Wildlife conservation in the cultural landscapes of the central Andes

LANDSCAPE AND URBAN PLANNING

ELSEVIER

Landscape

and Urban Planning

38 (1997) 137-147

Wildlife conservation in the cultural landscapes of the central Andes Kenneth R. Young Department

of Geography,

Uniuersiv

*

of Maryland Baltimore County, Baltimore, MD 21250, USA

Abstract The inhabited areas of the central Andean highlands are cultural landscapes, long dominated by land uses associated with subsistence agriculture. The wildlife remaining in these landscapes includes rare and endangered species, useful species, and those that can cause economic losses to local residents. Despite a general lack of relevant scientific investigations, it is important that wildlife concerns be included in rural development and nature reserve projects. This is best done by a three-prong approach that (1) improves the coverage, protection, and management of national parks and nature reserves; (2) integrates the control and management of wildlife into development projects focused on Andean communities; and (3) promotes species-specific programs that target endangered, useful or nuisance species of special concern. 0 1997 Elsevier Science B.V. Keywords:

Andes: Cultural

landscapes;

Subsistence

agriculture;

Endangered

1. Introduction The Andes Mountains were occupied by humans at the end of the last glacial period and the beginning of the Holocene (Rick, 1988; Bruhns, 1994). Buming, hunting, and gathering were undoubtedly the first anthropogenic impacts that began to alter conditions for wildlife. Several large mammals, including giant ground sloths and gomphotheriids, were extinct by the early Holocene, perhaps because of climatic and/or human influences (Martin and Klein, 1984; Webb, 1992). Over the next 10 millennia, the development of agriculture, ceramics and settlements followed (Burger, 1992; Bruhns, 19941, with increasingly important consequences for the characteristics and distribution of wildlife habitats, at least as can be

* Corresponding

author.

0169.2046/97/$17.00 0 1997 Elsevier Science B.V. All rights reserved. PZZ SO169-2046(97)00029-7

species; Nature reserves

surmised from palynologic and archaeological evidence. For example, Hansen and Rodbell (1995) recorded a dramatic increase at 5000-4000 years BP of pollen of weedy species probably associated with agriculture in a currently unoccupied area of the humid eastern cordillera of northern Peru; Johannessen and Hastorf (1990) found evidence in archaeological excavations of selective use of tree and shrub species as fuelwood beginning 2000 years ago in the Mantaro River valley of central Peru; and Wheeler et al. (1976) traced a 2000-year history of camelid hunting that then showed evidence of domestication and husbandry of llamas and alpacas during the next 4000 years. By the time of European contact in 1526 AD, most of the landscapes of the central Andean highlands had been modified to at least some extent by the land use practices of different social groups

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(Denevan, 1992a). This included often extensive terracing in steep areas, irrigation systems, and the drainage or other modification of wetlands (Donkin, 1979; Knapp, 1991; Treaty, 1994). After European contact, these landscapes were affected by dramatic declines in the sizes of native American populations, often on the order of 50-80% (Denevan, 1992b). This was followed by the slow rebuilding of rural populations (e.g., Cook, 1981) first over three centuries as colonies of Spain, and then during another century as the independent countries of Ecuador, Peru and Bolivia. The last three decades of the 20th century have seen some of the most rapid changes in the natural and cultural landscapes of the central Andes due to rapid urbanization in some areas, and out-migration in others as rural residents move to the cities. The consequences for wildlife of this long and complicated history have been inadequately investigated. Proposals for wildlife conservation in inhabited areas of developing countries need to be crafted in ways that harmonize with and build on the traditional land-use systems of rural populations. Although considerable effort has been expended by the governments and people of Ecuador, Peru and Bolivia to create national park and nature reserve systems, there has been virtually no holistic planning efforts for the wildlife remaining in the bulk of the region that falls outside of those protected areas. It is my goal here to describe important conservation concerns for the vertebrates found in the central Andean highlands that have long been occupied by people: the cultural landscapes of the high tropical Andes. I first review issues involving native biological diversity in these landscapes and then offer a three-part proposal for a planning process that is meant to be sensitive to both the demands of modem biological conservation practices and to the needs for integrated and culturally sensitive rural development.

