Effects of Livestock on Guanaco Lama Guanicoe Density, Movements and Habitat Selection in a Forest–Grassland Mosaic in Tierra Del Fuego, Chile

Home , Guanaco

Effects of livestock on guanaco Lama guanicoe density, movements and habitat selection in a forest–grassland mosaic in Tierra del Fuego, Chile

C LAUDIO A. MORAGA,MARTÍN C. FUNES,J.CRISTÓBAL P IZARRO C RISTÓBAL B RICEÑO and A NDRÉS J. NOVARO

Abstract Locally abundant ungulates often come into con- Keywords Camelid conservation, Chile, Lama guanicoe, flict with human activities. After a population collapse that livestock, partial migration, sheep ranching, sub-Antarctic reached its nadir in the s, the guanaco Lama guanicoe forest, ungulates population in Tierra del Fuego, Chile, recovered and is now in conflict with sheep ranching and commercial logging. We studied the effects of livestock density and en- Introduction vironmental factors on guanaco abundance and spatial ecology, using seasonal counts and radio-telemetry in a pri- ocally abundant ungulates often have strong effects on vate protected area (Karukinka) and neighbouring ranches plant communities, sometimes leading to conflict with – L in a forest grassland mosaic in Tierra del Fuego. Guanaco human activities (Graham et al., ; Rutherford & density was highest in low-elevation areas with more grass- Schmitz, ). Understanding the processes that are re- land cover and little snow accumulation in winter. In low- sponsible for increases and declines in ungulate abundance, elevation areas, guanaco density decreased with increasing and ultimately for their ecosystem effects, is necessary to livestock density. Radio-tracked guanacos exhibited a partial guide conservation of ungulates and their habitats (Bolger migration pattern: two individuals migrated seasonally, se- et al., ; Gordon, ; Graham et al., ). lecting grasslands and avoiding forests mainly in summer, The guanaco Lama guanicoe has been the only native whereas six sedentary individuals used habitats according ungulate on Tierra del Fuego Island, at the southern tip of to their availability. Migratory guanacos spent the summer South America, for the last , years (Sarno et al., ). in Karukinka and winter on nearby ranches. High sheep Until large-scale colonization by Europeans and their live- densities and poor range condition on the ranches reduce stock began in the late th century, guanacos were the key forage resources available to guanacos and may promote main source of protein for the indigenous people, the use of forests by guanacos, affecting forest regeneration and Selk’nam, who followed the guanaco’s seasonal migration increasing conflict with logging. Current guanaco harvest by between forested highlands and grassy lowlands (Bridges, loggers may fail to reduce the impact of guanacos on logged- ). The number of sheep in Chilean Tierra del Fuego forest regeneration if guanaco spatial ecology and sheep peaked at  million in the early s (Martinic, ). management are not considered. Our results provide insight The Selk’nam were driven to extinction through coloniza- into the interactions among guanacos, forests and livestock tion of their land and persecution. By the mid s, as a ranching, and may be used to reduce conflicts and guide result of hunting, competition from sheep, and habitat conservation in the Fuegian ecosystem. degradation, the guanaco population had collapsed to c. , on the Chilean side of the island (Raedeke, ). Since then, guanaco numbers have recovered to . , as a result of hunting restrictions and reduced sheep num- † CLAUDIO A. MORAGA* (Corresponding author), J. CRISTÓBAL PIZARRO and bers (Skewes et al., ; CONAMA, ). CRISTÓBAL BRICEÑO‡ Karukinka, Chile Program, Wildlife Conservation Society, Punta Arenas, Chile. E-mail [email protected] Their recovery, however, has brought guanacos into con-

MARTÍN C. FUNES Patagonian and Andean Steppe Program, Wildlife flict with another human activity expanding in Tierra del Conservation Society, Junín de los Andes, Neuquén, Argentina Fuego: commercial logging of the sub-Antarctic forest. ANDRÉS J. NOVARO INIBIOMA-CONICET and Patagonian and Andean Steppe This is the forest that occurs at the highest latitude in the Program, Wildlife Conservation Society, Junín de los Andes, Neuquén, southern hemisphere and it is being logged, mostly for tim- Argentina ber export, at an annual rate of c. , ha, throughout the *Current address: School of Natural Resources and Environment, and Department of Wildlife Ecology and Conservation, University of Florida, USA Magallanes region (Wildlife Conservation Society, unpubl. †Current address: Department of Environment and Resource Studies, data; Donoso & Lara, ; Huber & Markgraf, ). University of Waterloo, Canada ‡ Unlike the rest of their range, where guanacos live in grass- Current address: Department of Preventive Medicine, Faculty of Animal and  Veterinary Sciences, Universidad de Chile, La Pintana, Santiago, Chile lands and shrublands (de La Tour, ), in Tierra del Fuego Received  November . Revision requested  April . much of the population lives in the transition zone between Accepted  August . First published online  October . forests and grasslands (Raedeke, ; CONAMA, ).

© 2014 Fauna & Flora International, Oryx, 49(1), 30–41 doi:10.1017/S0030605312001238 Downloaded from https://www.cambridge.org/core. IP address: 170.106.33.14, on 23 Sep 2021 at 18:05:34, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605312001238 Effects of livestock on guanacos 31

