
 

Rosane Vera MARQUES1,2; Marta Elena FABIÁN2 
1. Unidade de Assessoramento Ambiental, Divisão de Assessoramento Técnico, Ministério Público do Estado do Rio Grande do Sul, 

Porto Alegre, RS, Brasil; 2. Pós-graduação em Biologia Animal, UFRGS, Porto Alegre, RS, Brasil. mfabian@ufrgs.br  
 

 
ABSTRACT: The maned wolf (Chrysocyon brachyurus) is a neotropical canid that is considered near 

threatened by the IUCN, vulnerable in Brazil and critically endangered in the state of Rio Grande do Sul.  The Araucaria 
High Plains have coverage of Atlantic forest mixed in with open areas and small wetland areas and are at one of the limits 
of the range of the maned wolf. The objectives of this article are to demonstrate that, although rare, the maned wolf does 
still occur in the Araucaria High Plains and to discuss the relevance of conservation efforts in areas in which the 
occurrence of threatened species is borderline. Data collection took place between March of 1999 and December of 2010 
with a total sampling effort of 11,431 trap-days. After 10 years in which there were no corroborated records, the species 
was caught by camera traps in the São Francisco de Paula National Forest in November of 2009.  This is a protected area 
with forest vegetation and is not considered the species’ typical habitat. The open areas in this region are being taken over 
by agriculture and silviculture of exotic species. 
 

KEYWORDS: Chrysocyon brachyurus. Canidae. Threatened species. Atlantic Forest. Brazil. Camera-trapping. 
 

 
INTRODUCTION 

 
Chrysocyon brachyurus (Illiger, 1815) (the 

maned wolf) is not typically a forest-dwelling 
species (CABRERA; YEPES, 1960; DIETZ, 1985) 
and is generally considered an inhabitant of open 
areas, but it does also occur in forested areas in the 
South of Brazil (CÁCERES et al,. 2007). It has been 
classified as near threatened by the IUCN because 
of drastic reductions in its habitat as a result of 
conversion to agriculture, causing fragmentation of 
habitats, isolation of subpopulations and road 
deaths. The presence of feral dogs inside 
Conservation Units (CU), exposes the maned wolf 
to the threats of hunting, increased competition and 
disease transmission (IUCN, 2008).  Within Brazil 
the species is defined as threatened with extinction 
(CHIARELLO et al,. 2008). Specifically in the state 
of Rio Grande do Sul (south Brazil), it is classified 
as critically endangered (INDRUSIAK; EIZIRIK, 
2003). 

The Brazilian Atlantic forest can be 
considered a “hotspot” on the basis of the great 
degree of biological diversity concentrated in 
relatively small areas. The most significant threats 
to conservation are deforestation, agriculture, 
silviculture of exotic species and hunting. This 

region has a long history of exploitation which 
makes it highly relevant to understanding the long 
term effects of habitat fragmentation on other 
tropical regions that are under threat (LAURANCE, 
2009). The Atlantic forest is a series of rainforest 
ecosystems that extends from the northeast to the 
south of Brazil (FONSECA et al., 2004). The 
Atlantic forest can be divided into at least eight sub-
regions, one of which is the Araucaria Forest, in the 
south of the country (RIBEIRO et al,. 2009). A large 
proportion of the work done on conservation in the 
Atlantic forest has been conducted in the southeast 
(VIEIRA et al., 2009; GALETTI et al., 2009) or 
northeast (LEAL et al., 2010) regions of Brazil, but 
the relevance to biodiversity of key areas in the 
south of the country has also recognised (PAESE et 
al., 2010). 

The Araucaria High Plains have some 
vegetation containing species associated with the 
Atlantic forest mixed in with fields, forming a 
mosaic pattern (PILLAR et al., 2009). They are 
located in the northeast of the state of Rio Grande 
do Sul, (RS), and are considered to represent the 
limit of the current range of the maned wolf (IUCN, 
2008; CHIARELLO et al., 2008). The species has 
been considered rare in RS since the beginning of 
the twentieth century. In the 1970s it was recorded 

Received: 22/05/12 
Accepted: 05/10/12 



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O Lobo-Guará...  MARQUES, R. V.; FÁBIAN, M. E. 

