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Foraging in the fallows: Hunting patterns across a successional continuum in the Peruvian Amazon
Michael C. Gavin*
Victoria University of Wellington, School of Geography, Environment, and Earth Sciences, P.O. Box 600, Wellington, New Zealand

A R T I C L E I N F O

A B S T R A C T

Article history: Received 27 April 2006 Received in revised form 21 July 2006 Accepted 28 July 2006 Available online 9 October 2006 Keywords: Conservation Development Sustainable harvest Secondary forest Tropical rainforest

Studies comparing hunting between secondary and mature forests are critical to understanding secondary forests potential as sustainable hunting grounds. I examined hunting across a successional continuum by surveying 67 households in three communities near the Cordillera Azul National Park, Peru, and analyzing the potential for sustainable hunting. Ninety-nine percent of households surveyed went hunting during the six-month study. Ninety-one species were recorded from seven vertebrate and invertebrate classes, with mammals the most hunted. Five percent of extraction events were species of concern on IUCNs Red List. Households extracted signicantly more, in terms of number of species, number of collection events, and biomass, from older forests (>20 y) than from young secondary (15 y) or old secondary forests (520 y). However, when extraction is measured per unit area (kg/km2 or collection events/km2), households extracted more wildlife from old secondary forest. Households consumed meat at rates below Amazon regional averages. However, because human population densities are well above carrying capacity, current low harvest rates are likely a relic of past overharvests and do not reect sustainability. Even if management focuses on source-sink dynamics with buffer zone hunting resupplied by protected area populations, long-term sustainability seems doubtful. As more agricultural clearing converts older forests to elds and fallows, the role of secondary forests in resource management plans will increase. The impact of high human population densities in the region means conservation and development programs should focus more on supplying alternative sources of protein and income and limiting immigration into the area. 2006 Elsevier Ltd. All rights reserved.

1.

Introduction

Both land conversion and hunting are major threats to tropical wildlife. Agricultural clearing has created a mosaic of much of the worlds tropical forests. Areas of mature forest are often surrounded by a patchwork of secondary forests, elds, and pastureland. Hunting has drastically reduced wildlife numbers in many areas of the tropics, and is considered the major cause of population decline in one-third of threatened bird and mammal species (Rosser and Mainka, 2002).

In addition, hunting has led to numerous local extinctions (Rao and McGowan, 2002 and references therein). In areas with larger human populations, hunting has emptied forests of game, dramatically altering forest species compositions, and leading to a cascade of ecological impacts (Redford, 1992; Chapman and Onderdonk, 1998; Wright et al., 2000; Kaiser and Jennings, 2001; Wright, 2003). A synergy exists between agricultural forest clearance and the impact of hunting on wildlife. Wildlife is a critical resource for rural poor throughout the tropics, with bushmeat

* Tel.: +64 4 463 5195; fax. +64 4 463 5186. E-mail address: michael.gavin@vuw.ac.nz. 0006-3207/$ - see front matter 2006 Elsevier Ltd. All rights reserved. doi:10.1016/j.biocon.2006.07.011