Guanaco herbivory affects regeneration growth patterns of The area includes steppe–forest ecotone and mountain southern beeches, particularly Nothofagus pumilio (Dodds, forest habitats (Pisano, ). Dominant plant communities ; Rebertus et al., ), limiting timber production include beech forests, matorral shrublands, Sphagnum bogs, (Martínez-Pastur et al., ; Cavieres & Fajardo, ). meadow–upland mosaics, and steppe grasslands. Tree spe- Guanaco effects on tree growth and regeneration, however, cies include the deciduous lenga Nothofagus pumilio and have not been disentangled from effects of livestock and log- ñirre Nothofagus antarctica and the evergreen coihue de ging, and it is not clear what effect guanacos have on forest Magallanes Nothofagus betuloides. Main shrub species are dynamics at the landscape scale (Martínez-Pastur et al., michay Berberis ilicifolia, calafate Berberis microphylla and ; Cavieres & Fajardo, ). This is a common de- romerillo Chiliotrichum diffusum. Dominant grasses are ficiency in traditional models of ungulate herbivory inter- Festuca, Poa and Alopecurus (Pisano, ). actions (Wisdom et al., ). Nevertheless, the Chilean government recently allowed the main timber company to begin harvesting guanacos to reduce their density and for Methods sale of their meat. Indirect evidence suggests that grasslands are the pre- At the landscape scale, we studied how environmental fac- ferred habitat of guanacos in Tierra del Fuego but that tors and human disturbances, including sheep numbers, af- they use forest patches because of displacement by sheep fect seasonal changes in guanaco density along an altitudinal (Raedeke, ). At a mainland site where guanacos were gradient from low-elevation grasslands in the north, pri- sedentary, sheep were shown to exclude guanacos from marily on ranches, to high-elevation forests and alpine mea- grasslands through resource competition (Baldi et al., dows in the south, mainly within Karukinka (Fig. , Table ). , ). The effects of sheep ranching on guanaco den- Densities of guanacos and livestock were estimated using the sity and habitat use and selection have not been studied line transect method (Buckland et al., ) by surveying  quantitatively in the grassland–forest mosaic of Tierra del – km transects seasonally along randomly chosen sections Fuego. In addition, some guanacos may still migrate season- of all available secondary roads. Transects were surveyed ally between forests and grasslands (Raedeke, ), as a from the austral spring of  to the spring of  ( sea- combined effect of the guanaco’s habitat requirements and sons in all). Surveys were conducted during daylight hours, competition with domestic ungulates. We thus need to have when guanacos are active, from an open pick-up truck, with a better understanding of seasonal changes in guanaco ecol- two observers standing in the bed (Baldi et al., ). − ogy and of any implications of migration for the conser- Travelling speed was – km h . Distances between gua- vation and management of the species (Berger, ; naco groups and the vehicle were recorded with a laser Bolger et al., ; Harris et al., ). rangefinder, and bearing to the transect line with a compass. Here we assess the effects of sheep density and other en- Guanaco and livestock densities were estimated using vironmental factors on the seasonal abundance of the gua- DISTANCE v. . (Thomas et al., ). naco and describe the species’ movements and habitat To assess vegetation cover types along transects we selection in a forest–grassland mosaic in southern Tierra plotted transects on a geographical information system del Fuego. To do this we combine a landscape-level analysis layer in ArcView v. . (ESRI, Redlands, USA), intersected of guanaco density patterns and a habitat-level analysis of this layer with a vegetation cover map (WCS, ) and es- guanaco movement patterns in a private protected area timated percentage cover of each vegetation type in a  m and on adjacent ranches. buffer zone around each transect. Cover types considered were grasslands (steppe, bushland mosaic, alpine), forests (deciduous and evergreen) and meadows (peat bogs, mea- – Study area dow upland mosaics). Effects of environmental and anthropogenic factors on Our study was conducted in southern Tierra del Fuego guanaco densities were analysed using a generalized linear Island, Chile, on the eastern edge of the Karukinka private model (GLM) with a Poisson distribution and log-link func- protected area and adjacent ranches used mostly for sheep tion (Dobson, ). The Wald statistic was calculated to grazing (Fig. ). Karukinka has had little or no livestock evaluate the relative effect of each independent variable. grazing since , and since  has been managed as a We developed a maximized generalized model for guanaco protected area, with no livestock ranching, by the Wildlife density to evaluate the overall effect of each independent Conservation Society (Saavedra et al., ). Monthly variable. The independent variables considered for the mean temperatures range between –°C in winter and GLM were year, season, percentage of grassland cover, per- °C in summer and mean annual precipitation is  centage snow cover and depth, and livestock density. All mm, with frequent snowfall during autumn and winter density values were natural-log transformed. We then con- (Tuhkanen et al., ). ducted a backward removal of variables from the maximized

FIG. 1 The location of the study area on Tierra del Fuego Island. The survey transects are shown along with the capture area for guanacos Lama guanicoe that were radio-tracked across open and closed habitats. The rectangle on the bottom inset shows the location of the main map in Chile.

model using P = . to assess which variable was more re- through direct observation whenever possible. After July lated to guanaco density. Because high-elevation transects  we did not confirm locations with direct observations had almost no livestock we also tested the effect of livestock and therefore excluded these locations from habitat-use on guanaco density by building a model with data from low- analyses, utilizing them only to assess shifts between elevation transects, including livestock density, snow depth, summer and winter ranges (see below). LOAS v. . and year as independent variables. Both GLM analyses were (Ecological Software Solutions, Sacramento, USA; Larson, carried out using Statistica v.  (StatSoft, Tulsa, USA). ) was used for bearing analyses. To study seasonal To examine the processes that led to seasonal changes in movements we divided the telemetry data into summer guanaco abundance at the landscape scale, we studied gua- (November–April) and winter (May–October) to account naco movements and habitat selection at a finer spatial scale for key seasonal changes in guanaco social behaviour using radio-telemetry. Our intensive study area corres- (Ortega & Franklin, ). To estimate seasonal home-range ponded to the central portion of the altitudinal gradient size and location for each guanaco we generated minimum along the border between Karukinka and the neighbouring convex polygons (MCPs) using the Animal Movement v. . ranches. Guanaco social structure includes family and bach- extension in ArcView (Hooge & Eichenlaub, ). We then elor groups (Franklin, ). We investigated primarily the plotted seasonal home-range sizes against number of loca- movements of family groups, which are usually territorial. tions to estimate the minimum number of locations re- Ten adult guanacos from family groups were captured and quired to achieve stable estimates of home-range size. To radio-collared with VHF transmitters (Advanced Telemetry quantify seasonal movements we calculated a centroid for Systems, Isanti, USA) between January  and January each seasonal home range and measured the distance be- , following capture procedures described in Karesh tween centroids for each guanaco, using radio-telemetry lo- et al. (), and radio-tracked until September . Data cations until September , and detected migration when for a guanaco that lost its transmitter  months after capture we observed a complete round-trip shift between seasonal were excluded from the analysis. We restricted the capture home ranges that were not used at other times of the year area (Fig. ) to maximize the likelihood that guanacos (Berger, ). were using a variety of habitats in Karukinka and the adja- For the habitat selection analysis we determined habitat cent ranches. Telemetry readings were taken during daylight use by assigning each guanaco location to a vegetation type. hours. Until July  we confirmed guanaco locations We determined vegetation types available in four types of

© 2014 Fauna & Flora International, Oryx, 49(1), 30–41 Downloaded from https://www.cambridge.org/core. IP address: 170.106.33.14, on 23 Sep 2021 at 18:05:34, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605312001238 https://www.cambridge.org/core/terms Downloaded from © 04Fua&FoaInternational, Flora & Fauna 2014 https://www.cambridge.org/core . https://doi.org/10.1017/S0030605312001238 Oryx 91,30 49(1), , . IPaddress: TABLE 1 Seasonal and annual variation in guanaco Lama guanicoe and sheep densities on the  survey transects in Tierra del Fuego (Fig. ) during – (S, summer; A, autumn; W, winter; P, spring; years are subscripted), with mean percentage grassland and snow cover and mean snow depth.

