Huemul Hippocamelus Bisulcus in Central Chile Anthony Povilitis Life Net, HCR Route 3, Box 3845, Willcox, A285643, USA

BIOLOGICAL CONSERVATION

ELSEVIER Biological Conservation 86 (1998) 97-104

Characteristics and conservation of a fragmented population of huemul Hippocamelus bisulcus in central Chile Anthony Povilitis Life Net, HCR Route 3, Box 3845, Willcox, A285643, USA

Received 7 September 1997; received in revised form 9 September 1997; accepted 11 September 1997

Abstract The future of the endangered huemul deer in central Chile rests with efforts to recover a single surviving population located in the 3000 km 2 Nevados de Chill/m Motmtains-Polcura Valley area. Field survey data for 1975-1997 were used to assess size, spatial distribution, and trend of the population. In 1997, it was estimated at 60 individuals at 12 sites separated by an average cross-valley distance of 8.0 kin. Data suggested a 58% population decline over two decades. Huemul had disappeared from at least five sites, and evidence for recolonization of vacant habitat was weak. A habitat evaluation and assessment of threats to the huemul, such as poaching and population fragmentation, help provide a basis for conservation action. Proposed measures include protection of two habitat core areas each consisting of 8-11 primary sites, conservation of connecting habitat between sites and core areas, land management practices limiting livestock, logging, and development impacts, and, if necessary, population augmentation. For long- term conservation of the huemul, renewed connectivity between central Chilean and Patagonia populations is recommended. © 1998 Elsevier Science Ltd. All rights reserved. Keywords: Hippocamelus bisulcus; Endangered deer; Central Chile; Population fragmentation; Reserve design and management

1. Introduction Requiring broad areas of habitat, the huemul is con- sidered an 'umbrella species' (Noss and Cooperrider, The huemul is one of four species of large South 1994; Hunter, 1996) whose conservation lends protection American deer, all of which are at risk of extinction to other Andean wildlife (Parques Nacionales de (IUCN, 1990; US Fish and Wildlife Service, 1994). Argentina, 1992). It is also a high profile 'flagship species' Because of poaching, habitat loss from forest clearing symbolizing the imperiled natural heritage of the south and human settlement, and livestock-related diseases, Andes and depicted with the threatened Andean condor huemul have vanished from the northern part of their Vultur gryphus on the Chilean coat-of-arms. historic range, except for a single population in central Huemul prefer steep, broken terrain with forest and Chile (Povilitis, 1983a; Parques Nacionales de Argen- shrub cover, and require access to lower elevations tina, 1992; Smith-Flueck and Flueck, 1995) (Fig. 1). where snow accumulation is light (Povilitis, 1979; Most huemul occur in Patagonia where they number Frid, 1994). Such topography characterizes the south ca. 1500 individuals. Andes, except for areas of dense lowland forest, open Huemul in central Chile were the focus of a major grasslands, and barren volcanic or glaciated terrain. The conservation campaign involving habitat acquisition, huemul'sselection of mountainous terrain appears to protection by wildlife guards, public education, and the have shaped its unique social organization, featuring creation of a regional recovery team (Aldridge, 1988; small group size and prolonged male-female pairing Espinosa, 1996). Geographically separate from other (Povilitis, 1983b, 1985; Montecinos, 1995). huemul, these animals may comprise a peripheral Because of human impacts, huemul populations population of special genetic and evolutionary impor- today are smaller, more subdivided, and more isolated tance (Lesica and Allendorf, 1995) in a distinct region of from one another than those of the past (Miller, 1980; Chile. Central Chile is of conservation priority as the Drouilly, 1983; Povilitis, 1983a). Consevationists have nation's center of biological diversity, human density, and responded by creating protected areas for small groups economic growth (Hoffman, 1993; Armesto et al., 1994). of huemul and by posting wildlife guards to prevent