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acting as a major protein source (Bennett et al., 2000; Rao and McGowan, 2002), and as a means of income (Bodmer et al., 1994; Eves and Ruggiero, 2000). Increased access to forest interiors, caused by migration to frontiers and more roads, facilitates forest clearing and leads to more hunting. With more people clearing forests, hunting is bound to increase in frequency. In addition, agriculture presents those species capable of adapting to a human-dominated landscape with food supplies in the form of crops. Crop raiding can cause substantial losses for farmers on the forest frontier, and these losses are often balanced with the gains from hunting in elds and fallows (Naughton-Treves et al., 2003; Smith, 2005). Some wildlife species, particularly those with short-lived individuals, shorter generation times, and higher intrinsic rates of increase, can actually increase in abundance in humandominated environments due to forest disturbance or even in response to harvesting (Bodmer et al., 1997; Lopes and Ferrari, 2000; Peres, 2000). Robinson and Bennett (2004) hypothesize that secondary forests may have a greater supply of wildlife than either mature forests or newly cleared elds, and that the secondary forest supply of meat may often be greater than the demand from hunting. Therefore, hunting in these secondary forests has the greatest potential to be sustainable in the long-term. Such a scenario would support conservation and development programs that seek to establish multiple use areas in buffer zones surrounding protected areas, where human inuences lead to more secondary forest cover. These zones might take advantage of the large numbers of anthropogenic species in fallows while still protecting mature forest obligates within adjacent protected areas. Such a strategy could meet development goals by providing local people with access to the secondary forest game species, which are generally not a major concern for the conservation efforts focused on mature forest species. However, the potential of these secondary forests to act as reservoirs for wildlife depends on numerous factors, including human population density, cultural norms, and ecology, which determine the ultimate impact hunting has on wildlife (Cuaron, 2000; Peres, 2000; Robinson and Bennett, 2000). Quantitative studies that compare hunting between secondary and mature tropical forests are critical to understanding the role secondary forests and buffer zones can play as sustainable hunting grounds. To date, no studies have examined wildlife use across a successional continuum in forests of different ages. Researchers have focused on hunting in elds/gardens versus mature forests and found mixed results with some studies showing more hunting in older forests (e.g., Naughton-Treves et al., 2003) and others the opposite trend (e.g., Smith, 2005). My research contributes to the literature on tropical hunting by examining the use of fauna in young secondary forests, old secondary forests, and older forests in the buffer zone of a national park in the Peruvian Amazon and analyzing the potential for sustainable hunting in this region.

2.

Methods

My research focused on three human communities in the northwestern section of the Cordillera Azul National Parks buffer zone. I selected communities with similar population

sizes and forest ecology. The communities sampled depend on slash-and-burn agriculture, hunting, and gathering of river and forest products for subsistence. All three communities are situated within lowland tropical moist forest between 200 and 500 m. I selected houses at random (community leaders pulled names from a hat) from the pool of potential candidates. Their homes had to be within two hours walk of the community center, they had to be permanent residents of the community, and they had to be willing to participate in the study (ve households in San Rafael, three in Nuevo Lima, and two in Alianza refused to take part). Due to the sensitivity of the information collected, particularly concerning illegal resource use, the names of the communities have been changed. I surveyed approximately one-third of all households in each community for a total of 67 households. The participating households represented the mixed ethnic heritage of communities in the northwestern sector of the parks buffer zone, with both indigenous Quechua Lamista families and non-indigenous colonists of Amazonian, Andean, and Pacic coast origins taking part in the study. Some households had a history in the forest dating back 500 years, others had been in the region six or seven generations, whereas still others arrived only months before the study began. Time since agricultural elds were abandoned was used to dene forest stages due to the consistent land use histories in the region. Subsistence agriculture has been the principal use of land by people in the region. Farmers in the region do manage fallow lands via occasional trips to collect fruits from planted crops and to encourage growth of planted species with weeding. The most accurate assessment of forest stages would combine information on oristics, forest structure, and ecosystem function (Guariguata and Ostertag, 2001), but this typology is not easily applicable in a cross-cultural setting. The ability of informants to differentiate among forest types was highly variable. Many indigenous households, especially those with long histories in the region, recognized a complex forest classication system, including seven or eight different forest types. Recent immigrants could often only differentiate three forest types based on their memory of land use histories. I relied on informants memory to classify forest stands into three categories: young secondary forests (forests 15 years old), old secondary forest (520 years old), and older forests (>20 years old). These stages are equivalent to the locally used categories of purma joven (young fallow), purma antigua (old fallow), and monte alto (tall bush). Because the histories of forests located beyond the communities boundaries were not known, the possibility existed that these forests had never been cleared and should be termed primary and not older forests. However, due to limits in the ability of some local informants to classify forest ages, all forests over 20 years were designated older forests. The use of forest fauna was recorded over six months covering both wet and dry seasons. Active participation by informants in the data collection process ensured contributions were made by both illiterate and literate community members, and also helped avoid informant fatigue. The use of participatory methodologies also engages local hunters in the early stages of monitoring and management of local wildlife populations, a critical step to successful community-based