– −2 −2 41 Mean guanaco densities ± SE km Mean sheep densities ± SE km 170.106.33.14 Mean snow Mean snow Mean transect (no. of groups); seasonsyears (no. of groups); seasonsyears Grassland cover ± SE depth ± SE altitude ± SE (m) Min Max Min Max cover (%) (%) (cm) ± ± ± ± ± ±

, on 175 2 2.3 0.5 (3); P10 34.8 5.8 (40); W07 0.0; S08,09,10 ,A08,09,10 ,W09,10 , 319.3 188.3 (21); P06 82.3 37.2 1.3 9.6 1.3 P

23 Sep2021 at18:05:34 09,10 169 ± 2 1.0 ± 0.1 (2); A10 21.8 ± 3.6 (25); W07 0.0; S10,A10,W10,P10 187.7 ± 70.8 (11); A08 55.3 34.2 ± 0.5 8.1 ± 1.2 167 ± 2 1.9 ± 0.3 (4); A10 36.9 ± 3.7 (36); A08 8.7 ± 3.5 (6); P09 221.8 ± 83.7 (13); A08 67.0 41.7 ± 2.2 11.8 ± 1.6 163 ± 1 6.8 ± 0.9 (20); W10 44.7 ± 6.3 (52); P06 0.0; S07,08,09,10 ,A08,09,10 , 61.6 ± 23.5 (6); A07 86.1 45.0 ± 1.7 11.8 ± 1.6 W07,08,09,10 ,P06,07,08,09,10 176 ± 2 4.8 ± 0.7 (14); W10 37.1 ± 4.9 (55); P08 0.0; S07, 08,09,10 ,A08,09,10 , 123.2 ± 47 (12); A07 70.6 52.1 ± 1.8 16.4 ± 1.8 W07,08,09,10 ,P06,07,08,09,10 , , subjectto theCambridgeCore termsofuse,available at 216 ± 2 8.2 ± 1.1 (34); A08 50.0 ± 9.2 (30); S10 * * 28.1 52.1 ± 1.3 12.5 ± 0.7 216 ± 1 3.3 ± 0.5 (12); A07 43.6 ± 10.8 (18); S07 * * 48.7 67.1 ± 1.4 18.2 ± 0.9 211 ± 1 1.4 ± 0.3 (1); P10 37.1 ± 6.9 (6); S10 * * 21.0 66.0 ± 1.3 15.0 ± 0.7 237 ± 2 1.5 ± 0.2 (2); A08 26.1 ± 6.5 (6); S07 * * 26.5 45.0 ± 0.4 6.7 ± 0.8 261 ± 2 0.3 ± 0.0 (4); W07 46.1 ± 7.2 (59); P06 * * 60.1 57.1 ± 0.4 25.1 ± 1.9 527 ± 6 0.0; A08,09 W07,09,10 7.3 ± 1.5 (16); S08 * * 14.7 65.8 ± 0.5 37.5 ± 2.1 202 ± 2 0.0; S06,07 W07,09,10 P06,07 0.8 ± 0.1 (3); P09 * * 17.6 62.5 ± 0.9 40.0 ± 1.9 338 ± 9 0.0; S09,10 A09 W07,09 3.2 ± 0.6 (7); S08 * * 3.9 59.2 ± 1.4 34.7 ± 2.3 563 ± 10 0.0; A08,09 W07,09,10 5.5 ± 1.1 (12); S08 * * 3.2 70.8 ± 1.4 42.5 ± 1.6 fet flvsoko uncs33 guanacos on livestock of Effects *No sheep ranching during any survey 34 C. A. Moraga et al.

ranges: ranges of sedentary guanacos that remained most of TABLE 2 GLM estimates of guanaco densities in Tierra del Fuego the year in Karukinka, ranges of sedentary guanacos that re- (Fig. ) for all transects and for low-elevation transects only (–  mained most of the year on the ranches, and winter and m). summer ranges of migratory guanacos. We plotted four Effect Estimate ± SE Wald statistic P MCPs with all locations in these four guanaco range types All transects and intersected them with the same vegetation map used ±  Intercept 0.35 0.11 10.21 0.001396 for the GLM analysis. Using the χ statistic, we performed a Grassland cover 0.01 ± 0.00 32.93 0.000000 habitat selection analysis, comparing proportions of locations Snow depth −0.05 ± 0.01 23.71 0.000001 recorded and expected according to available vegetation Scale 1.00 ± 0.00 cover. When selection was significant, we used Bonferroni Low-elevation transects only ± confidence intervals to test for differences between use and Intercept 108.25 45.05 5.77 0.016264 Year −0.05 ± 0.02 5.66 0.017355 availability of each habitat. To represent the magnitude of − ± ’ Snow depth 0.02 0.01 14.63 0.000131 the selection we calculated Manly s standardized selection Sheep density −0.03 ± 0.02 5.01 0.025188 index (Bi) for each habitat, which is based on subsequently Cattle density −0.08 ± 0.04 3.95 0.046813 standardized selection ratios (Manly et al., ). Scale 0.49 ± 0.00 Standardized ratios provide indices of selection that are di- rectly comparable between seasons. Independence of locations for analyses of home-range size and habitat selection Guanaco seasonal movements and habitat selection was ensured by obtaining telemetry readings every  days (on average), at different times of the day. All but one (ID ) of the radio-tracked guanacos were members of family groups. Visual confirmation of group – Results status was obtained on occasions per individual (Table ). From these groups we detected only two mi-   Factors affecting guanaco density gratory guanacos. They were both females ( and ) and migrated from Karukinka to sheep ranches in summer − Guanaco densities averaged . km in low-elevation and winter, respectively (Table , Figs  & ). These mi- transects with abundant grass cover (mean .% grassland) gratory guanacos spent summers in Karukinka as part of − and sheep grazing throughout the year, . km in mid- family groups (Table ), and winters either entirely or par- elevation transects with intermediate grass cover (mean tially on sheep ranches. Guanaco  ranged in winter onto − .% grassland) and limited livestock grazing, and . km a ranch in neighbouring Argentina with similar land use to in high-elevation transects with dominant forest cover (mean Chilean sheep ranches and returned to the  summer .% forest) and no ranching (Table ). Guanacos were pres- home range in the summers of – (Fig. ). ent in high-elevation transects only during spring and sum- Migratory guanacos expanded their ranges in winter: mer. Sheep density in low-elevation transects peaked in mean home-range size was  times larger in winter (. −   spring (mean . ± SE . km ) and autumn (mean . ± ± SE . km ,n=) than in summer (. ± SE . km , − − SE . km ) and was low in summer ( ± SE . km )and n=; Table ). Guanaco  died at the end of the  win- − winter (. ± SE . km ). Mean annual density of cattle was ter, when it was moving towards its previous year’s summer − low in the first part of the study (. ± SE . km in  and range (Fig. ). A necropsy indicated poor nutritional con- − −  km in ) and increased to . ± SE . km in  dition (based on marrow fat content) and a widespread sar- − and . ± SE . km in . Horse density was stable and coptic mange infection. − averaged . ± SE . km during the study. Cattle and Six other radio-tracked guanacos were considered seden- horses occurred mainly on low-elevation transects. tary because they did not make significant shifts between The model indicated that guanaco densities in the study seasonal ranges: mean distance between the centroids of area were positively associated with grassland cover and their summer and winter ranges was . ± SE . km negatively associated with snow depth during autumn and (Table , Fig. ). Five of the sedentary guanacos were part winter (Table ). The analysis that included only transects of family groups every summer. Male guanaco  was at low elevation indicated that guanaco density was nega- part of a family group when captured but joined a bachelor tively associated with livestock density (Table ). There group  months after capture. Four (three males and one fe- was an overall decline in guanaco density in the area during male) of the five sedentary guanacos in family groups had the study, as indicated by the significant negative effect of –% of their home ranges inside Karukinka, whereas year in the GLM. Between  and  mean annual gua- the remaining one (male )had% of its range within naco density across all transects declined by .%. The gua- a neighbouring ranch (Fig. ). This particular ranch had a − naco decline occurred as sheep numbers declined (%) and high density of sheep (– km ) only during autumn cattle increased.  and no livestock during the remainder of the study.