0006-3207/98/$19.00 © 1998 Elsevier Science Ltd. All rights reserved PII: S0006-3207(97)00161-4 98 A. Povilitis/Biological Conservation 86 (1998) 97-104 poaching or disturbance of huemul over broader areas. south (Fig. 2). The nearest known huemul population This approach may work if dispersing individuals occa- occurs 350kin to the south in Nahuel Huapi National sionally move between existing huemul groups and park, Argentina (Serret, 1992). where local extinction, if it occurs, is balanced by reco- Between 1975 and 1997, 85 field surveys were con- Ionization. Conversely, where huemul groups become ducted in late spring or summer to evaluate huemul entirely disjunct and dispersal does not occur, efforts to population distribution, trends, and size in the Nevados protect limited habitat areas will not be adequate, and area. Surveys covered 30 distinct areas of mountain more extensive habitat protection and restoration will terrain which were identified by topography and vege- be needed. tation (Povilitis, 1979) as potential huemul sites. Thir- Population characteristics, threats, and habitat of teen of 19 areas where huemul occurred were huemul in central Chile are evaluated by addressing the periodically re-examined. The work was conducted by issue of population viability. Based on the analysis, teams of 2-20 people trained to detect huemul, and measures for huemul recovery are proposed. normally lasted from 2-7 days depending on team size and terrain. Teams systematically surveyed along ridges and mountain tops, descending for 100-300 m to exam- 2. Study area and methods ine preferred huemul locations: ravines, rocky areas and cliffs, and associated Nothofagus spp. forest and shrub Central Chile's huemul population is located in the (Povilitis, 1979). Nevados de Chill/m Mountains-Polcura Valley area Minimum counts of huemul were derived from sight- (hereafter referred to as the Nevados area; Fig. l) The ings and indirectly through track analysis. The typically area covers 3000 km 2 bordered by Chile's central valley open volcanic soils of ridges and mountain tops in the on the west, Argentine steppe highlands on the east, and Nevados area aided in tracking huemul. Distinct track the Rio iquble and Rio Laja valleys to the north and sets and fresh discontinuous track sets separated by more than 1 km were recorded as separate animals.

o Huemul groups were defined as sets of individuals in 75° 7O I I spatial proximity and presumed to be interactive. Hue- i • Santiago mul groups typically consisted of an adult female and t dominant adult male often with associated juveniles / 35 °_ and/or subdominant males (Povilitis, 1979, 1983b, 1985). i J 70=30' 71 ° 15" Area I I

Chile 36o30 , -- Argentina ~'~4)6 ~ 40°~ ~6/e C

"13 C ¢0 N -- ~ .,,,,/~ ~.Gei"trudis~ 36° 45~ _ /,- tO t i s <( 4s°... oo : I o Chill~n"~ o@ 400 km I I I O- Ig" Ren~ / N i

Former range 37°00 S _ (- - ~/J t Ponce"') C~koados ~.~ \ xt 10 km Infiernillo" / ~ 50° __ C.-" L,a~lnas /c.~nposa s ,, / ( Atrav~d~ 0..~ (/~___henque

'Tierra 37° 15" -- vi,,~0~.~ ~ / CoJor~do) ..,, f/ I t

I o 170o 75

Fig. 1. Geographic range of huemul in the south Andes .(based on Fig. 2. Distribution of primary huemul sites in the Nevados de Osgood, 1943; Cabrera and Yepes, 1960; Serret, 1992; Diaz, 1993; Chill~.n area of Chile. Sites believed occupied by huemui in 1997 are Frid, 1994). hatched. A. Povilitis/Biological Conservation 86 (1998) 97-104 99