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conservation efforts (Bodmer and Robinson, 2005). The data informants gathered included the name of the species collected and the quantity, the forest type where the product was collected, and whether the animal product was sold (along with its price). Additional information, including exact size and quantities of products, was gathered in weekly semistructured interviews. Informants identied where each collection event took place: on their land, on a neighbors land, or on open-access land. I estimated average weights of the quantities collected for each species from interviews or based on sample weighings. Because I could not observe each collection event during all phases of this study, the identities of all fauna were recorded using local classication schemes (i.e., folk names). Folk species do not always coincide with scientic species (e.g., perdiz refers to multiple species of tinamou (Tinamus spp.)). Collection areas were estimated separately by forest type for products from an informants own land, those from a neighbors land, and those from open-access land. In semistructured interviews I asked participants to list all their land holdings and the ages of all forest types they owned. These forest ages were cross-referenced against major events in the lives of each participating family (births, deaths, illnesses, etc.). When informants indicated a forest product as collected from a neighbors land, the neighbors land holdings were also estimated. I dened open-access lands as areas not claimed by any community member. Open-access lands were found both mixed with private holdings in the agricultural zones, as well as outside the core agricultural areas. The open-access lands outside the agricultural zones of the communities were all older forest. I accompanied informants on several hunts to estimate the average collection area within the open-access lands outside the agricultural zones. Hunting occurred from established camps and hunters tended to radiate out from these camps in several different directions. During hunting forays, I recorded the average linear distance from camp informants travelled and used this as the radius of a circular collection area for each camp. The average linear distance travelled was four km, and thus the catchment (collection area) for each camp was 50 km2. Inside the agricultural zones, hunting began at the community center from which hunters tended to hunt in a variety of different directions. Again, the average linear distance travelled was four km, and the catchment (collection area) was 50 km2, including both open-access and private lands. I estimated the extent of open-access land by subtracting the area of all private holdings from the total catchment area (50 km2). I determined the composition of open-access lands within the agricultural zone by rst calculating the average percent cover of each forest type in the private land holdings. These percentages were then multiplied by the extent of open-access lands to produce the nal area of each forest type. I compared the number of species collected, number of species collected in only one forest type, number of animal collection events, and the biomass extracted from each of the three forest types by the 67 participating households using Friedman tests with post hoc Nemenyi pairwise comparisons.

3.

