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Studies on Neotropical Fauna and Environment Vol. 43, No.

1, April 2008, 19

ORIGINAL ARTICLE Habitat use by guanacos (Lama guanicoe, Camelidae) in northern Patagonia (Mendoza, Argentina)
Silvia Puig*, Fernando Videla, Monica I. Cona and Virgilio G. Roig
Unidad Ecologa Animal, Instituto Argentino de Investigaciones en Zonas Aridas (IADIZA, CONICET), Mendoza, Argentina (Received 15 February 2005; accepted 9 May 2007) Strategies of habitat use of the guanaco were analyzed in early and late summer, when parturition and lactation increase nutritional needs and predation risk. The daily distribution of guanacos was followed by scan-sampling, relief and soil were measured in each habitat, and vegetation was analyzed by the point-quadrat method. Guanacos showed a selective habitat use, preferring open areas with grasslands, gentle terrain and low hiding cover. The density of guanacos was positively associated with the intensively eaten grass Panicum urvilleanum, and negatively associated with rocky soils, high shrublands and avoided plants. A weakening of the selective habitat use occurred when the abundance of guanacos increased in the study area during late summer, ceasing avoidance of poor and risky habitats. Keywords: arid environments; habitat selection; spatial patterns; South American camelids; ungulates

Introduction Spatial patterns in topography and forage distribution frequently influence habitat selection by large herbivores. Abiotic factors such as slope or distance to water are considered primary determinants of herbivore distribution, and constrain selection based on biotic factors such as plant structure or productivity (Senft et al., 1987; Bailey et al., 1996). Habitat use by ungulates may reflect different foraging decisions at different spatial scales (Kotliar & Wiens, 1990). The spatial hierarchy developed by Senft et al. (1987) for large herbivores can be summarized as a selection of home ranges within a region, selection of feeding areas within a landscape, and selection of food items within a plant community. Habitat decisions of ungulates may operate based on minimizing the risk of predation at a large scale, and on maximizing energy intake and nutrient balance at a small spatial scale, as was observed in the mule deer (Belovsky & Schmitz, 1994; Kie, 1999; McShea et al., 2001). Environmental factors noninteractive with herbivores (such as climate, water availability and physical features) would be expected to become more important in the decision-making process at a large scale (e.g. a region), as geomorphological and climatic factors are determinant of environmental heterogeneity (Senft et al., 1987). On the other hand, interactive factors such as plant biomass and structure should dominate habitat selection at a small scale (e.g. a micropatch), together with patch size and aggregation (Kotliar & Wiens,

1990; Stuth, 1991). A more selective habitat use is to be expected during the reproductive season, considering that parturition and lactation usually increase nutritional needs and predation risk in ungulates (Berger, 1991; Barten et al., 2001). The wide geographical range of the guanaco (Lama guanicoe Muller, 1776) includes a great diversity of environments. The most abundant populations are located in Patagonia (Wheeler, 1991; Puig, 1995). Open habitats prevail (Raedeke, 1982; Cajal, 1989) and mountain wetlands (vegas), where available, are preferred by guanacos (Jurgensen, personal communication; Lawrence, personal communication). Low competition with livestock, high food availability and presence of escape terrain are important habitat attributes for guanacos (del Valle et al., 1997). A negative correlation between abundance of guanacos and livestock was found in Central and Northern Patagonia (Puig et al., 1997; Baldi et al., 2001). The guanaco population of La Payunia Reserve stands out as one of the most abundant in a protected area (Torres, 1992). The region of La Payunia constitutes the northernmost District of the Patagonian Biogeographic Province. It is of interest to evaluate habitat preferences by guanacos in this region where vegas are absent: the availability of habitat and food for guanacos showed heterogeneity at several spatial scales, conditioned by an arid climate and a strong volcanic activity. The distribution of guanacos in the region was associated mainly