Based on previous research on habitat selection by ! 6 huemul in the Nevados area (Povilitis, 1979), primary habitat sites (Fig. 2) were defined as predominantly O O north-facing, generally 30--40 degrees in slope, and attaining at least 1750m elevation. Upper south-facing ); 18 ~ • Huemul group slopes within 500m of such terrain and major ridges 0 i extending southward were included as potential summer } ~ Multiple groups habitat. To qualify as primary, sites needed to include at '~, 0 Vacated site least 100 ha of north-facing terrain below 1500 m elevation, 8~•'* ,~ Shortest cross-vaJley where winter snow accumulation would be light. Habitat ,q route between groups 0 sites were delineated on 1:50,000-scale topographic maps. 6 ...... Shortest highland Habitat sites were assessed by size, quality, and spa- 4 rb~ route between groups tial configuration. Habitat size was ranked according to total map area and potential winter range (north-facing slopes below 1500 m elevation). Sites of > 2000 ha were assigned a value of 1; those > 1200-2000ha, a 2; and those <1200ha, a 3. Similarly, for winter range, sites Fig. 3. Dispersion of huemul in the Nevados de ChillOn area, Chile, with > 500 ha were assigned a 1; those > 300-500 ha, a based on recent field surveys. Numbers indicate shortest cross-valley 2; and sites with <300ha, a 3. Total site size and winter distances (kin) between nearest groups. range scores were averaged to assign overall size rank as follows: score of 1.0=rank 1; 1.5=rank 2; 2.0=rank 3; Of 13 sites surveyed at least twice, the total minimum 2.5 =rank 4; and 3.0=rank 5. count from the latest surveys was 58% less than earliest Habitat quality was ordinally ranked in which surveys (23 versus 55) (Table 2). Minimum counts A = excellent; B = very good; and C = good, on the basis dropped at 10 sites, were essentially unchanged at 1, of combined habitat criteria: degree of rockiness; abun- declined and partially recovered at 1, and increased at 1 dance of cliffy sites and ravines; extent of shrub and (p < 0.01, Wilcoxon signed rank test). forest cover; and cover type heterogeneity (Povilitis, The number of huemul for primary sites was esti- 1979). Rankings were based on visual assessments, and mated at 52 individuals, based on the minimum count on sites descriptions by Lopez (1994). of 40 huemul for the latest surveys (all sites, Table 2) plus 31%--an average difference between minimum counts and probable number derived from repetitive 3. Results and discussion surveys and visual identification of huemul at three sites (Povilitis, 1979). Given that some additional animals 3.1. Distribution of groups were likely present as dispersers in secondary habitat, I suggest an overall population of about 60 huemul for Huemul groups or group clusters were scattered with the Nevados area in 1997. shortest distances between them averaging 8.0 km across valleys and 11.4 km along ridge and mountain routes 3.3. Habitat evaluation (excluding the outlying Palo group for which no direct highland connection exists; Fig. 3). Although cross- The Diguillln and lqiblinto sites are the largest, best valley distances are shorter, huemul may seldom enter quality sites for huemul in the Nevados area (Table 3). valleys as only one Nevados huemul was recorded in a Further, they help bridge north and south site clusters, valley (Victor Sfinchez, provincial chief, CONAF, pers. comm). During 437 h of observing huemul in Patagonia Table 1 (Povilitis, 1985), only once were huemul observed Vacated huemul sites at the Nevados de Chiil~in area, Chile based on advancing to a valley bottom, from which they shortly periodic field surveys retreated to mountainous terrain. Site Years present Years absent

3.2. Population trend and size Chillhn 1975, 1976, 1980, 1983, 1987 1988, 1990, 1992, 1995, 1997 Jos6 1975, 1976, 1989, 1991 1992, 1994 Huemul in the Nevados area have disappeared from Gertrudis 1975, 1976, 1980, 1986, 1993 1996, 1997 at least five sites (Table 1). The best documented case Capados 1982, 1983, 1990 1993, 1997 involves the Chill/m site which was occupied by huemul Blanquillo 1983, 1990 1994, 1997 for at least 13 yr prior to 1988 when only old tracks and Infiernillo 1994 1995a pellets could be found. By 1990, huemul were clearly Gato 1994 1997a absent from the site. a Multiple surveys are required to confirm huemul absence. 100 A. Povilitis/BiologicalConservation 86 (1998) 97-104