2. Cultural

landscapes

of the central Andes

When physical and biological characteristics of landscapes are modified extensively by humans, it is common to refer to them as cultural landscapes, because social and economic decisions and processes are predominant in determining the extent and spatial

patterning of landscape features such as agricultural fields and rangeland, and of the remaining forests and wetlands. Because of the ancient and extensive impact of humans in Europe, Birks et al. (1988) dedicated a volume to studies concerning the development of cultural landscapes; Lepart and Debussche (1992) recently characterized the nature of human impact on landscapes, flora and fauna in the Mediterranean. It was Ellenberg (1979) who so prominently made the comparison between cultural landscapes in the Mediterranean region of southern Europe and similar patterns and processes in the tropical Andes. Much earlier, von Humbodlt and Bonpland (1955) recognized that natural ecosystems were distributed in the tropical Andes in relation to the environmental gradients of elevation and humidity. Troll (1968) extended this perspective to include the three-dimensional distribution of human land use superimposed on the natural heterogeneity of physical and biological patterns. Because some mountains reach well above permanent snowline, and because humidity regimes can vary from arid to perhumid (Gomez Molina and Little, 1981; Parsons, 1982) the spatial heterogeneity that characterizes both natural and cultural landscapes of the central Andes is remarkable and probably the most salient feature that affects both wildlife and human livelihood. At these latitudes, the tradewinds move most moisture from east to west. As a result, the western slopes of the central Andes are dry or seasonally dry (Valencia, 1992) except for northern Ecuador where a maritime influence creates conditions appropriate for the wet and rain forests of the Choc6 (Gentry, 1982). The eastern slopes were originally covered by near continuous humid-to-very-humid montane (1500-3500 m) and premontane (500-1500 m) forests, except in several areas where topographic rain shadows create drier conditions (Young, 1992). The highlands themselves are characterized by a north-south drying trend (Troll, 1968) with those in Ecuador much moister than those in southern Peru and Bolivia. In addition, the highlands include extensive rocky slopes; deep and narrow inter-Andean valleys; and large, rolling plateaus, especially from central Peru southward. The composition and structure of the original vegetation of much of these highland areas is relatively unknown, but climatic data and remnant vege-

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tation suggest to many that there was once much more extensive forest cover (e.g., Ellenberg, 1958; Laegaard, 1992; Kessler, 1995). For example, a tropical dry forest with a tall deciduous canopy and including numerous tree and shrub species can still be found in uninhabited valleys in southern Ecuador and northern Peru; most similar sites elsewhere are dominated by scrub or cacti. Many forest patches found at 2000-3500 m in Ecuador, Peru and eastern Bolivia appear to include tree species capable of covering much larger areas, perhaps at one time forming continuous forests. Even timberline, often at about 3500 m, is no absolute limit to Andean forests, given the adaptations of trees in the rosaceous genus, Polylepis, which are capable of forming closed forests to 4200 m or even higher (Simpson, 1979). Most elevations above 3500 m, but below snowline, are dominated by bunch grasses of the genera Calamagrostis, Festuca and Stipa. However, dominance at the highest elevations shifts to cushion plants, and wetlands are often are dominated by rushes and sedges. Traditional rural Andean communities chiefly consist of subsistence farmers who maintain a diverse set of crop species and varieties in fields located at different elevations. When this is not possible, barter or some other economic device is used to acquire needed resources from other ecological zones (Murra, 1972; Brush, 1976; Masuda et al., 1985). Construction materials often come from eucalypts (Dickinson, 19691, which were introduced more than a century ago from Australia and are now the most conspicuous trees in many central Andean landscapes. Fuelwood must be harvested from the forest remnants remaining in ravines or on hillsides. Households above timberline use peat, shrubs, or animal dung as fuel (Winterhalder et al., 1974; Reynel Rodriguez, 1988). Domesticated animals include imports from Eurasia, such as chicken, pigs, sheep, cattle and horses, and also indigenous species including llamas, alpacas and guinea pigs. Large areas of scrub and grassland are used as rangeland; usually the only preparation for grazing is occasional burning. Modernization since the 1960s has often meant declining rural populations (Donoso de Baixeras, 1992; Southgate and Whitaker, 19941, especially in parts of the highlands where such needs as good

Table 1 Selected protected

areas in the central Andean highland?