*Corresponding author. S. Puig, IADIZA, CC. 507, 5500, Mendoza, Argentina. Fax: +54-261-5244001. Email: spuig@lab.cricyt.edu.ar
ISSN 0165-0521 print/ISSN 1744-5140 online # 2008 Taylor & Francis DOI: 10.1080/01650520701461319 http://www.informaworld.com

S. Puig et al. their subdivision into geomorphological units, were identified within La Payunia according to recurrent patterns of relief, soil and vegetation, based on 1:50 000 aerial photography, geological cartography (Gonzalez Daz, 1972) and plant cartography (Martnez & Dalmasso, 1993). Seasonal availability and use of plant species eaten by guanacos in different landscapes of La Payunia are described in Puig et al. (1996, 1997). The study area (15 km2) was located in Zaino, a landscape of La Payunia Reserve representative to the guanaco habitat in Northern Patagonia with availability of good observation points, long distance visibility, a diversity of geomorphological units, a high abundance of guanacos, low human impact, and very few livestock. Poaching impacts can be discarded because of the presence of game wardens in La Payunia protected area during the present study. Five geomorphological units, potential feeding areas for guanacos, were identified in the selected study area (Table I): a wide sandy plain, an extensive piedmont, a hillside, a basaltic scarp, and an extensive plateau. Field design Daily activities of guanacos in early and late summer (December 1994 and February 1995) were followed from the top of the Zaino hill with the aid of a 20240680 Galileo Triog telescope. Observations were carried out from 09:00 to 19:00 h using scan samples (Altmann, 1973), recording every hour the number of guanacos per geomorphological unit. Total number of guanacos observed in all five units on each occasion was considered an estimator of their abundance in the study area. The area occupied by each geomorphological unit within the study area was estimated from a satellite image. Vegetation, relief and soil were analyzed at three sampling sites in each geomorphological unit (two in the basaltic scarp unit because of its narrow extent). Ten 30-m transects

with vegetation abundance and composition, presence of livestock (Videla & Puig, 1993; Puig et al., 1997) and illegal hunting (Puig, 1986). The puma (Puma concolor), being the main predator of South American camelids (Sarno et al., 1999), also may affect habitat selection by guanacos in La Payunia. This paper analyses the strategies of guanacos for habitat use in Northern Patagonia during the reproductive season, at the landscape scale. The objectives are: (1) to test selectivity in habitat use through the density of guanacos in the geomorphological units, as potential feeding areas; (2) to evaluate whether changes in the total number of guanacos in the study area affect the selective use of geomorphological units; and (3) to identify the relative importance of environmental variables in habitat selection by guanacos, such as: (a) type of relief and soil, (b) plant cover and diversity, (c) availability of preferred food items, and (d) hiding cover.

Materials and methods Study area and habitat characteristics The study was conducted in La Payunia Reserve (Mendoza, Argentina, 36u109S, 68u509W, 2500 km2), an environment representative of the northernmost District of the Patagonian Biogeographic Province. The xerophyllous vegetation is a Patagonian shrub steppe (Cabrera & Willink, 1980), and the climate is a continental desert type (Consejo Federal de Inversiones, 1977). Mean seasonal temperature ranges from 6uC in winter to 20uC in summer, and annual precipitation averages 255 mm (Puig et al., 1996). The zone presents signs of strong volcanic activity and of wind and water erosion. The relief consists of gentle slopes and large plains, interrupted by basaltic scarps and hill groups, where soil and vegetation have been influenced by geomorphological processes (Gonzalez Daz, 1972). Landscapes, and

Table I. Mean (SD) of relief features, soil types and dominant plant communities in the five geomorphological units.
Units* Hillside (20%) Piedmont (22%) Plain (45%) Basaltic scarp (5%) Plateau (8%) Elevation (m) Slope (degrees) 157122 154531 151724 148214 15008 134 31 32 93 12 Soil type Rocky Rocky to sandyrocky Sandy Volcanic rocky Sandy-rocky to sandy Dominant plant community Stipa spp. grassland with dense high Neosparton aphyllum or low Mulinum spinosum shrubland Dense high Larrea nitida shrubland to Panicum urvilleanum grassland with open high Neosparton aphyllum shrubland Panicum urvilleanum and Poa spp. grassland with open low Ephedra ochreata shrubland Dense very low Acantholippia seriphioides shrubland with Panicum urvilleanum grassland Panicum urvilleanum grassland with a high Anarthrophyllum rigidum shrubland

*Proportions of surface in the study area are in parentheses.

