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Eur J Wildl Res (2005) 51: 199–206

DOI 10.1007/s10344-005-0092-1

O R I GI N A L P A P E R

Thierry Bonaudo Æ Yvonnick Le Pendu


Jean François Faure Æ Darcisio Quanz

The effects of deforestation on wildlife along the transamazon highway

Received: 23 November 2004 / Accepted: 22 March 2005 / Published online: 10 May 2005
 Springer-Verlag 2005

Abstract Hunting, although prohibited, is widely prac- settlement, development of a viable plan for game
ticed by the rural inhabitants settled along the Tran- management and forest preservation is of great impor-
samazon highway. In 1997 and 2000, we investigated tance.
subsistence hunting in Uruará, a township located on an
Amazonian pioneer front (Brazil). We analyze hunting Keywords Subsistence hunting Æ Amazon region Æ
practices, game yield, hunting efficiency and their rela- Game harvest Æ Pioneer settlement Æ Deforestation Æ
tion to forest coverage rate. The hunting methods were Hunting techniques
stand hunting (55.5%) and beating (44.5%). Paca
(Agouti paca), tatus (Dasypus novemcinctus and D. sep-
temcinctus), and collared peccary (Pecari tajacu) were
the most frequently hunted species, supplying 68% of Introduction
the gross game weight. Beating was significantly more
efficient than stand hunting (3 vs. 1.9 kg/hunter/h, In Brazil, vast colonization programs were started dur-
Mann–Whitney U test, P=0.02). Hunting territories ing the second half of the twentieth century with the
were classified in three categories according to forest settlement of thousand colons along axes of penetration
coverage rate. The higher the forest coverage rate the called pioneers fronts. Anthropogenic degradation is
larger was the harvested species and the more efficient more pronounced in border zones between wooded
the hunter (Kruskal–Wallis test P=0.01). Considering ecosystems and agricultural areas. Cattle ranching and
the ecological and demographic changes in this pioneer lumbering are the main causes of both deforestation and
the building of these new rural areas. The pasture would
cover 80% of all deforested surfaces (Santana et al.
T. Bonaudo (&)
1997). The National Institute for Space Research sug-
Département TERA-Green, CIRAD, gests that from 1978 to 2000, 65 millions hectares of
TA 60/15, 73 rue JF Breton, forest were lost in the Brazilian Amazonian region, i.e.
34398 Montpellier Cedex 5, France more than the surface of Germany and Italy together
E-mail: b_thierry@yahoo.com (INPE 2003).
Tel.: +33-467-615966
Fax: +33-467-614415 Redford (1993) estimates that rural Amazonian
populations in Brazil hunt 14 million mammals yearly.
Y. Le Pendu
Universidade Estadual de Santa Cruz (UESC),
Peres (2000) increased the number to 23.5 million
Departamento de Ciências Biológicas, mammals killed every year. Under this circumstance,
Rodovia Ilhéus/Itabuna km 16, game meat consumption would be up to 89.224 tons
CEP: 45662-000 Salobrinho, representing a value of US$ 190 million per year (Peres
Ilhéus, Ba, Brazil 2000). In spite of its ecological, economic and food
E-mail: yvonnick@uesc.br
supply importance, fauna exploitation is not submitted
J. F. Faure to management or valorization. Even though legally
Unité de Service ESPACE,
IRD Centre IRD d’Orléans Technoparc,
forbidden, hunting is widely practiced. The majority of
5 rue du Carbone, 45072 Orléans Cedex 2, the rural population practices subsistence hunting. In
France fact, game is the most accessible source of animal pro-
D. Quanz
tein. At the same time, hunting is a pastime maintaining
Embrapa, CEP: 68 140-000 Uruará, social relations with neighbours. Consequently, it is
Pará, Brazil necessary to put in place new ways of exploiting faunal
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resources while meeting local population needs and


preserving Amazonian biodiversity. This study analyzes Methods
hunting practices on a pioneer front and tests the rela-
tion between game yield, hunting efficiency and forest Study Site
cover.
We also compare our results with those of a similar The survey took place in the Uruará township (Pará,
study conducted in the same region at the beginning of Brazil), 180 km west of Altamira on the Transamazon
its colonization (Smith 1976) in order to detect any highway, between 0251 S¢ and 0416 S¢ and 5309 W¢
change in the hunting methods and game yield. and 5417 W¢ (Fig. 1). Since 1970, this township has

