|Macuxi hunting patterns and game choice in Roraima
Joel N. Strong1 and José M. V. Fragoso2
1824 10th Avenue North, Saint Petersburg, FL 33701, USA; email: firstname.lastname@example.org
Wildlife and hunting play an important socio-economic role to rural peoples of the neotropics, both as a source of protein and as a way of life (Redford and Robinson 1987; Robinson, Redford et al. 1999; Peres 2000). Furthermore, the most important and heavily hunted game species tend to play important roles in the structuring and maintenance of ecosystems (Redford 1992; Fragoso 2005) as seed dispersers (Fragoso 1997; Strong and Fragoso 2006) and seed predators (Fragoso 1998). Overhunting by rural indigenous and non-indigeneous peoples is one of the most serious threats facing wildlife in the neotropics (Robinson and Redford 1991; Redford 1992; Bodmer, Eisenberg et al. 1997) and potentially affects up to 98.8 % of the 3.7 million km2 Brazilian Amazon (Peres and Lake 2003). Within the Brazilian Amazon, approximately 20 % of the land area is designated as indigenous reserves. Wildlife of these areas sustain large populations of indigenous people, but the value of these lands as conservation tools remains debatable as some tribes are known to deplete their natural resources through overhunting, overfishing, and overharvesting of timber (Peres 1994). Therefore, understanding the impacts of hunting on game populations and developing management strategies for sustainable wildlife use are important, particularly on lands controlled by indigenous peoples (Peres and Terborgh 1995). This is especially true in Roraima, as nearly 50 % of the area is designated as indigenous land inhabited by the Macuxi, Wapixana, Ingarikó, Yanomami, Waiwai, Yekuana, Waimiri-Atroari, and Taurepang tribes. Thus, understanding the hunting patterns and behaviors by indigenous peoples and the response by wildlife populations is key to sustainable wildlife use in this region and to maintaining the important socio-economic and ecological role of wildlife.
Numerous hunting studies in the neotropics report wildlife harvest rates and the sustainability of hunting by forest-dwelling peoples in the Amazon Basin (Vickers 1991; de Souza-Mazurek, Pedrinho et al. 2000; Mena, Stallings et al. 2000; Peres 2000; Hill, McMillan et al. 2003; Bodmer and Robinson 2004; Hurtado-Gonzales and Bodmer 2004), yet similar studies focusing on savanna and forest-edge inhabiting peoples are considerably less common (Leeuwenberg and Robinson 2000). Although many of the game species hunted may be common to both habitats (i.e. Tapirus terrestris, Tayassu pecari, T. tajacu, Mazama sp., Dasyprocta sp., Agouti paca, Cebus sp., Ateles sp., Geochelone sp., Crax sp., and Tinamus sp.) hunting patterns and harvest rates may vary considerably. We present data on game species harvested in the Mangueira Macuxi Indigenous Reserve and discuss hunting patterns by the Macuxi in the Rio Branco-Rupununi Savanna.
The Macuxi are sedentary, garden-scale agriculturalists who rely on game and fish for protein. Over 20,000 Macuxi live in Roraima, occupying 2,582,566 ha within the state. The Macuxi also extend into bordering Guyana and Venezuela where they inhabit the same savanna biome, which includes low-lying, seasonally flooded grasslands, forest islands, and gallery forests. In Brazil, the Macuxi use certain Western technologies, including the use of motorcycles, motorboats, and shotguns for hunting and fishing. Mangueira Macuxi Indigenous Reserve (Mangueira) is a 4,238 ha indigenous reserve in north-cenral Roraima, located 5 km to the south of Maracá Island Ecological Reserve (Figure 1). Terra firme, lowland, evergreen rainforest covers approximately 60 % of the reserve with the remainder being composed of savanna habitat, where the village is located. The forested portion is contiguous with a vast expanse of forest extending into Maracá (Figure 1). Manguiera is composed of ten families totaling 65 individuals. Some livestock and poultry are raised for trade and consumption, but the majority of dietary protein is procured from hunting (J.M.V. Fragoso, pers. obs.). There is no market for bushmeat in this region of Roraima, and thus, hunting is for subsistence only. Additionally, there is some subsistence farming of corn, rice, and manioc by households on the reserve as well as a small coffee plantation as a cash crop.