TABLE 3 Home range of the  radio-tracked guanacos in Tierra del Fuego (Fig. ) in successive seasons during January –September , with age and gender, type of social group, mean distance between home-range centroids, number of sightings with social group, period of radio-tracking, size of annual (or summer and winter) home range, and percentage of home range within Karukinka protected area.

2 Mean distance ± SE No. of Home range, km (no. of locations) Age & Type of between centroids, km sightings Tracking % within Guanaco gender group (no. of seasons) with group period Annual Summer Winter Karukinka 1014 Adult Family 0.58 ± 0.18 (4) 12 Jan. 2008– 2.10 (22) 94 male June 2009 1045 Adult Family 0.57 ± 0.16 (3) 17 Jan. 2008– 2.07 (28) 75.1 male June 2009 1083 Adult Family 1.67 ± 1.12 (5) 8 Jan. 2008– 1.22 (25) 100 male June 2009 1123 Adult Family 1.31 ± 0.51 (5) 4 Jan. 2008– 6.08 (28) 5.9 male June 2009 1161 Young Family 2.66 ± 1.0 (5) 6 Jan. 2008– 5.67 (29) 99.7 adult June 2009 female 9451 Young Bachelor 3.54 ± 0.48 (3) 23 Sep. 2007– 12.23 (24) 100 adult June 2009 male 984 Young Family 11.62 ± 5.34 (4) 10 Jan. 2008– 4.70 (18) 100 adult Oct. 2008 female 843 Adult Family 8.87 ± 2.36 (8) 25 Jan. 2007– 0.35 (27) 39.04 (13) 28 female with Oct. 2007 chulengo (calf) Nov. 2007– 0.41 (6) (2)3 Oct. 2008 Nov. 2008– 4.34 (8) (3)3 June 2009 8842 Young Family with 6.50 (2) 25 Jan. 2007– 1.86 (20) 17.51 (26) 88.5 adult chulengo Aug. 2007 female (calf)

 Bachelor captured in a family group initially  Died during study  Home range not calculated because of small sample size

 Annual home range averaged . ± SE . km for sedentary indicated that during summer migratory guanacos guanacos from family groups and was three times larger for selected the small grassland area within Karukinka about bachelor  (Table ). five times more often than deciduous forests and four Female guanaco  behaved as sedentary in the summer times more often than meadows. Conversely, during winter, and winter of  but in the spring of  moved perman- when much of the grassland habitat in the ranches was oc- ently c.  km to the north-west (Fig. ). During the summer cupied by sheep, migratory guanacos selected meadows and winter of  she also behaved as sedentary in her new about four times more often than grasslands and forests range, which encompassed Karukinka and an adjacent (Table ). ranch devoted to logging. Migratory guanacos occurred in areas of higher livestock There were significant differences in the proportions of density during winter (mean . sheep and . cattle − habitats used by migratory guanacos in summer and winter, km ), whereas the sedentary guanacos (,  and but not by sedentary guanacos (Table ). The Bonferroni ) used the ranches with few or no livestock after autumn confidence limits indicated that migratory guanacos  towards the east of Karukinka (Fig. ). Although there used forests less than expected in summer but did not was no significant habitat selection by sedentary guanacos significantly select or avoid any particular habitat during that lived mostly within Karukinka, in summer they tended winter (Table ). Manly’s standardized selection index to avoid forests (Table ).

FIG. 2 Seasonal home range of migratory guanacos (,  and  in (a), (b) and (d), respectively) and (c) annual home range of sedentary guanacos (, , , ,  and ) followed by radio-tracking in Tierra del Fuego, including minimum convex polygon areas by season (S, summer; W, winter) and year (e.g. W). No winter  data were available for guanaco . The stars in (a) show the positions of guanaco  in winter , and the star in (d) shows the last position of guanaco  in winter .

Discussion Increases in guanaco density at high elevations during summer and at mid and low elevations during spring and Factors affecting guanaco seasonal density autumn are consistent with altitudinal movements of other mountain-dwelling ungulates that migrate seasonally, We observed that a large proportion of the guanaco popu- such as black-tailed deer Odocoileus hemionus columbianus, lation studied in the forest–grassland mosaic had a prefer- elk, and red deer Cervus elaphus (Schoen & Kirchhoff, ; ence for grasslands, as reported elsewhere where forests are Hebblewhite et al., ; Pépin et al., ). These increases absent (Puig et al., ). Firstly, guanaco density at the land- in guanaco density at high elevations during summer were scape scale was highest throughout the year where grassland mostly associated with movement of family groups and soli- cover was highest (Tables  & ). Secondly, guanaco density tary guanacos in Karukinka. This relationship between den- during summer, when all habitats were available because of sity and elevation has also been observed for guanacos in the absence of snow, was lowest at high elevations where the Torres del Paine National Park during summer (Ortega & proportion of grassland cover was lowest (Table ). These Franklin, ; Young & Franklin, ). patterns are consistent with those of other ungulates with Snow depth and cover limited the density of guanacos at mixed diets that inhabit grassland–forest mosaics, such as high elevations during winter. A similar pattern has been elk Cervus canadensis, white-tailed deer Odocoileus virginia- observed at the landscape scale for other mountain ungu- nus and mule deer Odocoileus hemionus hemionus, which at lates, including the Alpine chamois Rupicapra rupicapra coarse spatial scales prefer open grasslands (Hebblewhite and elk (Boldt & Ingold, ; Hebblewhite et al., ). et al., ; Massé & Côté, ; Laundré, ). Even at low elevations snow cover and depth during winter

TABLE 4 Habitat selection by migratory and sedentary radio-tracked guanacos in Tierra del Fuego from January  to July  (habitat χ ’ preference is shown by the significance of the test), with the number of locations used, Manly s standardized selection index (Bi; Manly et al., ), the available proportion of each vegetation type, and the proportion of vegetation type used (with % Bonferroni confidence interval).