Fig. 1 Location of the study


site
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grown rapidly with the immigration of farmers coming 2. Reports: hunters filled out activity report sheets every
from southern and northeastern regions, with a limited time they hunted, even when hunting was not suc-
knowledge of the Amazonian environment. Actually, cessful. A hunt was considered successful when at
13,000 urban and 32,000 rural inhabitants live on the least one animal was killed. A scientist (T.B.) visited
10.666 km2 of the township (IBGE 2000). Human den- each hunter weekly in order to help him fill out the
sity is low, about 4 inhabitants/km2, and is concentrated hunting activity report sheets.
along the communication axes (Transamazon highway 3. Direct observations: The same scientist accompanied
and trails). each hunter, two to three times to validate the data
Farming is the main activity, with more than 6,500 collected during the interview.
rural households. More than 70% of the agricultural
These methods allowed collecting individual data on
exploitations have an area of less than 150 ha (Veiga
methods, frequencies and duration of hunting, as well as
et al. 1996). Production is divided between annual (rice,
the number of participants and the game yield.
cassava, corn, bean) and perennial crops (pepper, coffee,
Consumed game species were allocated to three
cacao), as well as extensive cattle ranching (Ferreira
groups according to body weight. ‘‘Small’’ species weigh
2001; Toni 2003). Game meat is an important part of the
less than 10 kg: armadillo, Dasypus novemcinctus and D.
diet for settlers who generally live in isolation. The
septemcinctus; paca, Agouti paca; Brazilian agouti,
average game meat consumption is 16.42 kg/individual/
Dasyprocta leporina; marail guan, Penelope marail.
year (Bonaudo et al. 2001). Lumbering is the second
‘‘Medium-size’’ species weigh from 10 kg to 20 kg: col-
economic activity in the township and employs 1,500–
lared peccary, Pecari tajacu and gray brocket deer,
2,000 people.
Mazama gouazoupira. ‘‘Large’’ species weigh more than
Anthropogenic degradation is still limited: in 1999,
20 kg: white-lipped peccary, Tayassu pecari; red brocket
the forest ecosystem represented more than 85% of the
deer, M. americana; capybara, Hydrochaeris hydrochae-
surface of the township and the yearly deforestation rate
ris. Jaguar (Panthera onca), margay cat (Leopardus
was only 1% from 1992 to 1999 (Venturieri et al. 2003).
wiedii) and coati (Nasua nasua) were also hunted but not
The climate is equatorial, warm and humid with a rainy
consumed.
season from December to May and a mild dry season
Game gross weight for each consumed species was
the rest of the year.
estimated by multiplying the number of animals killed
by the mean gross weight of the species. The values used
were provided by Fonseca et al. (1996) for mammals and
Study period and hunters
Thery et al. (1992) for guan marail.
Hunter efficiency was defined as the mean gross
This study focuses on rural hunters, for whom game is
weight of consumed game harvested by a hunter during
an important nutritional part of their diet. We studied
an hour. Values were compared according to hunting
the practices of 40 hunters living along 15 trails. The
method (non parametric Mann–Whitney U test) and to
surveys were conducted in two periods, from May 27th
forest recovery rate of the hunting territory (non para-
to September 29th of 1997 (127 days), and from April
metric Kruskall–Wallis test and Dunn’s Multiple Com-
28th to July 18th of 2000 (81 days). The surveys repre-
parisons test).
sented 133 and 94 hunting expeditions, respectively.