Name and location

Year established

Ecuador Cotacachi Cayapas Ecological Cotopaxi National Park Sangay National Park Podocarpus National Park

Area (km*)

1968 1975 1979 1982

2040 334 5177 1463

Peru Pampas Galeras National Reserve Manu National Park Huascaran National Park Salinas y Aguada Blanca National Reserve Rio Abiseo National Park Yanachaga-Chemillen National Park

1967 1973 1975 1979 1983 1986

65 15 328 3400 3669 2745 1220

Bolivia Ulla Ulla National Reserve Amboro National Park Eduardo Avaroa National Reserve Tariquia National Reserve

1972 1973 1973 1989

2500 6376 7148 2469

aPulido,

199 1; Marconi,

Reserve

1992; Mena Vasconez,

1995.

communications, roads, electricity, education and health care are limited or lacking. Some large highland cities, such as Quito, Cajamarca, Cusco and La Paz, are growing rapidly, with the conversion of nearby hillsides and valleys into suburbs or communities producing products for the new urban markets. The last three decades have also seen the development of a series of national parks and nature reserves in the central Andean highlands (Table 1). In most cases, these were tracts of land owned or expropriated by the national governments. In some cases, such as Huascaran National Park in Peru, the protected areas include lands still used by rural people for the grazing of livestock.

3. Biological diversity

of the central Andes

Indirect and partial evidence suggests that the biological diversity of the central Andes is of global importance. Henderson et al. (1991) found that the floristic diversity of the northern Andes was as great or larger than that found in the entire Amazon basin. The estimates of plant species richness for Ecuador (25 000; Jorgensen et al., 1995), Peru (17 000; Brako

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and Zarucchi, 19931, and Bolivia (18 000; Solomon, 1989) suggest that the central Andes also constitute a significant portion of neotropical plant diversity, especially considering the high endemism found in the arid and semi-arid western Andes and coastal plain (Leon et al., 19961, and the humid montane and premontane forests of the eastern Andes (Leon et al., 1992; Young, 1991, 1996). Given these facts, the international attention paid to the Amazon lowlands appears to be disproportionate. Mares (1992) made this very point using data on the distribution and diversity of mammals in South America. He found that the majority of species were associated with dry or montane habitats; this was especially true for species with restricted distributions. He pointed out that conservation efforts focusing on the Amazon lowlands were thus disregarding important components of tropical diversity. Others that document important diversity in the highlands include Dinerstein et al. (1995) in general, and Fjelds% and Krabbe (1990) for birds, Duellman (1993) and Rodrfguez et al. (1993) for amphibians, and Ortega (1992) for fishes. To this perspective, I would add that the biological diversity found in the cultural landscapes of the central Andes is also critically important. The ancestors and wild relatives of Andean crops, such as the potatoes, are often associated with highland scrub, montane, or tropical alpine vegetation (e.g., Iltis, 1988; National Research Council, 1989; Ochoa, 1990). The same is true for the domesticated camelids (Wheeler, 1995) and guinea pigs (Morales, 1995). Many other species are useful or important economically, or are of concern because of their rarity. 3.1. Useful wildlife species The occasional taking of game is common in traditional communities of the central Andes. Often one or several men in a given community specialize in hunting white-tailed deer (Odocoileus uirginianus), or, less frequently, other game such as ducks (Anatidae), doves (Columbidae), viscachas (Lagidium spp. ), or rabbits (Silvilagus brasiliensis). A substantial limitation is the cost and availability of ammunition and weapons. Although no published study quantifies the game taken, presumably more isolated communities are more dependent on game.