Results

Ninety-one animal folk species were recorded from seven different classes. Although I recorded all uses of fauna, the analysis presented here remains consistent with the denition of hunting put forth by Robinson and Bennett (2000), and thus is limited to those species of mammals, reptiles, and birds that are captured by humans. I have not included data in this analysis from the species collected in the four other classes, including eight species of sh, seven species of insects, two species of gastropods, and one amphibian. Forty-ve of the 73 folk species recorded in the three focal classes, and 14 of the 25 most hunted species were mammals (Table 1). Overall, the hunters in the Cordillera Azul use a large number of the mammal species available in the Amazon. Of the 192 species of mammals in the Amazon, a review of the literature by Fa and Peres (2001) found 28% (54 species) were hunted. Fortyve of these 54 species were harvested in the three communities of the Cordillera Azul over the course of this six-month study. Fifty-one (5%) of the collection events in the Cordillera Azul involved the extraction of animals listed as species of concern on the IUCN Red List. These 51 events involved ten different species ranging in conservation status from near threatened to endangered: jaguar (Panthera onca, near threatened), puma (Puma concolor, near threatened), common woolly monkey (Lagothrix lagothricha, near threatened to vulnerable), night monkey (Aotus sp., vulnerable), white-bellied spider monkey (Ateles belzebuth, vulnerable), giant anteater (Myrmecophaga tridactyla, vulnerable), bush dog (Speothos venaticus, vulnerable), Brazilian tapir (Tapirus terrestris, vulnerable), pacarana (Dinomys branickii, endangered), and giant armadillo (Priodontes maximus, endangered). Sixty-six (99%) of the households surveyed went hunting at least once during the six-month study, and 44 households (66%) hunted on more than ten occasions. The number of collection events per household ranged from four to 143, with a median of 51. Of the 1071 collection events recorded by all households, only 35 did not involve killing the animal. These 35 events involved collection of animals for pets and pet trade. Eighty-four percent of the animals collected were used for food. Households used the majority of the animals in all use categories for subsistence, with only 11% sold to market. The households surveyed extracted signicantly more animals, in terms of number of species, number of species collected in only one forest type, number of collection events, and biomass, from older forests than from young secondary or old secondary forests (p 6 0.001; Table 2). However, the households surveyed extracted more wildlife per unit area (i.e., per km2), both in terms of biomass (kg/km2) and collection events (events/km2), from old secondary forest (Table 2). The majority of the collection events (63%; Table 2) occurred on private lands. Of the 20 most used animal species, households collected 16 of them signicantly more in older forests and only two signicantly more from young secondary (Brazilian rabbit, Sylvilagus brasiliensis; common possum, Didelphis marsupialis) (Table 1). For comparison with other studies and sites across the Amazon, I calculated consumption rates, expressed as the

Table 1 Twenty most collected animal folk species by 67 households in the Cordillera Azul, Peru Scientic name Class English name Local name Number of collection events Percent collected per forest type (collection events/km2) Older forest
65***(0.49) 77***(0.36) 92***(0.28) 93***(0.27) 72***(0.19) 92***(0.20) 71***(0.14) 90***(0.17) 10 (0.02) 26 (0.04) 87***(0.12) 94***(0.12) 97***(0.11) 100***(0.12) 81***(0.09) 88***(0.06) 30 (0.02) 36 (0.02) 100***(0.06) 100***(0.05)

Old secondary
19 (3.76) 12 (1.47) 2 (0.16) 6 (0.46) 15 (1.02) 2 (0.12) 13 (0.66) 10 (0.49) 16 (0.69) 18 (0.69) 3 (0.11) 3 (0.10) 3 (0.09) 0 (0.00) 8 (0.24) 0 (0.00) 23 (0.42) 36 (0.57) 0 (0.00) 0 (0.00)

Young secondary
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Dasyprocta fuliginosa Agouti paca Tayassu tajacu Mazama americana Dasypus sp. Penelope spp. Tinamus spp. Crax tuberosa Sylvilagus brasiliensis Didelphis marsupialis Potos avus Geochelone denticulata Sciurus igniventris Tayassu pecari Leopardus pardalis Amazona spp. Ortalis motmot Lontra longicaudis Tapirus terrestris Lagothrix lagothricha * Sig. difference <.05. ** Sig. difference <.01. *** Sig. difference 6.001

Mammalia Mammalia Mammalia Mammalia Mammalia Aves Aves Aves Mammalia Mammalia Mammalia Reptilia Mammalia Mammalia Mammalia Aves Aves Mammalia Mammalia Mammalia

Black agouti Paca Collared peccary Red brocket deer Armadillo Guan Tinamous Currassow Brazilian rabbit Common possum Kinkajou Amazon red squirrel White-lipped peccary Ocelot Parrots Variable Chachalaca Neotropical otter Brazilian tapir Woolly monkey

An uje Majas Sajino Venado colorado Carachupa Pucacuga Perdiz Paujil Conejo Zorro Choshna Motelo Ardilla Huangana Tigrillo Loro catalino Manacaraco Lobo Sachavaca Mono choro