Table 2 Table 3 Minimum counts of huemui recorded from occupied huemul sites, Evaluation of primary habitat sites for huemul in the Nevados de Nevados de Chili/m area, Chile 1975-1997. Figures separated by a Chillfin area, Chile comma represent results from two different surveys Overall rank Size and Cluster Corridor c Huemul Site 1975-1976 1980-1989 1990-1993 1994--1997 General quality rank a component b presence a trend a Highest Cato N. - - - 2 - Diguillin 1A xx X X Cato S. - - 4 - - /qiblinto IA xx X X Damas 1 0 4 - + 3 Juan 2A x X C_rertrudis 3b 3,3 3 0,0 -3 Damas 3A xx X Cabras 6 b 3,1 2 3 -3 Infiernillo 3A xx lqiblinto 9 - 6 4 -5 Cato N. 2B xx Gato - - - 3,0 -3 Cato S. 3B xx Jos6 3b 4 0,0 - -3 Pelado 3B x Chilhln 3b 3,1 0 0,0 -3 Catalinas 4B xx Renegado - 2,3 2 2,2 0 Atravesado 5A x Diguillln - 8,6 10 6 -2 Chillfin 4B Infiernillo - - - 2,0 -2 Renegado 1C Catalinas - 12 8 4 - 8 Jos~ 4C Capados 1,1 1,0 0 - 1 High Blanquiilo - 2 2 0,0 -2 Ponce 3C xx Juan - - - 6 - Blanquillo 4C xx Pelado - - - 2 - Capados 5C xx Atravesado - - - 2 - Alico 5C xx Palo - - - 1 - Gertrudis 4B x Cabras 4B x a Figures are the difference between most recent and earliest counts. Gato 4B x b Figures represent median counts of multiple surveys: Gertrudis n = 3; Cabras n = 7; Jos6 n = 5; Chill~n n = 11. Moderate Colorado 1? x Palo 3C x namely groups of primary habitat sites in relative Mariposas 3B Chenque 4B proximity (Fig. 2). Eight other sites were 'highest' Toro 5C x ranked as key duster components. The Chill~n, Rene- Villagrfin 3C gado, and Jos6 sites were also top ranked because of their position between north and south site clusters. a Size rank: 1 =largest, 5=smallest; habitat rank: A=excellent, B = very good, C = good. Seven 'high' ranked sites fill out the site dusters: b x = Cluster of 3 sites, xx = cluster of 4 or more sites. Alico, Gertrudis, Cabras, and Gato to the north, and c Linking north and south site clusters. the tightly-grouped Ponce, Blanquillo, and Capados d Based on the most recent survey at each site. sites in the south. Remaining primary huemul sites in the Nevados area ranked 'moderate' as they are impor- tant for population recovery particularly along the Rio (Frid, 1994; Smith-Flueck and Flueck, 1995). Based on Polcura (Fig. 2). track records, the density of puma in the Nevados area appeared low. Culpeo fox were common, probably 3.4. Threats because of an abundance of introduced hare Lepus europaeus as prey. 3.4.1. Predation Poaching of Nevados huemul has not been docu- 3.4.2. Disease mented since the 1970s when two huemul where cap- Huemul show vulnerability to disease and parasitism. tured as a curios by local people. Nevertheless, the Three captive huemul died apparently from coccidiosis intense alarm response to people observed in the field acquired from domestic sheep (Texera, 1974). Eight suggested harassment; huemul not normally pursued by others were believed to succumb from bladderworm people show a remarkable lack of wariness (Povilitis, Cysticerus tenuicollis and liver fluke Fasciola hepatica 1979; MacNamara, 1982; Monteeinos, 1995). Harass- infections (M. Durand, private zoo owner, Santiago, ment is likely to involve domestic dogs which accompany pers. comm.) These animals all originated from Pata- most livestock herders and backcountry travelers. Dogs gonia (MacNamara, 1982). In the Nevados area, wild have been observed chasing huemul from lower moun- huemul and free-ranging domestic cattle and goats all tain slopes (Povilitis, 1983a), and they could kill fawns. had light infections of coccidia and strongles, while Natural predators include puma Felis concolor and tapeworm Moniezia sp. occurred in both huemul and potentially culpeo fox Pseudalopex culpaeus on fawns cattle (Povilitis, 1979). A Nevados fawn that died in A. Povilitis/Biological Conservation 86 (1998) 97-104 101 captivit) was severely infested with goat louse Bovicola 3.5. Population integrity caprae. For population viability, small groups of animals 3.4.3. Habitat degradation require at least occasional interchange of individuals, Land settlement, recreational development, and and recolonization of unoccupied sites. Gilpin (1987), energy pipeline construction have dramatically Hanski and Gilpin (1991) and Harrison (1991) discuss increased human presence in the Nevados area since the these requirements in a metapopulation context. 1970s. Logging and road construction have increased, Interchange among huemul groups in the Nevados reducing protective cover and allowing human access to area is in serious doubt because of the diminished num- formerly remote sites. The extensive conversion of ber of individuals and fragmented distribution of the native forests to exotic pine plantations in central Chile overall population. Individuals dispersing from scat- now extends into the lower Andes (Burschel et al., 1991; tered groups may be lost from the population by failing Manzur, 1993). Livestock, present at most huemul sites, to find another group. Of special significance is the lack impact soils and vegetation through overgrazing and of clear evidence for recolonization of sites from which trampling. Herders burn