Studies on Neotropical Fauna and Environment were distributed randomly on each sampling site, separated from one another by more than 100 m. Altitude, slope, type of soil and height of plant species were determined for each transect, and plant cover, proportion of layers, diversity, and abundance of plant species were estimated in early and late summer using the point-quadrat method (Daget & Poissonet, 1971). Throughout the study period, 88 scan samples were recorded, and 280 transects were traversed along the study area. Statistical analyses Densities of guanacos were log-transformed to normalize their distribution and were compared between dates and among hours with the Fisher analysis of variance for repeated measures, using the Tukey test for multiple comparisons (Zar, 1984). Significant differences between the number of guanacos observed and expected in each geomorphological unit were detected by the Chi-square test (Zar, 1984). Expected numbers were calculated from the proportion of study area occupied by each unit. Preference or avoidance of geomorphological units by guanacos was detected using Bailey confidence intervals based on Bonferroni inequality (Cherry, 1996). Differences in soil, relief and vegetation among geomorphological units and between dates were detected by MANOVA with absolute variables logtransformed and proportions transformed by arcsine of the square root. The ShannonWiener function (H9; Colwell & Futuyma, 1971) was used to estimate plant diversity. Three plant layers were distinguished: a herbaceous layer (mostly composed of grasses, occasionally forbs), a low shrub layer (chamaephytes and phanerophytes,1 m tall), and a high shrub layer (phanerophytes.1 m tall). Preferred (Poa lanuginosa, Panicum urvilleanum, Ephedra ochreata) and avoided (Stipa spp.) plant species, as well as tall phanerophytes which constitute potential hiding cover (Berberis grevilleana, Larrea spp., Anarthrophyllum rigidum, Neosparton aphylum) were included in the analysis of variance. A factor analysis procedure using principal component extraction (Hair et al., 1999) was applied in order to detect relationships among environmental variables and density of guanacos.

and 712 individuals (SD5188) in late summer. The two-factor ANOVA including repeated measures for the log-transformed density of guanacos detected significant differences between early and late summer (F539.13, P,0.001), while differences among hours of observation were not significant (F50.23, P50.993). The temporal differences corresponded to an increase in the density of guanacos from early to late summer, particularly in the piedmont, basaltic scarp and plateau units (Figure 1). Guanacos showed a selective use of geomorphological units in early summer (x25192.19, P,0.001) as well as in late summer (x25211.79, P,0.001). According to Bailey (1980) confidence intervals, guanacos preferred the plain unit, made a proportional use of the plateau unit, and avoided the hillside unit throughout the summer. The avoidance for the piedmont and basaltic scarp units in early summer shifted to a proportional use in late summer (Figure 2). Environmental variables in the geomorphological units Significant differences among geomorphological units were found in abiotic and biotic factors noninteractive with herbivores (features of relief and soil, vegetation structure and hiding cover, Wilks lambda50.06, F529.85, P,0.001). Biotic factors potentially interactive with herbivores also differed among units, such as the availability of plant species eaten with preference and avoidance by guanacos (Wilks lambda50.06, F540.213, P,0.001). The altitude differed among all units (Table I) in a slight gradient from northeast (1750 m) to southwest (1400 m), without relation to habitat preferences by guanacos. The slope and the proportion of rocks in soil was significantly higher in the hillside and basaltic scarp units, which were avoided by guanacos (hillside throughout the summer and basaltic scarp during early summer). The vegetation structure differed among units (Table II), as the proportion of the herbaceous layer was higher in the plain and plateau (units intensively used by guanacos), the proportion of the high shrub layer was higher in the piedmont (unit proportionally used), and the proportion of the low shrub layer was higher in the hillside and basaltic scarp (avoided units). The herbaceous layer mostly reflected differences in the proportion of grasses, as the proportion of forbs was very low throughout the study area. The plant diversity was higher in piedmont than in the other geomorphological units. The relative proportion of vegetation occupied by high phanerophytes, considered as hiding cover, was significantly higher in units proportionally used as