Hunting territory
Data collection
Each hunter had two to five favorite hunting locations
This study was completed successfully thanks to the
situated at a distance of 0–20 km from their house.
collaboration of Embrapa Amazônia Oriental agents
However, more than 95% of the hunting is carried out
and rural development technicians having the trust of
on foot, at a maximum range of 5 km from the house.
the population. This allowed us to accompany them and
Therefore, each hunting territory was defined as a disk
present this research to the communities. We worked
of 10 km in diameter centered on the house of the
with the most interested colons. The first hunters who
hunter.
joined the project led other hunters to collaborate. The
The forest coverage rate of each individual hunting
fact that one of the scientists (T.B.) lived in the township
territory was calculated from a 1999 satellite picture
during several months prior to the study and knew the
(Landsat ETM+ scene). After correction, the channels
hard realities of pioneers’ life also facilitated to build
3, 4 and 5 of the picture were organized by automatic
relations of trust.
algorithms, according to a semi-supervised method
Three methods were used to study hunting practices:
elaborated from fit together masks (ERDAS Imagine
1. Interviews: hunters were questioned about their software).
practices, i.e. hunting methods, hunting frequency Knowledge of the study site and the analysis of the
according to season, harvested game species and satellite picture helped us to isolate main structures like
locations. water bodies, vegetation, roads, and bare soils. Then,
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those structural entities were divided in subtypes corre- According to the interviews, hunters go hunting on
sponding to different ecosystems: forest, fallows, pas- an average 3.6 times a month during the dry season and
tures, etc. Forest recovery rate of each hunting territory only once every 2 months during the rainy season (an-
was determined by superimposing the disks on the nual mean=2). During our survey, hunters went out on
transformed image. These rates were then allocated to an average of 2.3 times a month (SD=1.8, min=0 and
one of the three following types of habitat: max=7.7 hunt/month). Only three hunters did not
shoot during our study period.
Type 1: 49–65% of forest (n=8 territories)
Type 2: 66–85% of forest (n=19)
Type 3: more than 86% of forest (n=10).
Hunted species

Pacas and armadillos represent 61% of the animals


Results killed, with 81 and 46 individuals, respectively (Table 1).
Collared peccaries and red brocket deer are also fre-
Hunting methods
quently hunted. Transformed to gross weight values
(Table 1), the same game species come first with paca
Beating and stand hunting are the two methods used
(664 kg, 26%) followed by armadillo (570 kg, 22%) and
locally. A hunter chooses the technique according to his
collared peccary (522 kg, 20%).
preference and the opportunity to have dogs. The search
for a specific kind of game and the ecological charac-
teristics of the hunting location do not influence his
choice. Game yield
Beating is done by one to six individuals with their
dogs (1–12) and is carried out throughout the year. It A mean number of 0.92 animals are killed during a hunt,
begins at dawn and ends at noon, so that dogs can corresponding to a mean gross weight of 6.33 kg of
use fresh tracks on the trails. Beatings represent 44.5% game (n=227 hunts). Beating is significantly more effi-
(101/227) of all hunting. Peccaries, felines and coatis cient than stand hunting: 3.01 versus 1.91 kg (Mann–
are killed exclusively during beatings. Hunting of Whitney U test, P=0.02; Table 2). Game species differ
brocket deer is mainly done by beating but deer are according to the technique used: medium and large
occasionally killed during stand hunting (Table1). species like deer and peccaries are mainly hunted during
Beating represents only 11% (n=5) of hunts con- beating whereas small species like pacas and armadillos
ducted in hunting territories of type 1 but is prefer- are killed during stand hunting.
entially practiced in more forested hunting territories The percentage of successful hunts is high in all types
(50 and 55% of hunts in hunting territories of type 2 of habitat: 65, 63.9 and 69.7% respectively in types 1, 2
and 3, respectively). and 3. Yet, hunter efficiency varies according to the type
Stand hunting is practiced alone, generally at night of habitat (Kruskall–Wallis test, P=0.01; Table 3): it is
during the dry season. The hunter hides and waits at the significantly higher in type 3 than in type 1 (2.84 vs.
bottom of a tree producing fruits that are sought by 1.53 kg/hunter/hour, Dunn’s Multiple Comparisons
game. This method represents 55.5% of all hunting (126/ Test, P<0.01) and in type 2 than in type 1 (2.53 vs.
227). Armadillos and pacas are mainly killed during 1.53 kg/hunter/hour, Dunn’s Multiple Comparisons
stand hunting at night (respectively 63 and 79%, Ta- Test, P<0.05). Table 1 details hunting efficiency for
ble 1). This method is mainly practiced in low forested each consumed species and habitat type during 100 h. It
hunting territories, representing 89% (n=41) of the is based on hunting of known durations of this study i.e.
hunts conducted in hunting territories of type 1. includes stand hunting and beating in the proportion
Therefore forest recovery rate seems to influence the showed in Table 2.
choice of hunting method.
Hunters use 0.16 to 0.36-caliber rifles and home-made
ammunition. Hunting methods and tools are rudimen- Prey size
tary.
The size of the killed animals depends on the forest
Duration and frequency of hunts coverage rate (Fig. 2). In hunting territories of type 1,
86% (31/36) of the animals killed are small and only 3%
Hunting duration is highly variable, from 30 min to (1/36) are large. In highly forested hunting territories of
12 h. Mean hunt duration is 3 h and 47 min (n=185 type 3, large species represent 25% (20/77) of the ani-
hunts of known duration) and does not differ mals killed. The higher the forest coverage rate, the
according to the method used (Mann–Whitney U test, higher the proportion of large species killed. The largest
P=0.38) or among habitat types (Kruskall–Wallis test, proportion of medium size prey is encountered in
P=0.78). hunting territories of type 2.
Table 1 Hunted species and number of animals killed according to the method used; relative importance of species hunted according to gross weight; harvested game by a hunter during 100 h according to the
forest coverage rate of the hunting territory by weight (227 hunts)