protective cover to improve huemul have disappeared (Table 1). At Cabras, mini- site access and grass production. mum counts of huemul dropped from six in 1976 to one in 1986 (Table 2), when only old huemul signs were 3.4.4. Low reproduction and recruitment detected. A lack of recent browsing was noted on A fawn-adult female ratio of ca. 1:2 was reported for favored plants such as Schinus sp. and Maytenus boaria huemul in the Nevados area (Povilitis, 1979). Since (Povilitis, 1979). However, minimum counts rose by 1980, fawns were detected on only 33% of surveys at 1996, and a change in summer range hinted at the pre- sites having huemul (Table 2). In south Chile, a female sence of a new huemul group at the site. Still, it is monitored from 1980-1986 raised only three fawns, one uncertain if the site was ever entirely vacated. of which died within six months. She became pregnant At Dumas, surveys suggested huemul presence in for at least five of those years (Aldridge, 1988). Her 1975, absence in 1980, and presence again in 1992. female offspring did not reproduce until age four. Hue- However, given the large size and difficult topography mul almost always gave birth to only a single fawn of the site, an immediate follow-up survey to the 1980 (Povilitis, 1979; Montecinos, 1995). work would have been required to confirm huemul absence. 3.4.5. Small and fragmented population Spatial separation of huemul groups is believed to be A huemul population of ca. 60 individuals risks behaviorally enforced by huemul fidelity to restricted extinction by virtue of its small size. Berger (1990) home range areas. Distinct annual ranges of 321 and found that bighorn sheep Ovis canadensis populations 358 ha were reported for huemul groups at Chillfin and (n = 129) below a threshold of 50 animals went extinct Cabras sites in 1975-1976 (Povilitis, 1979). There was within 50 yr. Bighorns, like huemul, are rugged terrain no evidence for movement by huemul from these and specialists that often occur in fragmented habitats. two other sites closely monitored during that time. Nei- Fluctuation in group size, resulting from random ther summer nor wintering areas of groups overlapped, demographic and environmental events, likely plays a key as seasonal movements by huemul to lower elevation role in the disappearance of small, isolated huemul groups involved single mountain-ridge systems. (Povilitis, 1979). For example, drought in central Chile Evidence elsewhere also suggests site affinity of hue- from 1993-1997 could account for low huemul numbers mul. A naturally marked female was regularly observed recorded at that time. Growing season (September- from 1980 until her death in 1986 at a 275ha site in March) precipitation fell 41% below a 30-yr average southern Chile (Luis Montecinos, wildlife guard, (Universidad de Concepcion, 1993-1996). The drought CONAF, pers. comm.) Her breeding season range appeared to reduce available forage and eliminated some (March-April) was restricted to 70 ha during two con- water sources. It may have been responsible for the loss secutive seasons of close monitoring (Povilitis, 1985). of huemul from the Gertrudis site in 1996-1997, where Two other huemul, a male and female, appeared to free water became largely unavailable during summer. occupy roughly the same area for at least 5 and 15 yr, Other concerns include potential inbreeding, low respectively. At another Patagonia site, five of six hue- genetic variability, and random loss of adaptive alleles mul, radio collared beginning in 1989, were reported at through genetic drift (Smith-Flueck and Flueck, 1995). their original capture area in 1995 (John Bahamondes, Loss of genetic variation can be particularly high for wildlife biologist, CONAF, pers. comm.). populations experiencing local extinctions (Gilpin, While fragmentation of the Nevados huemul popula- 1991). Genetic factors may contribute to disease sus- tion is evident, I believe that interchange between hue- ceptibility and poor reproduction (Allendorf and Leary, mul groups can be restored through an enhanced 1986; Gilpin, 1987; Frankham, 1995) in huemul. conservation program. Huemul seem capable of moving 102 A. Povilitis/Biological Conservation 86 (1998) 97-104 between sites which are in comparative proximity. In the Nevados area, they were infrequently located beyond primary habitat sites: 1.5 km from Jos6 in 1975; 3.0km from lqiblinto in 1993; 3.0 km from Catalinas in North Core 1995; and 1.5 km from Palo in 1996. The limited huemul Area sign found in each case suggested individual animals either dispersing or wandering from their home ranges. One or two unfamiliar adult males and a subadult ( N female-male pair appeared briefly at a study site in Patagonia (Povilitis, 1985) indicating similar transient t movement. 1Okra A capability to disperse between nearby sites is con- ! ! sistent with the huemul's robust physical build and South Com -1,Z Z'3 ~1 ~ observed mobility (Osgood, 1943; Povilitis, 1979). On Area several occasions, for example, I tracked huemul within / , primary sites for 3-7 h, meandering through 2-3 km of / I I Polcura rugged terrain. Restoration Area