Native fishes are small in size in the highlands. They were once harvested by net or by hand; the fisheries of Lake Titicaca were especially important at one time (Villwock, 1994). The introduction of exotic fishes, particularly rainbow trout (Oncorhynchus mykiss), has changed these practices. Most trout are captured by hook and line; others are raised in tanks or in netted enclosures set in lakes. Domestication processes have produced guinea pigs (Cauia porcellus), alpacas (Lama paces), and llamas (Lama glama) in the central Andes. Because of the unexplored state of the genetic diversity found among these domesticates, it is likely that the most important value currently is that of the potential worth of that genetic material, which could be used to improve and develop breeds (Hodges, 1990). The wild species are present, although often poorly protected or managed, in high elevation habitats: wild guinea pigs (Cauia spp.), vicufias (Vicugna uicugna, see Wheeler, 1995), and guanacos (Lama guanicoe). Only the vicuiias have received special attention, and that after populations had crashed catastrophically earlier this century. Today their wool is collected yearly by a number of Andean communities in Peru (INRENA, 1994). Finally, many national and foreign tourists are willing to pay to see wildlife and wildlife habitats. This ecotourism represents an economic and cultural interchange that has been inadequately documented or examined for the Andean highlands, although it is, by all appearances, on the increase. 3.2. Nuisance

wildlife species

Subsistence farmers live on the margin and so are especially susceptible to natural calamities such as drought or hail. Because they typically have only small herds or flocks of domesticated animals, loss to predators can represent a tremendous economic cost. Common predators for horses, cattle and sheep are puma (Felis concolor) and Andean fox (Pseudalopex culpaeus). Weasels (Mustela frenata), opossums (Didelphis sp.), and raptors (Accipitridae, Falconidae) can be destructive to poultry, which are typically allowed to roam freely near dwellings. A host of other species are ready to damage crops if vigilance is not maintained. Granivorous birds (e.g., Emberizinae) and parrots (Psittacidae) can de-

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stroy crops such as maize and wheat; the granivorous birds also steal recently planted seed. Skunks (Conepatus spp.) dig up and consume potatoes. It is a common sight to see young boys deployed with slingshots, attempting to protect crops during their most susceptible stages. Although nuisance wildlife may not be numerically or quantitatively important in many communities, there is little doubt that they are viewed with considerable concern by local people. The loss of one horse to puma attack can set back the plans of a small land holder for several years. Overnight, one skunk can significantly affect the potato harvest of a family. In addition, some wildlife species can serve as health risks, typically due to diseases and parasites shared with related domesticated species, but in some cases, these can also affect humans directly. An example of the latter is rabies, while the former includes anthrax and distemper. 3.3. Threatened

wildlife species

Many of the wildlife species most at risk are no longer found among the cultural landscapes of the central Andes. Some, such as the Andean bear (Tremarctos ornatus), were once fairly widespread, but have been severely affected by hunting (Peyton, 1980). Others, such as the yellow-tailed woolly monkey (Lagothrix flauicauda; Leo, 1984) and the mountain tapir (Tupirus pinchaque), are specialists of the wet montane forests that are least suitable for human occupation. However, in some communities, there is an active deforestation front associated with the expansion of the agricultural frontier. Here, habitat loss and hunting can affect and possibly threaten remaining populations of species that are of concern to conservationists. Forest-dwelling species at risk in this manner include large birds, such as the cracids (Crux, Ortalk, Penelope), toucanets ( Aulacorhynchus), and trogons (Pharomarcus, Trogon). Smaller birds mentioned by Kattan (1992) for the Colombian Andes include woodcreepers (Dendrocolaptinae), ovenbirds (Funariinae), antbirds (Thamnophilidae), contingas (Cotinginae), and parrots (Psittacidae). Hunting is the primary cause of population declines of the wild camelids, and several high-elevation deer species (Hippocamelus antisensis, Mazama