175 108 72 68 60 51 45 43 38 34 31 29 27 27 26 17 16 14 13 12

16 (2.25) 11 (0.95) 6 (0.35) 1 (0.06) 13 (0.63) 6 (0.25) 16 (0.58) 0 (0.00) 74***(2.26) 56*(1.53) 10 (0.25) 3 (0.07) 0 (0.00) 0 (0.00) 11 (0.23) 12 (0.16) 47 (0.60) 28 (0.32) 0 (0.00) 0 (0.00)

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Table 2 Number of folk species and quantities extracted from different forest types in the Cordillera Azul, Peru Forest type Total species
45 49 81 91

Unique speciesa
4 (2) 3 (0) 26 (15)

Biomass collected kg
279 616 6 370 7 265

Collection events (Events/km2) Private land

48 92 540 680 (18) (76) (17) (19)

kg/km
48 84 54 54

Open-access
54 (5) 56 (6) 281 (1) 391 (1)

Total
102 (7) 148 (15) 821 (3) 1 071 (4)

Young secondary Old secondary Older forests All forest types

a Unique species are those only occurring in that forest type. Numbers in parentheses refer to species unique to that forest type represented by more than one collection event (non-singletons).

number of individuals of a species collected per consumer year. Redford and Robinson (1987) dene consumer years as: (365/ number of study days) (number of people consuming the animal products). Table 3 shows the ten most harvested mammals. Three are rodents, three are ungulates, two are carnivores, and there is one edentate and one lagomorph. Three of the most heavily harvested mammals in the Cordillera Azul, black agouti (Dasyprocta fuliginosa), Brazilian rabbit (Sylvilagus brasiliensis), and kinkajou (Potos avus), were consumed at rates higher than the average regional consumption rates calculated by Fa and Peres (2001) based on data from the entire Amazon basin. Various models exist which allow comparisons between actual harvests and estimates of sustainable yields, but most require either long-term monitoring of harvests or detailed biological information on local populations of wildlife (Bodmer and Robinson, 2005), data which are often not available

for remote tropical sites in need of rapid management decisions. Two exceptions are production models, of which the most commonly cited is that of Robinson and Redford (1991), and estimates of human carrying capacity. Actual harvest rates from the Cordillera Azul (Table 3) were less than the potential harvest rates as calculated by Robinson and Redford (1991). I also compared actual harvest rates and current human population densities to three other sustainability estimates: Robinson and Bennetts (2000) estimate of sustainable harvest rates at 152 kg/km2, Hill and Padwes (2000) estimation of potential sustainable yields when 5 km2 are available per consumer, and Robinson and Bennetts (2000) prediction of sustainable yields with human population densities 61 person/km2. In all forest types, harvest rates in the Cordillera Azul were lower than the sustainable estimate calculated by Robinson and Bennett (Table 2). However, when I combined the population sizes of the human communities

Table 3 Harvest and consumption rates of 10 most hunted mammals by 67 households in the Cordillera Azul, Peru Scientic name Local name English name Cordillera Potential Private lands Open-access Amazon regional lands average Azul harvest harvest rated consumption consumption ratec (ind/km2/y) (ind/km2/y) rate (p.c.p.a.)a consumption rate (p.c.p.a.) rate (p.c.p.a)b
0.80 0.40 0.18 0.09 0.20 0.11 0.17 0.26 0.26 0.11 0.66 0.81 0.61 0.31 0.67 3.01 1.30 0.91 0.64 0.82 8.98 1.31 2.41 0.67 5.19

Dasyprocta fuliginosa An uje Agouti paca Majas Tayassu tajacu Sajino Mazama americana Dasypus sp. Venado colorado Carachupa bombero Conejo Zorro Choshna Ardilla colorada Huangana

Sylvilagus brasiliensis Didelphis marsupialis Potos avus Sciurus igniventris Tayassu pecari

Black agouti Paca Collared peccary Red brocket deer Great long-nosed armadillo Brazilian rabbit Common possum Kinkajou Northern Amazon red squirrel Whitelipped peccary

0.19 0.16 0.15 0.09

0.02 0.02 0.02 0.06

0.0004 0.248 0.01 0.14

0.54 1.04 0.43 0.38

N/A N/A N/A 44.83

0.02

0.13

0.83

0.36

0.83

a p.c.p.a., per capita per annum. b From Fa and Peres (2001) except numbers for Tayassu pecari and Didelphis marsupialis from Redford and Robinson (1987) average consumption rates based on studies throughout the Amazon basin. c Harvest rate from Cordillera Azul is a total from private and communal lands. d From Robinson and Redford (1991), rates calculated using data from multiple studies across the neotropics.