Vegetation type No. of locations Bi Available proportion Proportion used (95% Bonferroni confidence limits)* Migratory, summer (χ2 = 19.643, df = 2, P , 0.001) Grassland/bush/steppe 8 0.65 0.038 0.140 (0.019–0.262) Meadow 21 0.22 0.294 0.368 (0.200–0.537) Deciduous forest 28 0.13 0.668 0.491 (0.316–0.666) Migratory, winter (χ2 = 15.956, df = 2, P , 0.001) Grassland/bush/steppe 20 0.17 0.504 0.435 (0.242–0.628) Meadow 9 0.66 0.058 0.196 (0.041–0.350) Deciduous forest 17 0.17 0.438 0.370 (0.182–0.557) Sedentary, in Karukinka in summer (χ2 = 2.697, df = 2, P = 0.260) Grassland/bush/steppe 20 0.40 0.416 0.513 Meadow 9 0.38 0.200 0.231 Deciduous forest 10 0.22 0.383 0.256 Sedentary, in Karukinka in winter (χ2 = 2.663, df = 2, P = 0.264) Grassland/bush/steppe 39 0.32 0.416 0.411 Meadow 21 0.36 0.200 0.221 Deciduous forest 35 0.31 0.383 0.368 Sedentary, outside Karukinka (χ2 = 3.942648, df = 2, P = 0.139) Grassland/bush/steppe 17 0.49 0.439 0.607 Meadow 7 0.33 0.270 0.250 Deciduous forest 4 0.18 0.291 0.143

*Vegetation type is used significantly less (value in bold) if the available proportion is larger than the upper limit of the Bonferroni confidence interval of proportion used (P , .).

affected guanaco habitat use; we often observed guanacos unpubl. data) and killing of guanaco  by poachers in feeding on sun-facing slopes where they were more likely . to find snow-free patches, as occurs with the Alpine chamois The decline of guanacos that we recorded may be a (Boldt & Ingold, ). Ortega & Franklin () suggested consequence of overgrazing by sheep as a result of decades that guanaco migration is related to snow accumulation in of poor range management. Range condition was already highland areas, which may cause a severe reduction of avail- poor on the ranches around Karukinka in the late sas able forage in winter and trigger migration. The same re- a consequence of overgrazing (Lara & Cruz, ). Between lationship between snow accumulation and migration  and  there was a high density of sheep and they appears to be common among other migratory mountain were moved little among paddocks in the largest ranch ungulates (Schoen & Kirchhoff, ; Boldt & Ingold, neighbouring Karukinka, leading to high mortality of ewes ; Hebblewhite et al., ; White et al., ). and lambs (C.A. Moraga, unpubl. data). In June  the The negative relationship between guanaco and sheep ranch manager was replaced and most remaining sheep densities at low elevations (Table ) confirms the effect of were removed but range condition was already poor. Poor sheep on guanacos (Raedeke, ; Baldi et al., ). At range management, with negative demographic effects on the landscape scale wild herbivores that coexist with live- sheep, explains the region-wide decline in range pro- stock tend to suffer density declines throughout the area ductivity and the collapse of the number of sheep in of overlap or concentrate in areas that are free from live- Patagonia (Aagesen, ; Texeira & Paruelo, ). stock, as in the case of chital Axis axis, gaur Bos gaurus, and Asian elephant Elephas maximus (Madhusudan, Seasonal movement patterns and habitat selection ). At the habitat scale, on the other hand, wild herbivores such as elk and mule deer also display strategies that Our study confirms the persistence of seasonal migration in include temporal avoidance of livestock (Stewart et al., the guanacos of Tierra del Fuego, and also that part of the ). Because of the regular patrols by rangers it was un- population is sedentary, as proposed by Raedeke (). This likely that guanacos suffered direct persecution by poachers mixed movement pattern has also been described for moose in Karukinka. Some level of persecution, however, occurred Alces alces and elk (Ball et al., ; Hebblewhite et al., ). outside Karukinka, as evidenced by the short flight- The home-range sizes of the migratory guanacos in distances observed on the ranches (C.A. Moraga et al., Karukinka were as small as in Torres del Paine National