Biosci. J., Uberlândia, v. 29, n. 3, p. 751-759, May/June 2013 

in larger areas in the centre, west and southwest of 
the state and in smaller areas in the northeast, north, 
south and southeast, the coastal plains and in the 
environs of the state capital Porto Alegre 
(INDRUSIAK; EIZIRIK, 2003). Specifically in the 
Araucaria High Plains, scats and sightings were 
reported at the end of the 1990s and start of the 
2000s, in the Aparados da Serra National Park 
(SANTOS et al., 2004). 

Recently, studies conducted using camera 
traps in the northeast of RS and the southeast of the 
state of Santa Catarina (SC) did not detect the 
maned wolf which makes the reports from the last 
decade inconclusive (FREITAS et al., 2009). 

The top priority for conservation projects 
focussed on the maned wolf was to conduct surveys 
of populations within and around protected natural 
areas throughout its geographical distribution 
(SILLERO-ZUBIRI; MacDONALD, 2004). The 
methods applied to determine where the species 
occurs include satellite habitat mapping, interviews 
with local people who spend a considerable time in 
the field (and who are capable of recognizing the 
species unequivocally) and camera trap images 
(SILLERO-ZUBIRI et al., 2004). 
 The objectives of this article are to 
demonstrate that, although rare, the maned wolf 
does still occur in the Araucaria High Plains and to 
discuss the relevance of conservation efforts made 
in areas in which the occurrence of threatened 
species is borderline. 
 
MATERIAL AND METHODS 
 
Study area 

The São Francisco de Paula National Forest 
– FLONASFP - is defined as a “Sustainable Use 
Conservation Unit” (Unidade de Conservação de 
Uso Sustentável) and is located within the municipal 
limits of São Francisco de Paula. It has a total area 
of 1,606.7 ha, at an altitude of 930 m and has 
canyons with depths of up to 100 m. The region’s 
climate is classified as Cfb according to the 
Köppen-Geiger system and is a humid subtropical 
zone (rains are well distributed and summers are 
mild) (CÁCERES et al., 2007). The native 
vegetation is either Mixed Rain Forest or Araucaria 
Forest. In addition to native forest, this CU also has 
managed stocks of native pine Araucaria 
angustifolia (Bert.) and exotic species (Pinus 
elliotii, Pinus taeda, Eucaliptus sp) 
(CADEMARTORI et al., 2002). The first studies of 
the mammals inhabiting the CU began in 1992 and 
by 2007 66 native mammal species and one exotic 

had been detected and 30 of the native species were 
medium to large animals (MARQUES et al., 2011).   

 
Fieldwork methods  

Records of medium and large mammals 
(body mass greater than 1 kg) were obtained on the 
basis of opportunistic direct observations (during the 
daytime and at twilight) and/or tracks and scats, 
which supplemented the primary method of using 
camera traps. Data collection with the traps took 
place between March of 1999 and December of 
2010 with a total sampling effort of 11,431 trap-
days. A total of 10 camera traps were set up in pairs 
in order to record both sides of animals and 
remained active twenty-four hours per day, during 
the daytime, at twilight and at night during all four 
seasons of the year.  

Sampling stations were unbaited and set at 
500 m intervals along trails between native forest 
and Araucaria angustifolia stands and off trail in the 
forest interior, covering an area of around 4.7 km2. 
The camera traps were equipped with active sensors 
and were triggered whenever an animal broke an 
infrared beam passing between an emitter and a 
receiver (MARQUES; RAMOS, 2001).  

All fieldwork was duly authorised by the 
Brazilian federal environmental authorities 
(Authorization numbers 14 and 15/2001/RS, 14 and 
15/2002/RS, 23 and 24/2003/RS, 25/2004/RS, 
34/2005-NUFAU, 21 and 22/2006-DITEC 
IBAMA/RS, SISBIO 11848-2/2008 and SISBIO 
26664-1/2011). Two security guards who worked at 
FLONASFP and reported having seen maned 
wolves were interviewed and asked to provide 
detailed descriptions of the animals they had 
observed to enable an assessment of whether or not 
they really were sightings of maned wolves. 
 