Results Abundance of guanacos and selective use of geomorphological units The abundance of guanacos in the study area averaged 370 individuals (SD588) in early summer,

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Figure 1. Density of guanacos in the geomorphological units in early and late summer. Confidence intervals were estimated for P50.05.

piedmont (represented by B. grevilleana and Larrea spp.) and plateau (A. rigidum). The availability of plant species selectively eaten by guanacos differed among geomorphological units. The preferred grass Poa lanuginosa showed a higher

availability in the plain (unit used with preference by guanacos) than in the other units. The avoided grass Stipa spp. presented a higher proportion in the hillside (unit used with avoidance), while P. lanuginosa and Panicum urvilleanum, other grasses

Figure 2. Availability proportion of geomorphological units and proportion of use by guanacos in early and late summer. Confidence intervals (CI) for the proportions of use were estimated using the Bailey formula. HI, hillside; PI, piedmont; SP sandy plain; BS, basaltic scarp; EP, extensive plateau.

Studies on Neotropical Fauna and Environment

Table II. Seasonal differences in mean (SD) plant cover, diversity and proportions of vegetation (grouped by layers and by categories) during early summer, and in parentheses during late summer.
Variable Plant cover (%) Plant diversity (H9) Herbaceous layer Low shrub layer High shrub layer Grasses Forbs Chamaephytes Phanerophytes Hillside 6911 (6310) 0.400.14 (0.390.12) 0.750.13 (0.760.11) 0.210.13 (0.190.11) 0.040.10 (0.050.09) 0.750.13 (0.760.11) 0.0040.010 0.050.07 (0.050.07) 0.200.13 (0.190.12) Piedmont 7813 (7118) 0.550.14 (0.510.17) 0.760.15 (0.770.18) 0.140.13 (0.080.09) 0.090.10 (0.160.16) 0.760.15 (0.770.18) 0.0020.004 0.000.01 0.230.14 (0.230.18) Plain 7619 (7517) 0.290.16 (0.300.15) 0.880.19 (0.930.10) 0.070.15 (0.050.08) 0.050.13 (0.030.07) 0.870.19 (0.930.10) 0.0020.005 0.010.01 0.120.19 (0.070.10) Basaltic scarp 6921 (6519) 0.320.15 (0.380.14) 0.800.18 (0.730.19) 0.170.20 (0.230.22) 0.020.06 (0.040.08) 0.800.18 (0.730.19) 0.0010.004 (0.0010.003) 0.110.16 (0.130.16) 0.090.09 (0.140.14) Plateau 8010 (7812) 0.340.18 (0.360.17) 0.930.12 (0.940.09) 0.010.03 (0.010.03) 0.060.12 (0.050.08) 0.930.12 (0.940.09) 0.0020.007 (0.0010.004) 0.070.12 (0.060.09)

intensively used by guanacos, showed lower availability. The preferred chamaephyte E. ochreata presented higher availability in piedmont in early summer, when this unit was proportionally used by guanacos. The availability of vegetation in the study area showed a significant decrease from early to late summer (Wilks lambda50.93, F52.192, P50.023). Plant cover particularly decreased in the hillside and the proportion of forbs (food item always preferred by guanacos) particularly decreased in the hillside and piedmont. Environmental variables and habitat selection by guanacos The principal component analysis explained 94.14% of the total variance in early summer and 93.97% in late summer (Table III). The density of guanacos and the proportion of P. urvilleanum (grass intensively eaten) showed a negative association with the first component of the rotated matrix for both dates. The altitude, proportion of rocks in soil, proportions of avoided or uneaten species (Stipa spp., Senecio filaginoides, Mulinum spinosum and Schinus spp.) presented a positive association with this component. The density of guanacos and the proportions of the preferred grasses Poa lanuginosa and Carex patagonum showed a negative association with the second component in early summer, when slope, proportion of rocks in soil and proportion of the avoided phanerophyte N. aphyllum were positively associated with this component. Proportions of P. lanuginosa, C. patagonum and