Park during summer (Young & Franklin, ). Defence of Conservation implications territories by dominant males from family groups during the reproductive season, in addition to greater availability The interaction between guanacos and forests is a conten- of good-quality forage, may explain the reduced size of sum- tious issue in Tierra de Fuego (Martínez-Pastur et al., mer ranges (Ortega & Franklin, ). Similar to migrants, ; Skewes et al., ; Cavieres & Fajardo, ). Our sedentary guanacos in our study used larger areas in winter study provides the first quantitative information on the ef- compared to summer (C.A. Moraga, unpubl. data). The fect of sheep ranching on guanaco density and use of forest death by starvation in late winter of migrant , however, habitats, a key factor in the conflict between logging, ranch- suggests that an increase in its home-range size was ing, and wildlife in Tierra del Fuego. Based on Raedeke’s not sufficient to compensate for reduced winter forage () seasonal counts and our study, migratory guanacos where the range was shared with abundant sheep. that move to grassland-dominated lower elevations in au- Differences in forage quality and quantity between summer tumn and winter may face competition from livestock. and winter probably contributed to the differences in season- Migratory guanacos had higher mortality from starvation al home-range sizes of guanacos as they did for roe deer than sedentary guanacos (Raedeke, ), although this Capreolus capreolus and white-tailed deer (Mysterud, was not confirmed by studying marked individuals. ; Lesage et al., ). However, the timing and condition at death of migratory The five radio-tracked males in our study were all seden- guanaco , as well as the negative effect of sheep on gua- tary. Young & Franklin () suggested that males estab- naco numbers, may support Raedeke’s conclusion. If mi- lish and defend territories for as long as they can, which may gratory guanacos that use predominantly forested areas explain the sedentary behaviour of males that we captured such as Karukinka and properties devoted to logging in and radio-collared while they held family groups (Sarno summer face high nutritional stress during winter, then ad- et al., ), with the exception of male . This guanaco justments in levels and timing of livestock stocking rates in displayed the typical transition from family to bachelor neighbouring areas may be needed to recover the range and group described for juvenile guanacos in Torres del Paine also maintain seasonal use by guanacos, particularly in win- Park (Young & Franklin, ). Females, on the other ter. Ultimately, high sheep densities and range degradation hand, are expected to display lower site fidelity than on ranches appears to be detrimental to logging interests as males after the reproductive season (Jurgensen, ). well as to guanacos, as they exacerbate guanaco use of forest Accordingly, females  and  migrated seasonally and habitats. returned to previously occupied ranges the following year The Chilean government has authorized six harvests in but female  was sedentary throughout the year. Female the southern area of Tierra del Fuego since , with a  left the range used during the first year, although this total of c. , guanacos harvested. In the last  years probably represented expulsion from a family group and the logging company in the area has leased the harvest of dispersal into a new range (Sarno et al., ). Both gua- guanaco for sustainable use. Nonetheless, aiming only for nacos that appeared to have been expelled from their orig- sustainable use may not necessarily favour a reduction of inal family groups were classified as young adults when they forest browsing by guanaco. For example, hunting guanacos were captured and collared (Table ). during winter in the forest–grassland mosaic may not be All the guanacos radio-tracked in our study utilized the targeting the guanacos that establish range in the logging forest but none selected this habitat type over others. areas during the growing season. In addition, hunting of ea- Specifically, reduced utilization of forests in summer by mi- sier targets in open areas could encourage surviving gua- grants and by sedentary guanacos in Karukinka suggests nacos to move inside forests, as human harassment can that forests were not a high-quality habitat for guanacos modify guanaco behaviour and increase their use of mar- during the reproductive season. Conversely, meadows ginal habitats (Donadio & Buskirk, ). Hence, current appeared to be important habitats for migratory guanacos guanaco management by loggers may increase forest use, during winter, a preference reported for guanacos in areas and may have unexpected negative effects on guanacos of the high Andes and southern Patagonia during summer and on forests unless it takes into consideration the spatial (Bank et al., ; Puig et al., ). Meadows in Tierra ecology and habitat requirements of the guanaco. del Fuego are highly productive habitats that are important The guanaco–forest interaction in Tierra del Fuego has for sheep during summer (Cingolani et al., ). As been studied exclusively at a local, patch-level scale we observed, meadows were not heavily utilized by (Martínez-Pastur et al., ; Pulido et al., ; Cavieres sheep in winter, perhaps as a result of the high humidity & Fajardo, ). In particular, the intensive use by gua- and the formation of ponds. The concentration of sheep nacos of recently logged forest patches affects the logging in- in grasslands during winter may have forced guanacos to dustry because browsing significantly slows tree growth and use the wetter meadows as a second-best choice after reduces seedling density (Martínez-Pastur et al., ; grasslands. Pulido et al., ; Riveros, ; Collado et al., ).

Interactions between ungulates and forests that are relevant BALDI, R., ALBON, S.D. & ELSTON, D.A. () Guanacos and sheep: for forest dynamics, however, must be assessed and managed evidence for continuing competition in arid Patagonia. Oecologia,  – at both local and landscape scales, as suggested for white- , .  BALDI, R., PELLIZA-SBRILLER, A., ELSTON, D.A. & ALBON, S.D. tailed deer in North America (Wisdom et al., ; () High potential for competition between guanacos and Tremblay et al., ; Rutherford & Schmitz, ). Our sheep in Patagonia. The Journal of Wildlife Management, , data provide insight into the interactions among guanacos, –. forests and sheep ranching, and can be used to guide the as- BALL, J.P., NORDENGREN,C.&WALLIN,K.() Partial migration by sessment of the effects of guanacos at the landscape scale and large ungulates: characteristics of seasonal moose Alces alces ranges in northern Sweden. Wildlife Biology, , –. the management and monitoring of guanaco populations. BANK, M.S., SARNO, R.J. & FRANKLIN, W.L. () Spatial distribution The Chilean authorities hope that the sale of guanaco of guanaco mating sites in southern Chile: conservation meat from legal harvests will be viewed as a contribution implications. Biological Conservation, , –. to the economy of the Magallanes region. This value may BERGER,J.() The last mile: how to sustain long-distance  – not be perceived, however, by most of the land owners migration in mammals. Conservation Biology, , . BOLDT,A.&INGOLD,P.() Effects of air traffic, snow cover and that sustain the guanaco population. Tierra del Fuego ran- weather on altitudinal short-term and medium-term movements of chers have shown little interest in sustainably managing female Alpine chamois Rupicapra rupicapra in winter. Wildlife guanacos and complaints from the forestry sector have Biology, , –. not declined since the legal harvest was initiated. Clearly, in- BOLGER, D.T., NEWMARK, W.D., MORRISON, T.A. & DOAK, D.F.  tegrated management of grasslands, forests and wildlife is ( ) The need for integrative approaches to understand and conserve migratory ungulates. Ecology Letters, , –. needed and it can only be successful if it is implemented BRIDGES, E.L. () Uttermost Part of the Earth. Holder and at the landscape scale. Integrated management in this Stoughton, London, UK. case requires combining knowledge of forest dynamics, gua- BUCKLAND, S.T., ANDERSON, D.R., BURNHAM, K.P., LAAKE, J.L., naco spatial and trophic requirements, and livestock and BORCHERS, D.L. & THOMAS,L.() Introduction to Distance range management, and the interactions among these. Sampling: Estimating Abundance of Biological Populations, nd Harvesting guanacos may be a useful tool to promote gua- edition. Oxford University Press, New York, USA. CAVIERES, L.A. & FAJARDO,A.() Browsing by guanaco (Lama naco conservation in parts of Tierra del Fuego but it should guanicoe)onNothofagus pumilio forest gaps in Tierra del Fuego, be part of an integrated management strategy and it must Chile. Forest Ecology and Management, , –. allow maintenance of key ecological processes such as sea- CINGOLANI, A.M., ANCHORENA,J.&COLLANTES, M.B. () sonal migration between forests and grasslands. Ultimately, Landscape heterogeneity and long-term animal production in  – conservation of Fuegian ecosystems depends on reconciling Tierra del Fuego. Journal of Range Management, , . COLLADO, L., FARINA, S., JARAS,F.&VARGAS,H.() Monitoreo the interests of livestock husbandry, guanaco conservation del estado de intervención y de la regeneración de Nothofagus and the timber industry. pumilio en un plan de manejo forestal en el ecotono estepa-bosque de Tierra del Fuego, Argentina. Bosque, , –. CONAMA () Ficha de antecedentes de especie Lama guanicoe   Acknowledgements (Miller, ). Reglamento de Clasificación de Especies Silvestres, to proceso de clasificación. Ministerio del Medio Ambiente, Gobierno de  We thank M. Uhart and C. Marull from the Global Health Chile. Http://www.mma.gob.cl/clasificacionespecies/ficha proceso/ fichas_actualizadas/Lama_guanicoe_PR-_RCE.doc [accessed Program of the Wildlife Conservation Society for their help  March ]. in capturing guanacos, and B. Saavedra, R. Muza, DE LA TOUR, G.D. () The guanaco. Oryx, , –. J. Sotomayor, M. Chacón, C. Silva, D. Droguett, DOBSON, A.J. () An Introduction to Generalized Linear Models, F. Repetto, M. Millán, S. Lorca, J. Gómez, A. Muñoz, nd edition. Chapman & Hall/CRC, Boca Raton, USA.  A. González, students and Karukinka staff for support. DODDS,P.( ) Efecto del ramoneo de guanacos (Lama guanicoe)sobre We also thank W. Franklin and an anonymous reviewer la regeneración de lenga (Nothofagus pumilio) en Russffin, Tierra del   Fuego. Forestry thesis. Universidad de Chile, Santiago, Chile. for their comments. Permits No. and were obtained DONADIO,E.&BUSKIRK, S.W. () Flight behavior in guanacos and from the Servicio Agrícola y Ganadero–Government of vicuñas in areas with and without poaching in western Argentina. Chile for guanaco captures, and protocols of the Global Biological Conservation, , –. Health Program of the Wildlife Conservation Society were DONOSO,C.&LARA,A.() Utilización de los bosques nativos en followed for animal handling. Funding was provided by Chile: pasado, presente y futuro. In Ecología de los bosques nativos de Chile (eds J.J. Armesto, C. Villagrán & M. Kalin Arroyo), pp. – the Wildlife Conservation Society. . Editorial Universitaria, Santiago, Chile. FRANKLIN, W.L. () Contrasting socioecologies of South America’s wild camelids: the vicuña and the guanaco. In Advances in the References Study of Mammalian Behavior (eds J.F. Eisenberg & D. Kleiman), pp. –. American Society of Mammalogists, Shippensburg, AAGESEN,D.() Crisis and conservation at the end of the world: USA. sheep ranching in Argentine Patagonia. Environmental GORDON, I.J. () What is the future for wild, large herbivores in Conservation, , –. human-modified agricultural landscapes? Wildlife Biology, , –.