RESULTS 

 
On the 1st of November of 2009 (which is 

spring in south Brazil) two traps photographed a 
maned wolf. The first picture was taken at 19:24 at 
coordinates 29°25’39.1”S 50°23’48.4”W and the 
second at 19:33 at coordinates 29°25’23.6”S 
50°24’06.8”W. Sunset was at 18:45 and so the 
records were defined as taken at twilight (between 1 
hour before and 1 hour after sunset) 
(THEUERKAUF et al., 2003). The animal appeared 
to be a healthy adult, with no obvious signs of worn 
teeth. Sex could not be determined (Figure 1). It had 
crossed the 500 m between the two traps in 9 min, at 
a velocity of around 0.93 m/s (3.33 km/h), 
apparently travelling without haste (Table 1). 
Around 15 days before, a partial maned wolf print 



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O Lobo-Guará...  MARQUES, R. V.; FÁBIAN, M. E. 

Biosci. J., Uberlândia, v. 29, n. 3, p. 751-759, May/June 2013 

had been found on the same trail where the animal 
was photographed.  

 

 

 
Figure 1. Maned-wolf caught on camera in November of 2009 in the Floresta Nacional de São Francisco de 

Paula, Rio Grande do Sul, Brazil. 
 
Table 1. Records of recent occurrences of Chrysocyon brachyurus in the São Francisco de Paula National 

Forest, RS, Brazil 
Records Camera traps Guards’ reports 

Date 01/XI/2009 18/VIII/2010 12/X/2010 
Season Spring Winter Spring 
Time 19:24                          19:33 17:30 08:00 
Period  twilight twilight daytime 
Velocity 0.93 m/s (3.33 km/h)   
Environment Road between  A. angustifolia 

plantations 
Pinus plantation 
and fields 

Road inside Pinus 
plantation 

Coordinates 29°25’39,1”S 
50°23’48,4”W     

29°25’23,6”S 
50°24’06,8”W 

29°23’36,9”S 
50°22’49,8”W 

29°24’41,7”S 
50°22’55,9”W 

 
The camera traps were left in place until the 

start of January 2010, but no more photographs of 
the species were taken on that trail. Later that year, 
two security guards who work at FLONASFP 
reported sightings of the species within the CU. On 
the 18th of August 2010, (winter), a single adult-
sized individual was seen at 17:30 (twilight) 
entering the FLONASFP perimeter close to the 
gates at coordinates 29°23’36.9”S 50°22’49.8”W 
after having followed a stretch of unpaved road 
outside the CU (Personal communication from Mr. 
João Luiz Machado Brando). On the 12th of 
October of 2010 (spring), an adult individual was 
seen at around 08:00 (daytime) with a cub about 
half of its size crossing an access road between 
Pinus elliotti plantations in the CU at approximately 
coordinates 29°24’41.7”S 50°22’55.9”W (Personal 
communication from Mr. Valdemar Scalcon) The 
distance between the site where the animal was 
photographed is around 4 km as the crow flies from 

the point where the individual was seen in the 
winter and the location of the third sighting is 
around 2 km from each of the first two. 
 
DISCUSSION 

 
The historic distribution of the maned wolf 

encompassed the pampas region and the high plains 
in RS. Natural grasslands in became degraded in 
areas in the North and Northeast of Argentina, 
Uruguay and the South of Brazil in which human 
populations reached high densities, leading to the 
species’ distribution limit reducing. There have been 
sporadic reports of maned wolves in the pampas and 
the high plains of RS between 1988 and 2008 
(QUEIROLO et al., 2011). The last records of the 
species in the Araucaria High Plains were in the 
1990s and the start of the 2000s (SANTOS et al., 
2004) and these animals had not been detected in 
the region since then (FREITAS et al., 2009). 



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Biosci. J., Uberlândia, v. 29, n. 3, p. 751-759, May/June 2013 

This long term data collection project in the 
field was designed to investigate forest-dwelling 
mammals and is not appropriate for species from 
open terrain. Despite this fact, the maned wolf was 
unequivocally recorded on camera, demonstrating 
that the species still exists in the Araucaria High 
Plains.    