Scientific Brazilian Number of % of total % of animals % of animals Unitary mean total gross % of total Number of animals Gross weight (kg) killed
Name Name* animals animals killed during killed during gross weight weight (kg) gross killed by a hunter by a hunter during 100 h
killed killed beating stand hunting (kg)* weight uring 100 h according according to the forest
to the forest coverage coverage rate of the
rate of the hunting hunting territory #
territory #

Type 1 Type 2 Type 3 Type 1 Type 2 Type 3

Agouti paca Paca 81 38,9 21 79 8.2 664.2 26.03 13.28 13.22 9.67 108.87 108.37 79.32
+
Dasypus sp. Tatu 46 22,1 37 63 2.9 570 22.34 5.64 6.22 9.39 16.35 18.05 27.24
Pecari tajacu Caititu 30 14,4 100 0 19 522 20.46 0.00 3.04 1.35 0.00 57.80 25.61
Mazama americana Veado-mateiro 18 8,6 87 13 29 210 8.23 0.00 1.82 3.02 0.00 52.66 87.44
Mazama gouazoupira Veado-catingueiro 8 3,9 63 37 16 200 7.84 3.58 0.35 0.16 57.25 5.66 2.55
Tayassu pecari Queixada 7 3,4 100 0 30 128 5.02 0.00 0.00 0.43 0.00 0.00 12.87
Dasyprocta leporina Cutia 5 2,4 33 67 2.2 133.4 5.23 0.00 3.61 3.68 0.00 7.95 8.09
Hydrochaeris Capivara 4 1,9 100 0 50 11 0.43 0.25 0.15 0.69 12.25 7.53 34.72
hydrochaeris
Penelope marail Jacu 4 1,9 0 100 1 4 0.16 0.00 0.31 0.74 0.00 0.31 0.74
Leopardus wiedii Gato-maracajá 3 1,5 100 0 3.2 9.6 0.38 0.00 1.17 0.00 0.00 3.73 0.00
Panthera onca Onça 1 0,5 100 0 94.5 94.5 3.70 0.00 0.24 0.00 0.00 22.77 0.00
Nasua nasua Quati 1 0,5 100 0 5.1 5.1 0.20 0.00 0.15 0.00 0.00 0.77 0.00
Total 208 2551.8 22.75 30.28 29.13 194.72 285.60 278.58
+
* From Fonseca et al. (1996) except Penelope marail, from Thery et al. (1992) D.novemcinctus and D. septemcinctus # Territories: type 1, 2 and 3: 49 to 65%, 66% to 85%, 86% and more
203
204

Table 2 Hunting output and hunter efficiency according to the method used

All hunts (n=227) Hunts of known duration (n=185)

Type of hunt Number of Mean number Mean gross Number Mean Mean number Mean gross Hunter
hunts of animals weight/hunt of duration of of animals weight/hunt efficiency
killed/hunt (kg/hunt) hunts hunt(hours) killed/hunt (kg/hunt) (kg/hunter/hour)