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chunyi, Pudu mephistophiles). In most cases, these animals are now so uncommon that few people deliberately hunt them, although they might be taken opportunistically. The exception is the vicufia, which is searched for and poached in Peru and Bolivia. Some of the most threatened vertebrates in the central Andes come from groups that receive little attention: fish and amphibians. Native fishes and frogs that have aquatic tadpole stages, can be systematically eliminated from lakes, ponds, and streams by the voracious feeding of trout (e.g., Flecker and Townsend, 1994; Villwock, 1994) which have seemingly been introduced to every body of water in the Andean highlands.

4. Conservation

strategies

Here, I propose a three-part approach to wildlife conservation. To date, most national and intemational attention and efforts have been focused on the establishment and reinforcement of national parks and other protected areas. At best, such protection can only cover a relatively small percentage of the region. What is more, however, is that there has always been a bias toward including what are viewed as ‘pristine’ habitats in these reserves, leaving unrepresented many highland sites influenced by humans. For that matter, good farmland has never been included deliberately within a reserve, for obvious reasons. This means that protected areas do not offer a complete solution to the conservation needs of wildlife, even if they were to function optimally. Instead, there should also be wildlife programs that are oriented specifically to the needs of rural residents. These should be community-based approaches tied to general development goals and implemented locally, or perhaps in reference to a watershed or some political entity such as a district or province. Yet, while these approaches cover protection of wildlife habitat and serve human needs, also required are projects directed at the particular species of most concern. For example, endangered species cannot be protected solely on the basis of their presence in one or more nature reserves. National regulations and laws protect these species regardless of their location. Other wildlife species might serve as sources of

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game if managed sustainably, or might need to be controlled if they cause losses to crops or livestock, or else represent a health risk. Below I elaborate on the different strategies required to cover each of these three general approaches. 4.1. Protected

areas

Protected areas that serve wildlife conservation needs are of several types in the central Andean countries, including some that permit resource extraction, such as hunting, and those designed for tourism and recreation (McNeely et al., 1994). As a generalization, those areas that are most important for wildlife are those that are managed for strict protection, such as the national parks and sanctuaries. Ironically, the strict protection required by law has often been taken to mean passive rather than active management. That is, the limits of the park or reserve are policed, but ‘natural’ processes are allowed to function within. This is fine for large expanses of pristine habitat, but can be counterproductive for small reserves or those that include degraded habitat or impoverished populations. More appropriate in those cases is an activist agenda that would include ecosystem restoration and the possibility of species reintroductions and population augmentation. Protection of reserve boundaries requires interactive management since the land-use practices of neighboring human communities often have implications for habitats or species within the reserve. For example, fires started outside can rarely be stopped before entering the reserve. Some diseases and parasites found among domesticated animals can infect wild populations of native species found within reserves. However, the reverse might also be the case: processes that originate within the reserve might damage or limit traditional agriculture or range management in adjacent areas. Wild animal populations can function as disease reservoirs; sometimes natural disturbances in protected areas can cause risks to infrastructure, dwellings, or human safety outside. Good management requires knowledge of and sensitivity to these issues. This, in turn, requires good relations with the reserve’s neighbors. This is best done with programs that address

mutual concerns and provide mutual benefits (Hough, 1988; Brandon and Wells, 1992). This does not always mean that such programs need to be carried out in the buffer areas of the park or reserve: it is only important that there exists a public relationship between a rural development project and the existence of the reserve. Southgate and Clark (1993) pointed out that many ‘park-people’ programs were mistakenly directing development to the very places where conditions were least suitable for human habitation and most critical for natural ecosystems. A simple example: it would be better to install a potable water system for a small town in return for support for a nearby reserve, than to promote alpaca husbandry in the grasslands adjacent to the reserve. The goal should be to direct land use and impacts away from the reserve (Young, 1993), and to make such displacement a permanent part of interactive park boundary management. A third strategy is to search for ways to provide spatial linkages between and among the different protected areas (e.g., Noss and Harris, 1986; Yerena, 1994). This regional approach limits the effects of insularization of the reserves (Saunders et al., 1991; Shafer, 1995) and promotes genetic interchange. Habitat corridors can be designed to be horizontal, that is, connecting similar habitats in different places; or vertical, connecting habitats across a range of different elevations or ecological zones. 4.2. Community-based