The study area is not the maned wolf’s 
typical habitat, since it tends to inhabit open terrain 
with tall grass, followed by cerrado savannah 
vegetation with little forest cover (JÁCOMO et al., 
2004). However, the lack of tall dense grass in the 
area, resulting from repeated burning 
(INDRUSIAK; EIZIRIK, 2003), means that there is 
a scarcity of sites that are appropriate for dens in 
which to rear young, whether above ground or 
sheltered below cracks in rocks, with bushy 
coverage, tall grasses or small dry hummocks 
surrounded by swamp or marsh (SILLERO-
ZUBIRI, 2009). 
 The maned wolf is omnivorous and its diet 
consists of around 50% vegetable and 50% animal 
material, and in cerrado environments the fruit of 
the wolf  Solanum lycocarpum St. Hil. is one of the 
species’ principal food resources (SILLERO-
ZUBIRI, 2009). This vegetable species does not 
occur in the Araucaria High Plains region 
(JARENKOW; BUDKE, 2009). However, 
Araucaria Forest does contain species bearing edible 
fruit (PAISE; VIEIRA, 2005). Dietary items of 
animal origin include small mammals, the majority 
rodents - sigmodontinae, guinea pigs and marsupials 
- some mammals with body mass over 1kg 
(armadillos and leporids), birds of the Tinamidae 
Family and eggs, reptiles (lizards and non-
poisonous snakes), arthropods (ants, grasshoppers 
and crickets, beetles, ticks, spiders) (ARAGONA; 
SETZ, 2001; JÁCOMO et al., 2004). These groups 
of animals are present in the Araucaria High Plains, 
both in forested areas and in open spaces 
(CADEMARTORI et al., 2002; CADEMARTORI 
et al., 2004; CADEMARTORI et al., 2008; 
DALMAGRO; VIEIRA, 2005; PEDÓ et al., 2010; 
MARQUES et al., 2001; FONTANA et al., 2008; 
DEIQUES, 2009; OTT; LISE, 2009; DIEHL, 2009). 
However, food availability is limited in areas used 
for pasture and where fire has been used, since these 
areas have significantly reduced abundance, 
biomass and diversity of the small mammals than 
areas that have been protected from these impacts 
(PEDÓ et al., 2010).  
 In central Brazil, female maned wolves 
come into oestrus once a year, between April and 
June, and the majority of births occur between June 
and September (the dry season) after a gestation of 

around 65 days. Litters range from one to seven 
cubs, but the average is three. Cubs are weaned after 
15 weeks, but receive regurgitated food from their 
parents for 4 weeks. Juveniles remain with their 
mother for around a year before dispersing 
(SILLERO-ZUBIRI et al., 2004). There is a greater 
abundance of rodents in the Cerrado Biome during 
the dry season (MELO et al., 2007). There is no dry 
season in the Araucaria High Plains, since rain is 
well distributed throughout the year, while the 
greatest seasonal differences are the lower number 
of hours of light per day and lower temperatures 
during the winter (June to August) when compared 
with the summer (December to February) 
(CADEMARTORI et al., 2002). The greatest 
abundance of sigmodontinae rodents is during 
August (the end of winter) in Araucaria Forest 
(CADEMARTORI et al., 2004). In general, fields 
are burnt illegally to clear them at the end of winter. 
Vegetation will re-sprout resulting in cleared fields, 
reducing the quality of the habitat for maned-wolves 
(INDRUSIAK; EIZIRIK, 2003). This lack of 
hospitable areas in the species’ preferred 
environments may be the reason why some 
individuals seek refuge in forest within a protected 
area, especially when rearing young. 
 The majority of studies of the ecology and 
behaviour of the maned wolf have been conducted 
in the southeast and central regions of Brazil, with 
emphasis on the Cerrado. This canid shows a strong 
preference for rocky grasslands, avoiding the 
Atlantic forest, and follows roads at night to move 
about (COELHO et al., 2008). The species tends to 
use areas with dense vegetation during the day, 
whereas during the night, when it is more active, it 
prefers open areas (COELHO et al., 2007). Maned 
wolves have been found in areas with cerradão, 
cerrado and eucalyptus plantations in patchwork, 
demonstrating the behaviour of a generalist species 
(LYRA-JORGE et al., 2008). A 6-month study of 
three GPS-collared maned wolves in an area of 
cerrado in Minas Gerais state demonstrated that this 
species is predominantly active at night, peaking at 
twilight, and that the female’s living area reduces 
drastically after cubs are born (MELO et al., 2007). 
They form monogamous pairs and share home 
ranges, but the pair maintain a mean distance of 
more than 500 m from each other when they are 
active, in order to avoid interfering with successful 
feeding (JÁCOMO et al., 2009). In canids, 
variations in home range size and sociability can be 
attributed to availability and dispersion of food and 
habitat resources (SILLERO-ZUBIRI, 2009). In a 
region that is borderline of distribution for the 
species and in the presence of impacts on its 



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Biosci. J., Uberlândia, v. 29, n. 3, p. 751-759, May/June 2013 

primary habitat, it is to be expected that home 
ranges will be large and reports will be rare and 
sporadic. A subpopulation that has adapted to a 
different environment from that found in the centre 
of Brazil should be investigated with conservationist 
objectives, even if it is rare and population densities 
are low (SOULÉ; KOHM, 1989).      