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surveyed along with the populations of neighboring communities, which also access the same hunting grounds, my estimates of km2 available per consumer are far from the sustainable estimates: hunting grounds from San Rafael had approximately 0.03 km2/consumer, Nuevo Lima vicinity had 0.05 km2/consumer, and the Alianza area had 0.02 km2/ consumer.

4.

Discussion

The results presented here provide information on where hunting is occurring, who is involved, what is being hunted, and how sustainable current hunting practices might be. Together these data have important implications for conservation strategies in the Cordillera Azul. In the Cordillera Azul hunters extract more animals and biomass from older forests (>20 y). The most hunted animals, even those typically associated with elds and young forests, were taken most often from older forests. However, old secondary forests (520 y), which cover the least area, are the greatest source of meat on a per unit area basis in terms of both biomass (kg/km2) and collection events per km2. This result provides support to Robinson and Bennetts (2004) hypothesis, as the relatively high extraction rates per unit area in fallows may be possible due to more available animal biomass in these secondary forests. In turn, these sites may demonstrate the best potential for sustainable harvesting. Although the quantities extracted differed greatly among participating households, nearly all (99%) hunted on at least one occasion during the course of the year. This differs from other ndings in Peru. For example, Naughton-Treves et al. (2003) found that only 49% of informants in the Tambopata region of southern Peru hunted. The variability in the quantities different households hunted reects an intracommunity heterogeneity in resource use found in many regions (Anderson and Ioris, 1992; Godoy et al., 1995; Takasaki et al., 2001). The fact that nearly all households hunted, indicates that any conservation and development efforts in the region will need to target this activity and its links to protein procurement and income generation. Any intervention must also account for the differences in resource dependence among households and the changes in resource extraction patterns over time as stakeholders adjust their livelihood strategies (Gavin and Anderson, 2006). As numerous studies in Amazonia and elsewhere have found (Robinson and Redford, 1991 and references therein; Vickers, 1991; Bodmer et al., 1994; Noss, 2000), mammals are the most hunted group. Redford and Robinson (1987) reviewed 24 studies of indigenous and non-indigenous hunting and found mammals to be the most hunted group followed by birds and reptiles. Among mammalian species, Bodmer (1995) found that hunters preferred ungulates and large-bodied rodents to primates or small rodents. The three most hunted animals in the Cordillera Azul were large rodents, and the fourth was an ungulate. The good news from a conservation perspective is that the most frequently hunted animals were those with high reproductive rates. Bodmer et al. (1997) suggest that animals with high intrinsic rates of natural increase, shorter longevities, and shorter generation times are less