Hunting interviews were conducted weekly between 15 March and 18 July 2004 with each household. Two male Macuxi assistants from Mangueira were responsible for conducting the interviews with the head of each household and J. N. Strong periodically checked reported hunting returns with the actual hunters to verify the data reported were accurate. Data were collected on the species, sex, and age of all harvested game species, as well as hunting method and approximate distance from the village where it was harvested. Biomass calculations for each species are based on the reported average weight of each species by
Results and Discussion
A total 192 animals (1,394.57 kg of biomass), representing 25 species were harvested in Mangueira over the four-month interview period (Table 1 and Figure 2). Numerically, reptiles were the most hunted taxa, comprising 45 % of the total, followed by mammals (42 %), and birds (13 %). The most important species were Geochelone carbonaria (N = 54), Dasyprocta leporina (N = 27), Dasypus novemcinctus (N = 25), Tayassu tajacu (N = 16), Caiman crocodilus (N = 15), and Geochelone denticulata (N = 11) comprising 71 % of all species harvested. In terms of biomass, mammals comprised 51 % of the total, followed by reptiles (46 %), and birds (3 %). The most important species, based on biomass, were Geochelone carbonaria (323.46 kg), Tayassu tajacu (280.32 kg), Caiman crocodilus (226.50 kg), Dasypus novemcinctus (88.60 kg), Mazama americana (78.30 kg) and Geochelone denticulata (64.68 kg), contributing to 76 % of the total biomass.
Although studies on hunting in the neotropics have shown that hunters prefer larger-bodied species (Peres 1990; Bodmer 1995) such as ungulates (i.e. Tapirus terrestris, Tayasu spp., and Mazama spp.) and large-bodied rodents (i.e. Hydrochaeris hydrochaeris and Agouti paca) to smaller species, actual harvests do not necessarily coincide with these preferences. Our results show that hunters took a greater proportion of smaller-bodied species such as Geochelone spp., Dasyprocta leporina, and Dasypus novemcinctus at Mangueira during this study compared to larger-bodied species such as Tayassu pecari and Tapirus terrestris. Increasing levels of hunting pressure lead to reduced numbers of large-bodied species and a higher proportion of small-bodied species (Peres 2000). This is likely the scenario at Mangueira. Small-bodied species with high intrinsic growth rates such as Dasyprocta leporina and Dasypus novemcinctus as well as some larger-bodied species such as Mazama americana and Tayassu tajacu can sustain relatively high levels of hunting and maintain relatively high populations due to their high production levels. The intrinsic growth rate of Geochelone spp. are unknown, but their ability to achieve high densities (Moskovits 1988; Aponte, Barreto et al. 2003) and abundance around Mangueira (J. Strong, unpublished data) suggest they can sustain chronic hunting pressure to some degree.
Compared to hunting studies from Amazonian hunting sites, the harvest profile at Mangueira has some unique differences. While Tayassu pecari is often near the top of the list for harvested species in other studies (Redford and Robinson 1987; Vickers 1991; de Souza-Mazurek 2001), only two were reported in Mangueira during this study. During trips into the forest around the village, tracks of this species were only seen in a Mauritia flexuosa swamp over 7km from the village (J. Strong, pers. obs.). This is a highly preferred species by the Macuxi and hunts were often organized for the following day if a hunter reported fresh sign or had encountered a herd. Although few individuals were reported during the present study, villagers reported a large herd moving through the village and a neighboring village over the course of several weeks where at least 13 individuals were harvested approximately a year later. Fragoso (2004), suggests that the apparent cycle of abundance and reduction in T. pecari populations is the result of epidemic outbreaks of disease that kill off most individuals in the herd and which may eventually be extirpated by local hunting, resulting in disappearance of the species in the area. This is followed by the reappearance of the species from the process of recolonization by distant herds moving over large geographical areas. The cycle may occur every 8-12 years in northern Roraima where hunting occurs. This may account for the lack of T. pecari in our data set and its importance in other data sets which span over long time periods or over broad geographical areas (Redford and Robinson 1987; Peres 2000).
The importance of Geochelone spp. in our study is one pattern consistent other published hunting studies. Geochelone denticulata was the second-most hunted species by the Waimiri-Atroari (de Souza-Mazurek 2001) and Redford and Robinson (1987) report that out of nine Indian studies where reptiles were recorded, Geochelone spp. ranked first in three of them. In a study examining the consumption rate of game vertebrates harvested in Brazilian Amazonia, Peres (2000) places Geochelone spp. as eleventh out of 53 species and first among reptiles. In our study, Geocheone carbonaria was by far the most hunted species, both numerically and in terms of biomass, by the Macuxi. The dominance of G. carbonaria in this study as opposed to G. denticulata is due to the difference in habitat preference by the two species. Geochelone carbonaria attains much higher densities in the forest-savanna ecotone of our study area than does G. denticulata, which tends to be restricted to the moist interior Amazonian forests as occupied by the Waimiri-Atroari.
Tayasu tajacu is another species consistently high on the list of game species harvested for studies in Amazonia. The relatively high intrinsic growth rate of this species and the behavior of small herds scattering when threatened afford them the ability to sustain high hunting pressures without being extirpated. Their relatively large body size also make them a highly preferred species over the smaller mammals and bird species. The use of dogs to hunt this species has made the Macuxi extremely efficient in hunting T. tajacu, yet populations appear to persist in the area as evidenced by numerous tracks both near and far from the village (J. Strong, pers. obs.).