4. Recommendations I Efforts to recover fragmented populations of large mammals like the huemul can draw on four tenets of biological restoration: (1) protection of habitat (population) core areas; (2) linkage of habitat areas; (3) conservation planning and management; and (4) Fig. 4. Proposed huemul reserve for the Nevados de Chillhn area of population augmentation (Caughley and Gunn, 1996; Chile. Primary sites with reserve status as of 1997 are hatched. Arrows Hunter, 1996). show principal habitat corridors.

4.1. (1) Habitat core areas (Shafer, 1990). Planning for habitat conservation is critical for this rapidly developing corridor area. The highest ranked habitat sites (Table 3) should be The land matrix around primary habitat sites also secured on a priority basis as strict reserves, i.e. where needs protection. Priority should be given to connecting threats such as poaching, livestock-related disease, and secondary habitats most threatened by development habitat degradation are entirely eliminated. To com- (housing construction, logging, roads, etc.). In addition, plete the core areas, other habitats should be added as corridors of natural cover extending north and south of strict reserves or as special conservation areas. Core the Nevados area should be preserved as natural areas areas should provide enough secure habitat to sub- (Fig. 4). stantially reduce the risk of population extinction. For huemul in the Nevados area, two habitat core 4.3. (3) Conservation management areas each consisting of 8-11 priority sites are recommended (Fig. 4). Chile's private conservation organiza- Conservation management plans are needed for tion (CODEFF), with help from the Frankfurt habitat which cannot be strictly protected. Private Zoological Society and others, has acquired the landowners in the northern part of the Nevados area lqiblinto site as a key building block for the north core have begun such planning (Espinosa, 1996; Militza area. Negotiations between CODEFF, Chile's forest Saavedra, landowner, pers. comm.). Plans should safe- service (CONAF), and local landowners are underway guard key water sources, escape terrain, foraging and to obtain the Diguillln, the top-ranked site for the south resting areas, and winter habitat of huemul. These core area. localities should be free of livestock, domestic dogs, wood cutting, fire, and other human disturbances. Plans 4.2. (2) Habitat connectivity may also include measures to actively restore vegetation cover at sites damaged by livestock and fire, and to North and south core areas should be linked by a protect other aspects of biological diversity, such as old- band of secure habitat that crosses the Josr, Chillfin, growth forest and wetlands. and Renegado sites (Fig. 4). The sites themselves need Conservation management for private lands is a key special protection as population 'stepping stones' to huemul recovery. As of 1997, only three primary A. Povilitis/BiologicalConservation 86 (1998) 97-104 103 habitat sites in the Nevados area (Capados, Blanquillo, Allendorf, F.W., Leary, R.F., 1986. Heterozygosity and fitness in and Catalinas) were protected public lands (Fig. 4) as natural populations of animals. In: Soule, M.E. (Ed.), Conservation part of a national forest reserve administered by Biology: the Science of Scarcity and Diversity. Sinauer Associates, Inc., Sunderland, pp. 57-76. CONAF (Ortiz, 1993); funding for additional reserve Armesto, J., Villagrfin, C., Donoso, C., 1994. La historia del bosque acquisitions in Chile was extremely limited. Potential templado Chileno. Ambiente y Dcsarrollo (Chile, marzo 1994), 66-72. benefits from tourism, education, and research (Lan- Berger, J., 1990. Persistence of different sized populations: an empiri- gholz, 1996) provide added incentive for landowners to cal assessment of rapid extinction in bighorn sheep. Conservation Biology 4, 91-98. manage huemul. Priority ranking for huemul site Burschel, P., Edens, J., MoreUo, J., 1991. Politica de manejo del bos- acquisition and conservation planning is provided in que nativo en Chile. Informe Ttcnico FO:TCP/CHI/0052(a). United Table 3. Nations Food and Agricultural Organization, Santiago. Cabrera, A.L., Yepes, J., 1960. Mamiferos Sud-Ameicanos. Ediar S.A. 4.4. (4) Population augmentation Editores. Buenos Aires. Caughley, G., Gunn, A., 1996. Conservation Biology in Theory and Practice. Blackwell Science, Cambridge. An augmentation project should be planned in case Diaz, N.I., 1993. Changes in the range distribution of Hippocamelus