GRAHAM, R.T., JAIN, T.B. & KINGERY, J.L. () Ameliorating PUIG, S., ROSI, M.I., VIDELA,F.&MENDEZ,E.() Summer and conflicts among deer, elk, cattle and/or other ungulates and other winter diet of the guanaco and food availability for a High Andean forest uses: a synthesis. Forestry, , –. migratory population (Mendoza, Argentina). Mammalian Biology, HARRIS, G., THIRGOOD, S., HOPCRAFT, J.G.C., CROMSIGT, J.P.G.M. & , –. BERGER,J.() Global decline in aggregated migrations of large PULIDO, F.J., DÍAZ,B.&MARTÍNEZ-PASTUR,G.() Incidencia del terrestrial mammals. Endangered Species Research, , –. ramoneo del guanaco (Lama guanicoe Muller) sobre la regeneración HEBBLEWHITE, M., MERRILL,E.&MCDERMID,G.()A temprana en bosques de lenga (Nothofagus pumilio (Poepp et Endl) multi-scale test of the forage maturation hypothesis in a partially Krasser) de Tierra del Fuego Argentina. Investigación Agraria de migratory ungulate population. Ecological Monographs, , –. Sistemas y Recursos Forestales, , –. HOOGE,P.N.&EICHENLAUB,B.() Animal Movement Extension RAEDEKE,K.() Population dynamics and socio-ecology of the to ArcView. Version .. Alaska Science Center, Biological Science guanaco (Lama guanicoe) of Magallanes, Chile. PhD thesis. Office, U.S. Geological Survey, Anchorage, USA. University of Washington, Seattle, USA. HUBER, U.M. & MARKGRAF,V.() European impact on fire RAEDEKE,K.() Habitat use by guanacos (Lama guanicoe) and regimes and vegetation dynamics at the steppe-forest ecotone of sheep on common range, Tierra del Fuego, Chile. Turrialba, , – southern Patagonia. The Holocene, , –. . JURGENSEN, T.E. () Seasonal territoriality in a migratory guanaco REBERTUS, A.J., KITZBERGER, T., VEBLEN,T.T.&ROOVERS, L.M. population. MSc thesis. Iowa State University, Ames, USA. () Blowdown history and landscape patterns in the Andes of KARESH, W.B., UHART, M.M., DIERENFELD, E.S., BRASELTON, W.E., Tierra del Fuego, Argentina. Ecology, , –. TORRES, A., HOUSE, C. et al. () Health evaluation of RIVEROS, J.C. () Efecto del cerco de exclusión de guanacos (Lama free-ranging guanaco (Lama guanicoe). Journal of Zoo and Wildlife guanicoe Müller) sobre el ramoneo en la regeneración de lenga Medicine, , –. (Nothofagus pumilio (Poepp. et Endl.)),  años después de su LARA,A.&CRUZ,G.() Vegetación del área de uso agropecuario de establecimiento en Russfin, Tierra del Fuego. Memoria de Ingeniería la XII Región, Magallanes y la Antártica Chilena. INIA (Instituto de Forestal. Universidad de Chile, Santiago, Chile. Investigaciones Agropecuarias), Santiago, Chile. RUTHERFORD, A.C. & SCHMITZ, O.J. () Regional-scale assessment LARSON, M.A. () A catalog of software to analyze radiotelemetry of deer impacts on vegetation within Western Connecticut, USA. data. In Radio Tracking and Animal Populations (eds J.J. Millspaugh The Journal of Wildlife Management, , –. & J.M. Marzluff), pp. –. Academic Press, San Diego, USA. SAAVEDRA, B., SIMONETTI, J.A. & REDFORD, K.H. () Private LAUNDRÉ, J.W. () Behavioral response races, predator–prey shell conservation, the example that the Wildlife Conservation games, ecology of fear, and patch use of pumas and their ungulate Society builds from Tierra del Fuego. In Biodiversity Conservation prey. Ecology, , –. in the Americas: Lessons and Policy Recommendations LESAGE, L., CRÊTE, M., HUOT, J., DUMONT,A.&OUELLET, J.-P. () (ed. E.B. Figueroa), pp. –. Editorial FEN–Universidad de Seasonal home range size and philopatry in two northern white- Chile, Santiago, Chile. tailed deer populations. Canadian Journal of Zoology, , –. SARNO, R.J., BANK, M.S., STERN, H.S. & FRANKLIN, W.L. () MADHUSUDAN, M.D. () Recovery of wild large herbivores Forced dispersal of juvenile guanacos (Lama guanicoe): causes, following livestock decline in a tropical Indian wildlife reserve. variation, and fates of individuals dispersing at different times. Journal of Applied Ecology, , –. Behavioral Ecology and Sociobiology, , –. MANLY, B.F.J., MCDONALD, L.L., THOMAS, D.L., MCDONALD, T.L. & SARNO, R.J., FRANKLIN, W.L., O’BRIEN, S.J. & JOHNSON, W.E. () ERICKSON,W.P.() Resource Selection by Animals: Statistical Patterns of mtDNA and microsatellite variation in an island and Design and Analysis for Field Studies, nd edition. Kluwer, Boston, mainland population of guanacos in southern Chile. Animal USA. Conservation, , –. MARTÍNEZ-PASTUR, G., PERI, P.L., FERNÁNDEZ, M.C., STAFFIERI,G. SCHOEN, J.W. & KIRCHHOFF, M.D. () Seasonal distribution and &RODRÍGUEZ,D.() Desarrollo de la regeneración a lo largo del home-range patterns of sitka black-tailed deer on Admiralty ciclo del manejo forestal de un bosque de Nothofagus pumilo: Island, southeast Alaska. The Journal of Wildlife Management, , . Incidencia del ramoneo de Lama guanicoe. Bosque, , –. –. MARTINIC,M.() Expansión colonizadora. In Historia de la Región SCHOEN, J.W. & KIRCHHOFF, M.D. () Seasonal habitat use by sitka Magallánica, Volume I (ed. M. Martinic), pp. –. Editorial black-tailed deer on Admiralty Island, Alaska. The Journal of Universidad de Magallanes, Punta Arenas, Chile. Wildlife Management, , –. MASSÉ,A.&CÔTÉ, S.D. () Habitat selection of a large herbivore SKEWES, O., GONZALEZ, F., OVALLE, C., RUBILAR, L., QUEZADA., M., at high density and without predation: trade-off between forage and JIMÉNEZ,A.etal.() Manejo Productivo y Sustentable del cover? Journal of Mammalogy, , –. Guanaco en Isla de Tierra del Fuego. Informe Final (Etapas II y III) MYSTERUD,A.() Seasonal migration pattern and home range of Universidad de Concepción, Servicio de Gobierno Regional XII roe deer (Capreolus capreoulus) in an altitudinal gradient in Región, Magallanes y Antártica Chilena, Chillán, Chile. southern Norway. Journal of Zoology, , –. STEWART, K.M., BOWYER, R.T., KIE, J.G., CIMON, N.J. & JOHNSON,B. ORTEGA, I.M. & FRANKLIN, W.L. () Social organization, K. () Temporospatial distributions of elk, mule deer, and cattle: distribution and movements of a migratory guanaco population in resource partitioning and competitive displacement. Journal of the Chilean Patagonia. Revista Chilena de Historia Natural, , Mammalogy, , –. –. TEXEIRA,M.&PARUELO, J.M. () Demography, population PÉPIN, D., ADRADOS, C., JANEAU, G., JOACHIM,J.&MANN,C.() dynamics and sustainability of the Patagonian sheep flocks. Individual variation in migratory and exploratory movements and Agricultural Systems, , –. habitat use by adult red deer (Cervus elaphus L.) in a mountainous THOMAS, L., LAAKE, J.L., REXSTAD, E., STRINDBERG, S., MARQUES,F. temperate forest. Ecological Research, , –. F.C., BUCKLAND, S.T. et al. () Distance .. Release . Research PISANO,E.() Fitogeografía de Fuego-Patagonia chilena. I: Unit for Wildlife Population Assessment, University of St Andrews, Comunidades vegetales entre las latitudes °y°S.Anales del UK. Http://www.ruwpa.st-and.ac.uk/distance/ [accessed  August Instituto de la Patagonia, , –. ].