Although not highly preferred, Dasyprocta leporina and Dasypus novemcinctus were the second and third most harvested species, respectively, after Geochelone carbonaria. Dasyprocta leporina is often hunted with the use of dogs, while D. novemcinctus is either dug out its burrow during the day or hunted with a headlamp or flashlight at night. Again, these are both species with high population production rates, whose populations can sustain high off-take levels with little risk of becoming extirpated. Their abundance in the harvest data supports the idea that Mangueira is a heavily hunted site with extremely low densities of the larger-bodied vertebrates. This was also evidenced by the very few tracks of Tapirus terrestris seen throughout the forest in Mangueira except beyond 7-9km from the village (J. Strong, pers. obs.).
In most Amazonian hunting studies primates are often a highly preferred taxa, whose populations quickly decline with increasing hunting pressure (Peres 1990; Peres 2000). The Macuxi, however do not prefer primates and will only hunt them if other game is scarce. This was supported in the harvest data with only one individual of Ateles belzebuth harvested over the four-month study. Troops of A. belzebuth and Cebus apella were often encountered in the forest around Mangueira and exhibited little fear of humans. Myrmecophaga tridactyla is another species that is important to some Indigenous cultures, but is not hunted by the Macuxi and is considered a taboo.
Hunting at Mangueira is done primarily in the forested portion of the reserve with full-day trips extending up to 10km into the forest along a main hunting trail. This forest is contiguous with Maracá (Figure 1), which likely acts as a game source, enabling source-sink dynamics to maintain populations to some degree within the Macuxi reserve. However as colonists continue to settle and encroach on the reserve, this connectivity is decreasing and may result in decreased movement of game species from source areas to the hunted forests around Mangueira. The Macuxi in Mangueira have voiced their concern for this and desire to secure more land around the reserve.
Hunting by the Macuxi in Brasil and Guyana is strongly tied to spacio-temporal shifts in game and fish movement. This is especially pronounced in the forest island-gallery forest-savanna mosaic in which the Macuxi inhabit. During the rainy season, the low-lying savannas often become flooded, pushing many game species into the higher-elevation forest islands. Game species become concentrated in these forested patches and the Macuxi capitalize on this trend by concentrating their hunting efforts during this time of year in order to maximize yields. Conversely, during the dry season when the waters recede, the Macuxi tend to focus on fishing as a source of protein and turn to the nearby Uraricoera River and its tributaries, as well as numerous ponds and lakes as important sources of food. Each household also maintains a small farm where they grow maniac, corn, bananas, papayas, and other crops that attract wildlife. Game is often harvested on and around these farms. A firearm is usually carried anytime work is done on the farm and game species are often encountered either en route or at the farm. Some Macuxi in other villages in Brasil and Guyana have farms that a quite distant from the village and thus, require overnight stays. Nocturnal species such as Agouti paca and Mazama spp. are often encountered during these overnight trips.
This study is one of the few that address hunting practices at the forest-savanna ecotone and shows many of the similarities in game species composition and harvest rates that exist between this site and other sites in the interior Amazon. Managing wildlife in this region to maintain ecologically and socio-economically healthy game population requires knowledge of the complex interaction between wildlife, the environment, and man. Understanding how wildlife species respond to spatio-temporal patterns in the complex northern Roraima landscape will do no good if it does combine that knowledge with hunting behavior and the needs of the people that rely on that resource.
Aponte, C., G. R. Barreto, et al. (2003). "Consequences of habitat fragmentation on age structure and life history of a tortoise population." Biotropica 35(4): 550-555.
Bodmer, R. E. (1995). "Managing Amazonian wildlife: Biological correlates of game choice by detribalized hunters." Ecological Applications 5(4): 872-877.
Bodmer, R. E., J. F. Eisenberg, et al. (1997). "Hunting and the likelihood of extinction of Amazonian mammals." Conservation Biology 11(2): 460-466.
Bodmer, R. E. and J. G. Robinson (2004). Evaluating the sustainability of hunting in the Neotropics. People in nature: wildlife conservation in South and Central America. K. M. Silvius, R. E. Bodmer and J. M. V. Fragoso. New York, Columbia University Press: 299-323.
de Souza-Mazurek, R. R. (2001). Kinja txi taka nykwa myrkase: Hunting and fishing among the Waimiri Atroari Indians of central Amazonia. PhD Dissertation. Chicago, University of Chicago. PhD: 115.
de Souza-Mazurek, R. R., T. Pedrinho, et al. (2000). "Subsistance hunting among the Waimiri Atroari Indians in central Amazonia, Brazil." Biodiversity and Conservation 9: 579-596.