Stand hunting 126 0.83 6.4 99 3 h 40’ 1.05 8.06 1.91


Beating 101 1.02 17.3 86 3 h 56’ 1.10 19.03 3.01

Amazon region focus on the sustainability of the activity


Discussion (see for example Bodmer and Robinson 2004). A few
studies investigated the hunting techniques used by non
According to various authors, small species like agouti, indigenous hunters, and more precisely the adaptation
armadillos (Naughton-Treves et al. 2003) or even col- of these techniques to the ecological characteristics of
lared peccaries (Altrichter and Boaglio 2004) resist well the environment.
to anthropogenic pressures such as hunting and eco- In a survey done in the same region in 1976, Smith
system transformation. Our study shows that the num- described the same techniques of hunting and ob-
ber of animals killed and hunting efficiency are related to served similar seasonal variations of hunting frequen-
the forest recovery rate of the hunting territory. The cies. Most of the hunted species are found in both
more deforested a hunting territory, the smaller the studies: armadillos, agoutis, peccaries and deer. The
species hunted and the less efficient the hunter. The small most remarkable difference is the tapir (Tapirus ter-
species hunted in degraded habitat are more prolific, restris) which provided 18.5% of game weight in 1976
more tolerant to second-growth mosaics (Peres 2001a) but was not killed during the present survey. The
and consequently better adapted to a changing envi- weak reproductive rate of the species makes it very
ronment. As game yield reflects the characteristics of the sensitive to overhunting and to the transformations of
habitat where hunting occurs, it can be used as an its natural habitat (Fragoso 1991; Bodmer 1997).
indicator of the degradation of wooded ecosystems. Consequently, it is one of the first species to disap-
Hunters also adapt their hunting techniques accord- pear. Other authors have demonstrated that large and
ing to the habitat and the game species present. In the medium size mammals are sensible to human impacts
most deforested areas where small game is abundant, the (Lopes and Ferrari 2000; Jerozolimski and Peres 2003)
hunters practiced preferentially stand hunting. The and forest fragmentation (Peres 2001a; Altrichter and
density of fruit trees is low in deforested areas and fauna Boaglio 2004). For example, there is a relation be-
concentrates on these rare feeding spots, turning them tween the size of mammal species and fragmentation
favourable stations for hunt. Only five beatings were in Brazilian Atlantic Forest (De Oliveira et al. 2004).
conducted in the most deforested type of hunting terri- Sudden local extinctions of large mammals (tapir and
tories. All were opportunistic huntings of isolated ani- white lipped peccaries) has been recorded in forest
mals previously seen in cultivated areas or pastures. fragment and poaching is suggested to override other
Lastly, stand hunting does not interfere with the agri- effects of fragmentation (Cullen et al. 2004). Peres
cultural work since it is generally practiced at night. (2001a) predicted that the extinction rate of verte-
In forested areas, hunters prefer to make beatings brates is a function of fragment size in Amazonian
because this technique enables them to kill larger prey. forests, and is aggravated by hunting pressure. Hunt-
Moreover, hunting efficiency (in terms of gross weight/ ing and fragmentation are linked. Fragmentation in-
hour/hunter) is superior during beating than stand creases the access to forest and hunting pressure as
hunting (12 and 30% in type 2 and 3, respectively). It is well. Hunting extirpates key vertebrate seed dispersers
worth noting that most research on hunting in the

Table 3 Hunting output according to the forest coverage rate of the hunting territory (type 1, 2 and3: 49 to 65%, 66% to 85%, 86% and
more)

All Hunts (n=227) Hunts of known length (n=185)

Type of Number of Mean number Mean gross Number of Mean duration Mean number Mean gross Hunter
hunting hunts of animals weight/hunt hunts of hunt(hours) of animals weight/hunt efficiency
territory killed/hunt (kg/hunt) killed/hunt (kg/hunt) (kg/hunter/hour)

Type 1 40 0.90 7.35 34 4 h 01’ 1.03 8.56 1.53


Type 2 111 0.86 10.25 83 3 h 37’ 1.05 12.33 2.53
Type 3 76 1.01 14.74 68 3 h 53’ 1.13 16.48 2.84
205

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