the Nevados huemul population fails to improve with bisulcus in Patagonia. Zeitschrift Sauget 58, 344-351. enhanced habitat protection. Translocation of huemul Drouilly, P., 1983. Recopilacirn de antecedentes biologicos y ¢cologi- from Patagonia (assuming suitable donor populations cos del huemul Chileno y consideraciones sobre su manejo., CONAF Boletln Trcnico No. 5 (Chile). exist) could strengthen huemul numbers and increase Espinosa, P., 1996. Intentan salvar a sesenta huemules de la extincirn. gene flow in the population (Griffith et al., 1989; E1 Mercurio (Chile, 11 de matzo). Santiago. Hedrick, 1995), especially in the Polcura Valley area Frankham, R., 1995. Inbreeding and extinction: a threshold effect. (Fig. 4). Translocation, however, should be carefully Conservation Biology 9, 792-799. planned to minimize risk of outbrceding depression (see, Frid, A., 1994. Observations of habitat use and social organization of a huemul Hippocamelus bisulcus coastal population in Chile. Biolo- for example, Hedrick, 1995; and Maehr and Caddick, gical Conservation 67, 13-19. 1995). Gilpin, M., 1987. Spatial structure and population vulnerability. In: The conservation measures described here may also Soule, M.E. (Ed.), Viable Populations for Conservation. Cambridge serve to restore scattered huemul populations in Pata- University Press, Cambridge, pp 125-139. gonia. For full species recovery, huemul should be rees- Gilpin, M., 1991. The genetic effective size of a metapopulation. Bio- logical Journal of Linnean Society 42, 165-175. tablished in other ecological regions where they Griffith, B., Scott, J.M., Carpenter, J.W., Reed, C., 1989. Transioca- formerly occurred, such as Nothofagus shrub-steppe of tion as a species conservation tool: status and strategy. Science 241, southern Argentina and Nothofagus-Araucaria forest of 477-480. south-central Chile (Povilitis, 1997). Long-term restora- Hanski, I., Gilpin, M., 1991. Metapopulation dynamics: brief history and tion should aim for continuity between central and conceptual domain. Biological Journal of Linnean Society 42, 3-16. Harrison, S., 1991. Local extinction in a metapopulation context: an Patagonian populations. These steps would greatly empirical evaluation. BiologicalJournal of Linnean Society 42, 73-88. improve the huemul's prospects for adaptive evolution, Hedrick, P.W., 1995. Gene flow and genetic restoration: the Florida and restore its former role as a widespread wild herbi- panther as a case study. Conservation Biology 9, 996-1007. vore and living cultural icon of the southern Andes. Hoffman, A., 1993. Conservacirn de la biodiversidad de Chile. Revista Vivienda y Decoracirn (Chile, 24 de abril), 53-55. Hunter, M.L., 1996. Fundamentals of Conservation Biology. Black- well Science, Cambridge. Acknowledgements International Union for the Conservation of Nature (IUCN), 1990. The IUCN red list of threatened animals. Gland, Switzerland. I thank the Corporacirn Nacional Forestal de Chile Langholz, J., 1996. Economics, objectives, and success of private nature reserves in sub-Saharan Africa and Latin America. Conserva- (CONAF), the Comit6 National Pro Defensa de la tion Biology 10, 271-280. Fauna y Flora de Chile (CODEFF), Earthwatch (USA), Lesica, P., Allendoff, F.W., 1995. When are peripheral populations Life Net (USA), and the Sierra Institute, University of valuable for conservation? Conservation Biology 9, 753-760. California, Santa Cruz USA for supporting field surveys Lopez, R., 1994. Estudio de la situaci6n actual de la poblacirn de for the huemul. I applaud the many hearty volunteers, huemules en la cordillera de la VIII Regirn del Bio-Bio, Chile. (noviembre de 1993 a mayo de 1994). Informe, CONAF Gerencia students, and wildlife guards who conducted the work. I Trcnica, Depto. de Patrimonio Silvestre, Concepcirn, Chile. extend my gratitude to the Stutzin family which so often MacNamara, M.C., 1982 Huemul: a deer of distinction. Animal hosted my stay in Chile. A. Gaines kindly helped illus- Kingdom, 85, 5-13. trate this paper. Maehr, D.S., Caddick, G.B., 1995. Demographics and genetic intro- gression in the Florida panther. Conservation Biology 9, 1295-1298. Manzur, M.I., 1993. Some relevant information concerning the destruction of the native forest huemul habitat in the VIII Region of References Chile. Serie Documentos. CODEFF Biodiversity Project. Santiago. Miller, S., 1980. Human influences on the distribution and abundance Aldridge, D.K., 1988. 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