E. ochreata presented a positive association with the second component only in late summer. The third component only showed a positive association with the preferred chamaephyte E. ochreata in early summer and with slope and proportion of N. aphyllum in late summer. Discussion Abundance of guanacos and habitat selection A selective habitat use was detected during early and late summer, as the distribution of guanacos among the geomorphological units was significantly different from that expected by chance. The intensity of use by guanacos seems to be stable along the day, as their density in each geomorphological unit did not change among hours of observation. The use of geomorphological units by guanacos seems to be focused to a selection among potential feeding areas, as more than 70% of guanacos in the study area were observed eating. The selective use of geomorphological units persisted from early to late summer, despite a significant increase in the number of guanacos that shared the study area, and a general decrease in plant cover and proportion of the preferred forbs. This decline in food availability and quality occurred mainly in the hillside, and justified that this unit was used with avoidance throughout summer. Plant cover did not decline despite the increased pressure of herbivory in the units most intensively used, the plain and plateau, where food availability could sustain throughout the summer the respective preference and proportional use of these feeding areas.

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Table III. Loading factors of the density of guanacos and habitat features in the principal components (Comp) of the rotated matrix.
Early summer Variables Density of guanacos Altitude Slope Rocks in soil Panicum urvilleanum Poa lanuginosa Carex patagonum Stipa spp. Senecio filaginoides Mulinum spinosum Ephedra ochreata Neosparton aphylum Schinus spp. Proportion of variance explained Comp 1 20.55 0.91 0.45 0.76 20.90 20.40 20.02 0.91 0.92 0.94 0.01 0.31 0.92 50.93 Comp 2 20.79 0.01 0.69 0.65 20.39 20.80 20.83 0.34 0.32 0.25 0.02 0.81 0.32 30.23 Comp 3 0.22 0.41 20.29 0.02 20.17 0.34 20.14 0.17 20.21 20.21 0.99 0.40 20.21 12.98 Comp 1 20.89 0.97 0.45 0.77 20.91 20.35 20.13 0.97 0.89 0.91 20.16 0.03 0.94 53.25 Late summer Comp 2 0.28 0.14 20.23 20.44 0.34 0.93 0.84 20.21 20.29 20.28 0.87 20.25 20.17 23.81 Comp 3 20.28 20.06 0.83 0.44 20.19 20.10 20.17 0.06 0.19 0.19 20.35 0.94 0.23 16.91

Nevertheless, a weakening of the selective habitat use seems to derive from the increase of guanacos in the study area, as the use of two units with important hiding cover, piedmont and basaltic scarp, shifted from avoidance in early summer to proportional use in late summer. A stronger intra-specific competition is expected from the increase in the abundance of guanacos combined with the decline in food availability, and could explain shifts in habitat selectivity. Indeed, expected consequences of increasing competition in ungulate species are a decline in their body condition (Kie & Bowyer, 1999), and a more risky foraging strategy with a higher use of habitats previously avoided (Scogland & Grovan, 1991; Kie, 1999). Environmental variables and habitat selection by guanacos Differences of altitude among geomorphological units seem to be too slight to affect habitat preferences of guanacos, although this environmental factor could influence the preferences of guanacos at a higher spatial scale. Pronounced slopes and rocky soils, factors that characterized the avoided geomorphological units, constitute physical obstructions for near visibility and movements, and could increase the risk of predation for guanacos. Visual obstructions enhance the habitat quality for ungulates with hider-type neonates such as the white-tailed deer (Bello et al., 2001). On the contrary, these obstructions would worsen habitat quality for ungulates with follower-type neonates affected by ambushing predators, as occurred in a Puna environment, where guanacos and vicunas were