prone to overharvesting. All of the top ten most hunted mammals t this description, and several of these species have been found to respond positively to harvesting through increased reproduction (Bodmer et al., 1997). Unfortunately, while the numbers of individuals extracted was low for the most vulnerable species, any hunting of these species, particularly those listed as threatened or endangered, will place them at additional peril. The willingness of participants to record the results of hunts for listed, and thus illegally obtained, species indicates the need for more enforcement, greater education to boost the awareness of local people regarding wildlife laws, and integrated conservation and development programming which provides incentives and alternatives to these species. Analyzing the sustainability of current hunting practices is a difcult task as it often relies on accurate information on local animal populations (including population densities, reproduction potential, and migration patterns) as well as on harvest rates. Making matters worse, the population dynamics of neotropical fauna can be highly variable in both time and space (Peres, 2000; Robinson and Bennett, 2004). Despite these difculties, comparison with regional consumption patterns and potential harvest rates can help pinpoint areas of possible conservation concern and highlight the need for further research. Households in the Cordillera Azul consumed meat of most species at rates well below regional averages. I also found that actual harvest rates were below the potential sustainable harvest estimates from the Robinson and Redford (1991) model. However, several caveats exist when applying numbers from the Robinson and Redford (1991) model. The sustainable harvest numbers calculated in that study used data taken from sites across the neotropics. Ideally, to make conclusions about sustainability in the Cordillera Azul, the model would need to be calibrated with data on local population dynamics for the species in question, but currently no research has been published from the Cordillera Azul for any of the species. In addition, the model assumes ideal conditions, with the population at 60% of carrying capacity, and production at the maximum levels recorded for this species, presumably under conditions where the species responds positively to lower densities. Because the species in the Cordillera Azul have been under hunting pressure for at least forty years and in many cases several hundred years, conditions are less than ideal and population densities in the region are undoubtedly low due to past use. This would mean the sustainable estimates used here are positively biased. Also, the model assumes a static population and does not account for sourcesink dynamics, which could allow local populations to sustain higher levels of harvesting thus indicating a negative bias in the Robinson and Redford estimates (Novaro et al., 2000). Finally, the Robinson and Redford model does not account for the ability of some species to positively respond to harvesting, which would greatly increase their sustainable harvest rates (Bodmer et al., 1997). The other three indicators of sustainability lead to conicting conclusions regarding the impact of hunting in the Cordillera Azul. Harvest rates in the Cordillera Azul (4884 kg/km2) were well below the sustainable estimate (152 kg/km2) of Robinson and Bennett (2000). This could indicate that either current hunting practices are not a major threat to wildlife in the

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region, or that past hunting pressure has drastically reduced wildlife densities. The hunting grounds in the Cordillera Azul currently only provide access to less than 0.1 km2 per consumer. Hill and Padwe (2000) estimate that sustainable hunting requires at least 5 km2 per consumer including both source and sink areas for hunted species. As in the Cordillera people in the Mbaracuya Reserve, where Hill Azul, the Ache and Padwe focused their research, rely on hunting, swidden agriculture, and domestic animals for subsistence; but it should be noted that the subtropical semi-deciduous forest of the Mbaracayu Reserve is a different forest type from the Cordillera Azul. Robinson and Bennett (2000) suggest that unless swidden agriculture systems provide substantial protein and assuming commercial hunting is limited, 1 km2/consumer could produce sustainable harvest rates. Although local people in the Cordillera Azul do not participate extensively in commercial hunting (only 11% of animals sold as compared with 86% in the Tamshiyacu-Tahuayo Communal Reserve; Bodmer et al., 1994), domestic animals provide some protein to all households surveyed in the Cordillera Azul (all participating households owned at least two chickens, 55% also owned a pig, and 15% had cattle). Therefore, it remains difcult to predict the exact sustainable human population density, but it can be assumed that a greater human population density can be sustained than if no domesticated animals were present. In reviewing studies of population densities in extractive reserves in Brazil, where people also hunt, keep domestic animals, and farm to a limited extent, Robinson and Bennett (2000) found that at sites with densities greater than one person/km2 wildlife populations were at low levels. Conditions vary across tropical forests in terms of biological carrying capacity, extent of commercial hunting, and degree of reliance on domestic crops and animals, making it difcult to apply particular sustainable human population density estimates across a large region. However, the current human densities in the study area in the Cordillera Azul appear to be well above any previous sustainability estimates. Another possibility is that the park adjacent to the communities surveyed serves as a source area for animals by resupplying the areas where hunting is intensive. If the source area supported a large enough population of key species, hunting might still be sustainable. Because data are not available on movement patterns of target species, I was unable to estimate the source area for the animals found in the hunting zones. However, based on 5 km2/consumer, in order for these hunting zones to be potentially sustainable, adjacent source areas must be between 4950 and 13 450 km2. Using the 1 km2/consumer estimate, the source areas would need to be between 990 and 2690 km2. Considering that the entire area of the national park covers 13 530 km2, and that the park is riddled with numerous barriers to animal movement including major rivers and mountains up to 2300 m, we can assume that insufcient source populations are available to match the current consumer demand. In addition, my estimates of local harvest rates are undoubtedly on the low end of the range, as they do not include hunting by non-locals. I noted non-local hunters, many with commercial interests in the region, on nearly a weekly basis during the study