TREMBLAY, J.-P., HUOT,J.&POTVIN,F.() Density-related effects YOUNG, J.K. & FRANKLIN, W.L. () Territorial fidelity of male of deer browsing on the regeneration dynamics of boreal forests. guanacos in the Patagonia of southern Chile. Journal of Journal of Applied Ecology, , –. Mammalogy, , –. TUHKANEN, S., KUOKKA, I., HYVÖNEN, S., STENROOS,S.&NIEMELÄ, J. () Tierra del Fuego as a target for biogeographical research in the past and present. Anales del Instituto de la Patagonia, , Biographical sketches –. WCS (WILDLIFE CONSERVATION SOCIETY)() Land cover map C LAUDIO M ORAGA’s research interests are wildlife ecology and inte- of Tierra del Fuego, Global Land Cover Facility Landsat  Dataset. grative conservation. He is currently studying guanaco–ranching inter- Wildlife Conservation Society, Living Landscapes Program and actions in Southern Patagonia. MARTÍN FUNES’s research focuses on Chile Program, Santiago, Chile. carnivore and raptor ecology, exotic species ecology, monitoring tech- WHITE, P. J., PROFFITT, K.M., MECH, L.D., EVANS, S.B., niques, and threats to wildlife in Patagonia. C RISTÓBAL P IZARRO’s CUNNINGHAM, J.A. & HAMLIN, K.L. () Migration of northern research focuses on human dimensions of wildlife conservation and Yellowstone elk: implications of spatial structuring. Journal of environmental education. He is currently studying human–bird rela- Mammalogy, , –. tionships in North America. C RISTÓBAL B RICEÑO is interested in WISDOM, M.J., VAVRA, M., BOYD, J.M., HEMSTROM, M.A., AGER,A. the conservation and health of threatened wildlife and their habitats, A. & JOHNSON, B.K. () Understanding ungulate herbivory— especially those confined to islands. A NDRÉS N OVARO’s research fo- episodic disturbance effects on vegetation dynamics: knowledge cuses on the impacts of hunting on wildlife and predator–prey interac- gaps and management needs. Wildlife Society Bulletin, , tions. He works with governments and the private sector to address –. threats from extractive industries, livestock grazing and poaching.