The maned wolf is considered a generalist 
species, because it can survive in altered 
environments, as long as there is natural vegetation 
mixed in among the agroecosystems (LYRA-
JORGE et al., 2009). Notwithstanding, in areas used 
for livestock farming, domesticated dogs are left 
free and can be predators or disease vectors for 
maned wolves and poisoned bait is often laid in 
fields (SILLERO-ZUBIRI et al., 2004), intended for 
the crab-eating fox Cerdocyon thous (Linnaeus, 
1766) and the pampas fox Lycalopex gymnocercus 
(G. Fischer, 1814) which are accused of killing 
lambs. In the Araucaria High Plains region, more 
than 20 years ago, some farmers used to order their 
farm workers to distribute small pieces of sausage 
containing strychnine to kill wild canids (Personal 
communication from Mr. Arcelino Pereira Moraes). 
Nowadays this poison is banned, as is its sale, but 
other toxic substances could have devastating 
effects of canids were they to be used.  
 The presence of the maned wolf in 
FLONASFP shows that protected and unprotected 
areas play complementary roles in nature 
conservation, since their boundaries are not 
absolute, but permeable. Strategies to achieve a 
compromise between human interests and 
conservation in unprotected areas are essential if 

management plans are to be successful (PRIMACK, 
2006). Small areas under cultivation, and 
silviculture in patchwork with fields could, in 
principle, allow the species to adapt as  long  as  
open areas are conserved. The following measures 
constitute some of the steps needed to restore the 
necessary conditions for survival for maned wolves 
and other species threatened with extinction. 
Legislation that enforces the responsibility of rural 
landowners to conserve a proportion of the natural 
ecosystems on their properties. Environmental 
licensing of high-impact activities that takes into 
account the need to maintain the permeability of 
environments for rare species. Publicity and 
implementation of alternative methods for avoiding 
predation of domesticated animals by the maned 
wolf (PAULA et al., 2008). Environmental 
education aimed at a range of different publics.  
 
ACKNOWLEDGEMENTS 

 
We would like to express our heartfelt 

gratitude to the electrical engineer Fernando de 
Miranda Ramos for his help with the fieldwork and 
for developing the camera traps. We are also 
grateful to the security guards João Luiz Machado 
Brando and Valdemar Scalcon and to the caretaker 
Arcelino Pereira Moraes for information used in this 
research. We would also like to thank the forestry 
engineer Artur José Soligo, the agricultural engineer 
Edenice Brandão Ávila de Souza and all of the staff 
at FLONASFP for their help and logistical support. 

. 

 
 
RESUMO: Lobo-guará (Chrysocyon brachyurus) é um canídeo neotropical considerado quase ameaçado de 

extinção pela IUCN, vulnerável no Brasil e criticamente em perigo no estado do Rio Grande do Sul.  Planalto das 
Araucárias tem vegetação de Mata Atlântica entremeada por campos e pequenas áreas úmidas, sendo um dos limites de 
distribuição do lobo-guará. Os objetivos desse trabalho são demonstrar que, apesar de raro, o lobo-guará ainda existe no 
Planalto das Araucárias e discutir a relevância dos esforços de conservação em áreas limite de ocorrência de espécies 
ameaçadas de extinção. A coleta de dados ocorreu entre março de 1999 e dezembro de 2010 com um esforço amostral total 
de 11.431 armadilhas-dia. Após dez anos sem registros inequívocos, essa espécie foi capturada fotograficamente na 
Floresta Nacional de São Francisco de Paula em novembro/2009.  Essa área protegida apresenta formação vegetal 
florestal, não considerada hábitat típico dessa espécie. O ambiente aberto nessa região está sendo substituído por cultivos 
agrícolas e silvicultura de exóticas. 
 

PALAVRAS-CHAVE: Chrysocyon brachyurus. Canidae. Espécie ameaçada. Floresta Atlântica. Brasil. 
Armadilhas fotográficas. 
 
 
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