period. Therefore, I propose that current low harvest rates are a relic of past overharvests which have limited current wildlife populations and do not reect sustainable use patterns. Data on local population densities and dynamics are required to conrm this hypothesis. Hunting appears not to be sustainable in the heavy use zones near human communities, but wildlife populations in more remote sections of the national park may remain relatively unaffected by current extraction activities. The ability of local communities to maintain current patterns of wildlife consumption may be limited unless hunters begin to penetrate deeper into the park, an outcome which would directly conict with conservation goals. My results have several important implications for conservation planning. Many conservation programs promote integrated conservation and development (ICDP) in buffer zones as a viable alternative to resource use inside protected areas (McShane and Wells, 2004). I found that while the secondary forests in the buffer zone of the Cordillera Azul National Park do provide more animal resources per km2, local inhabitants have a heavy reliance on older forests often inside the park, where they extract the majority of their wild meat. As communities grow, however, we can expect more older forests to be cleared for agriculture and a greater percentage of land to be covered in secondary forests. Because of the potential of these secondary forests to support higher biomasses of animals (Robinson and Bennett, 2004), I would predict that an increasing proportion of the total meat captured would come from secondary forests. The fact that most of the animal species hunted have high reproductive rates indicates the potential for many of the secondary forest animals to sustain high harvest rates. Unfortunately, the human population density in the communities surveyed near the Cordillera Azul National Park is well above sustainable levels, even when accounting for the large uninhabited protected area bordering the communities. Relatively low current levels of use appear to reect past overuse of wildlife and the current high population densities in the region. Even with ICDP projects focusing on establishing a source-sink dynamic with hunting in the buffer zone resupplied by protected animal populations in the park, the best information available seems to indicate that long-term sustainability of use near communities is unlikely. ICDP programs based on sustainable extraction in the region will need to be bolstered by projects which provide alternative protein options and income sources for local residents, and which seek to limit immigration into the area. Information on hunting, and resource use patterns in general, across an ecological succession can provide data critical to both local land use planning and to conservation and development programs. The results presented here indicate that while older forests remain a vital source of animal biomass for hunters, secondary forests provide more resources per unit area. As human populations, and in turn humandominated landscapes near protected areas grow, the role of secondary forests in resource management plans will only increase. However, the inclusion of sustainable harvesting into management regimes must be tempered by the realization that only low human population densities are likely to support signicant amounts of hunting.

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Acknowledgements
Financial support was provided by the Conservation, Food, and Health Foundation, Switzer Foundation, US Fulbright Programs, Garden Club of America, Conservation and Research Foundation, Ronald Bamford Endowment of the Department of Ecology and Evolutionary Biology, and the Graduate School and the Department of Ecology and Evolutionary Biology of the University of Connecticut. For logistical support and advice I thank the staff of the Centro de Conservacion, Investigacion, y Manejo de Recursos Naturales, Carlos Rengifo and Rafael Linares of the Centro de Desarrollo e Investigacion de la Selva Alta, Carlos Linares and Guillermo Vasquez of the Instituto de Investigaciones de la Amazonia Peruana. Gracias to Miguel Vasquez, Melita Ozambela, and Luis Garcia for their invaluable assistance in the eld; and to the communities who welcomed us and supported the research effort in countless ways. I thank G. Anderson, R. Chazdon, J. Silander, and Z. Cardon for support and advice during planning stages of the eldwork. R. Dunn and J. Solomon, as well as two anonymous reviewers, provided useful comments and suggestions on drafts of the manuscript.

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