Fragoso, J. M. V. (1997). "Tapir-generated seed shadows: Scale-dependent patchiness in the Amazon rain forest." The Journal of Ecology 85(4): 519-529.
Fragoso, J. M. V. (1998). White-lipped peccaries and palms on the Ilha de Maracá. Maracá: The biodiversity and environment of an Amazonian rainforest. W. Milliken and J. A. Ratter. New York, Johy Wiley & Sons: 151-163.
Fragoso, J. M. V. (2004). A long-term study of white-lipped peccary (Tayassu pecari) population fluctuations in northern Amazonia. People in Nature: Wildlife Conservation in South and Central America. K. M. Silvius, R. E. Bodmer and J. M. V. Fragoso. New York, Columbia University Press: 286-295.
Fragoso, J. M. V. (2005). The role of trophic interactions in community initiation, maintenance, and degredation. Biotic interactions in the tropics; their role in the maintenance of species diversity. D. Burslem, M. Pinard and S. Hartley. New York, Cambridge University Press: 310-327.
Hill, K., G. McMillan, et al. (2003). "Hunting-related changes in game encounter rates from 1994 to 2001 in the Mbaracayu Reserve, Paraguay." Conservation Biology 17(5): 1312-1323.
Hilty, S. L. (2003). Birds of Venezuela. Princeton, New Jersey, Princeton University Press.
Hurtado-Gonzales, J. L. and R. E. Bodmer (2004). "Assessing the sustainability of brocket deer hunting in the Tamshiyacu-Tahuayo Communal Reserve, northeastern Peru." Biological Conservation 116: 1-7.
Leeuwenberg, F. J. and J. G. Robinson (2000). Traditional management of hunting by a Xavante community in Central Brazil: The search for sustainability. Hunting for sustainability in tropical forests. J. G. Robinson and E. L. Bennet. New York, Columbia University Press: 375-394.
Mena, P., J. R. Stallings, et al. (2000). The sustainability of current hunting practices by the Huarani. Hunting for Sustainability in Tropical Forests. J. G. Robinson and E. L. Bennet. New York, Columbia University Press: 57-78.
Moskovits, D. K. (1988). "Sexual dimorphism and population estimates of the two Amazonian tortoises (Geochelone carbonaria and G. denticulata) in northwestern Brazil." Herpetologica 44(2): 209-217.
Peres, C. A. (1990). "Effects of hunting on Western Amazonian primate communities." Biological Conservation 54: 47-59.
Peres, C. A. (1994). "Indigenous reserves and nature conservation in Amazonian forests." Conservation Biology 8(2): 586-588.
Peres, C. A. (2000). "Effects of subsistence hunting on vertebrate community structure in Amazonian forests." Conservation Biology 14(1): 240-253.
Peres, C. A. and I. R. Lake (2003). "Extent of nontimber resource extraction in tropical forests: Accessibility to game vertebrates by hunters in the Amazon." Conservation Biology 17(2): 521-535.
Peres, C. A. and J. W. Terborgh (1995). "Amazonian nature reserves: An analysis of the defensibility status of existing conservation units and design criteria for the future." Conservation Biology 9(1): 34-46.
Redford, K. H. (1992). "The empty forest." BioScience 42(6): 412-422.
Redford, K. H. and J. G. Robinson (1987). "The game of choice: patterns of indian and colonist hunting in the neotropics." American Anthropologist 89: 650-667.
Robinson, J. G. and K. H. Redford (1986). "Body size, diet, and population density of Neotropical forest mammals." The American Naturalist 128(5): 665-680.
Robinson, J. G. and K. H. Redford, Eds. (1991). Neotropical wildlife use and conservation. Chicago, University of Chicago.
Robinson, J. G., K. H. Redford, et al. (1999). "Wildlife harvest in logged tropical forests." Science 284(5414): 595-596.
Strong, J. N. and J. M. V. Fragoso (2006). "Seed dispersal by Geochelone carbonaria and Geochelone denticulata in northwestern Brazil." Biotropica 38(5): 683-686.
Vickers, W. T. (1991). Hunting yields and game composition over ten years in an Amazon Indian territory. Neotropical Wildlife Use and Conservation. J. G. Robinson and K. H. Redford. Chicago, The University of Chicago Press: 53-81.
Table 1. Number and biomass of game species harvested between 15 March and 18 July
004 in Mangueira Macuxi Indigenous Reserve, northern Brazil.
Figure 1. Location of Mangueira Macuxi Indigenous Reserve and Maraca Island Ecological Reserve in northern Roraima
Figure 2. Total number (A) and biomass (B) of the most important species harvested by the Macuxi in Mangueira Macuxi Indigenous Reserve