approaches

Most central Andean communities are not located near a protected area, so wildlife conservation initiatives need to originate with development projects directed at all or a part of the community. Potentially, any project that involves renewable natural resources could include a wildlife component. At the least, such projects could incorporate an awareness of implications for wildlife. For example, the forest cover needed to protect the sources of water used for potable water systems will also serve as wildlife habitat. If done responsibly, and if the forest or tree plantation is sufficiently large, this could also be a site where firewood and game could be extracted sustainably. If this is developed as a community resource, then traditional or other measures are needed so the extracted products

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are shared or divided without leading to overexploitation (Goodland et al., 1989; Feeny et al., 19901. Often the goals of reforestation projects are quite limited or poorly evaluated. This partly explains why so many projects have been carried out in the Andes using exotic tree species. Planting such species for specific purposes is appropriate. However, reforestation goals that include reestablishment of original forest cover require the planting of native species. This is more difficult technically because of the paucity of information about the silvics of native Andean species. It is more difficult practically because of the need to gather ripe seeds and to distribute them in a timely fashion to participating tree nurseries. But the use of native species would serve ecological restoration goals, while also enriching the soil, maintaining native forest understory plants, and encouraging associated wildlife. Wetlands are often drained for agriculture and near urban areas. This can be shortsighted for hydrological reasons, because such changes increase the risk of flooding in times of abundant rainfall, and decrease stream discharge in times of drought. Maintaining wetlands will also provide habitat for native frogs and fishes, in addition to migrating waterfowl. The use of range is extensive, but not often intensive in the central Andes. This fits with the available technology and resources because little infrastructure or investment is required to graze cattle or sheep on unimproved rangeland. A disadvantage for wildlife is that such land-use systems leave the livestock exposed to predators and susceptible to disease and malnutrition over large tracts of land. This invites inevitable retaliation, often by poison or gunshot, when predators such as Andean fox or puma kill or scavenge domesticated animals. Solutions include intensifying range use by establishing improved pastures, by planting forage species, and by closer monitoring of livestock. Virtually unstudied is the practicality and productivity of mixed systems that could include both wild and domesticated herbivores, such as deer and cattle or vicuiia and sheep. Indeed, the vicui?a is a special case because of the great value of its fine wool, and the historical precedence of herding and shearing wild populations once a year. With the proper technical and legal backing, traditional Andean communities have successfully

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and sustainably used vicufia in South America (Rabinovich et al., 1991). The biggest threat is poaching by outsiders. While the solution is increased enforcement, it also would be helpful if demand could be undercut by an international refusal to purchase vicuiia wool that comes from dubious sources. Additional community-based approaches that could include wildlife also exist among health, education and infrastructure projects. Certainly, for example, road construction and maintenance projects should consider implications for wildlife (Young, 1994). The professionals involved in the design and implementation of development projects need to ask themselves when and if wildlife concerns can be addressed, at least as a secondary concern or goal. 4.3. Species programs Rare and threatened species are typically identified by the respective national governments using, as criteria, the sizes and distributions of their populations in the country, in addition to their presence on international lists, such as those of CITES. Unfortunately, only a few high-profile species have had recovery plans developed and implemented. Such planning needs to consider how well the species is protected in the existing park and reserve system, whether current legislation and enforcement is adequate and appropriate, and to what extent habitats not found in protected areas are critical for the survival and stabilization of populations of the species of concern. Additional strategies might include habitat-focused conservation programs for wetlands, and other sparse or disappearing ecosystems, incentives or fines to protect certain species, and reintroductions (e.g., Barrera-Rodrfguez and Feliciano-Caceres, 1994). Integrated planning for these species will need to be regional, national, and/or international in scope and will necessarily include both protected and nonprotected areas. In some cases, an endemic species can make a compelling symbol for local or regional development projects that include natural resources. Some rare species can also be nuisances. Although implementation might be difficult, fairness requires the development of either a compensation program when such species cause losses or damages

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to crops or livestock, or a relocation or extermination program for individual animals that cause losses. Species in this category might include the Andean bear and the raptors. Economic loss should be shared by the country or international community rather than being borne exclusively by the farmer or herder that happens to live near the place where the rare species is being maintained. Other programs directed at particular species may have exploitation or control as a primary goal. For example, white-tailed deer are found the length of the tropical Andes, but are seldom managed despite considerable expertise elsewhere (Hudson et al., 1989; Vaughan and Rodgriguez, 1991). Strategies to develop sustainable hunting regimes require information on habitat needs and reproductive rates. This in turn requires that more range and forestry programs in the universities include wildlife management in their curricula.

5. Implementation Interests in conservation and development motivate many social actors, including governmental organizations (NGOS), agencies, nongovernmental and unorganized groups (Murphree, 1994; Young and Leon, 1995). Planning and financing is often at regional, national, or even international levels, while the most efficient applications are generally at the grassroots level, in local communities or associated with particular protected areas. Because of the low profile of wildlife concerns among rural development programs, it would be helpful if more successful case examples were to be analyzed and promoted (e.g., Homewood and Rodgers, 1991; Western and Wright, 1994). Difficulties in implementation include the need to act for now with little information and few active professionals. There are few relevant baseline studies, including faunistic inventories, and funding for such research is difficult to obtain. There is a general lack of institutional structure and capacity to carry out either research or the development projects themselves. However, given the universal appeal of some kinds of wildlife and the demonstrated importance of Andean wildlife in terms of neotropical biological

diversity and in relation to rural livelihoods, it is important that wildlife concerns be given their due. Wildlife can be a primary or complimentary focus to many kinds of development projects designed for the Andean highlands. The institution building required in the universities and other organizations would likely have many additional benefits.

6. Conclusions By the time of European contact with the people of the central Andes, there existed a matrix of human-dominated landscapes imposed upon a mountainous region, diverse naturally with extremes in precipitation from arid to perhumid and in elevation from the Andean foothills to tropical alpine grasslands and permanent snow. In the last four centuries, these cultural landscapes have continued to change as a function of the size of rural populations and the type and intensity of land use. The last three decades have also brought rapid urbanization and the establishment of national park and nature reserve systems. Conservation of wildlife in these cultural landscapes requires a better understanding of the consequences of a long and complicated history. In the meantime, however, research might be productively focused on answering the following questions: How have wildlife habitats been modified? Did this modification result in local extinctions? How are the remaining species adapted to conditions found among the cultural landscapes? To what degree do the parks and nature reserves protect the species of concern? How are rural human populations affected by wildlife species? Conservation strategies should be targeted at (1) national parks, nature reserves, and surrounding human communities; (2) rural development projects that include manipulation or restoration of wildlife habitats and populations; and (3) species of concern due to their rarity, usefulness, or destructiveness. Currently, few nongovernmental organizations or government institutions are constituted such that wildlife concerns can be addressed simultaneously on these three fronts. At the very least, these social actors should be aware of the possibilities and importance of integrated planning for wildlife conservation in the central Andes.

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Acknowledgements For comments and information, I thank Asunci6n Cano, Rogelio Cueva, Blanca Le6n, Abel Salirrosas, Ctsar Salirrosas, and Carmen Ulloa Ulloa. For financial support I am grateful to the John D. and Catherine T. MacArthur Foundation, Pew Charitable Trusts, and the National Science Foundation @ES-87 13237 and EAR-9422423).

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