predated by pumas mainly in gorges (Cajal & Lopez, 1987). The avoidance of ravines and steep slopes in a Subantarctic population of guanacos also was related to difficulties in territory control by family males, and in detection of predators by females with juveniles (Ortega & Franklin, 1995). Ravines would become advantageous for guanaco females with juveniles as hiding cover from humans, in populations affected by poaching (Franklin, 1983). Dense shrublands also constitute hiding sites for predators and are avoided by ungulates, as was observed in antelopes (Underwood, 1982; Kie, 1999). High plant cover and diversity in the geomorphological units with dense shrublands, such as the plateau and piedmont, could account for a proportional use of these units, and justify that shrublands were less avoided than physical visual obstructions. Synthetic variables of vegetation, such as plant cover, structure and diversity, seem to have relative low relevance in the habitat selection by guanacos, at least in summer. Such variables could be more important for the selection of feeding areas by guanacos in other seasons, when the food availability for herbivores becomes scarcer. A high availability of plant species eaten with preference by guanacos (Puig et al., 1997) occurred in the geomorphological units positively selected. The dominance of a herbaceous layer, and especially grasses intensively used such as Panicum urvilleanum, and preferred grasses such as Poa lanuginosa, accounted for the stable preference for the plain unit, and the proportional use of plateau. Grasses and forbs were major food items for guanacos in La Payunia, especially in summer (Puig et al., 1996). The availability of complementary species eaten with

Studies on Neotropical Fauna and Environment preference by guanacos is also associated with a proportional use of geomorphological units, such as the chamaephyte E. ochreata in piedmont during early summer. A prevalence of avoided species such as Stipa spp. characterized the hillside, a unit avoided throughout the summer. The geomorphological plain unit should be considered an optimal feeding area, as it combined a high availability of preferred food and a low risk of predation, due to the scarce obstructions to near visibility and movements. Sandy soils, gentle terrain and vegetation dominated by the herbaceous layer characterized the units most intensively used in the summer. Camelids are usually associated with open spaces (Raedeke, 1982; Cajal, 1989), and female guanacos selected vegas with a low peripheral hiding cover in a Subantarctic population (Lawrence, personal communication). The preference for open grasslands by the pronghorn antelope and the blue wildebeest was related to a strategy to detect predators from afar, and to escape if necessary after detection (Barnes et al., 1991), and could also justify such preference by guanacos. Relative importance of variables in the habitat selection by guanacos Non-interactive factors (proportion of rocks in soil, prevalence of hiding cover and strong slopes) as well as foraging factors (availability of preferred food items, prevalence of avoided or uneaten plant species) seems to concern the selection among feeding areas by guanacos. A combination of non-interactive and interactive variables affecting herbivore decisions is expected at intermediate spatial scales, such as the feeding habitats of wild ungulates (Quintana et al., 2002). A high relevance of food quality for the habitat selection by guanacos in summer, at least at the scale of geomorphological units, was shown by the association of the density of guanacos with a good availability of preferred food items (Panicum urvilleanum, Poa lanuginosa, C. patagonum), and with a low prevalence of avoided food items (Stipa spp., Schinus spp.) and uneaten plants (S. filaginoides, M. spinosum). Physical features such as low altitude, sandy soil, and low prevalence of hiding cover also appeared as important variables selected by guanacos, although the relevance of soil was difficult to isolate from the plant community, because low shrubs and Stipa spp. were the dominant vegetation in rocky soils. The abundance of guanacos in the area would affect the relevance of several features in the selection of habitats, considering that associations detected in

early summer disappeared in late summer as the number of guanacos increased in the study area. The association between the density of guanacos and a good availability of preferred grasses only persisted for Panicum urvilleanum in late summer, but disappeared for Poa lanuginosa and C. patagonum. The prevalence of hiding cover, represented by strong slopes and tall shrubs such as N. aphylum, was only relevant for habitat selection by guanacos in early summer. Concluding remarks The strategy of habitat use by guanacos during the reproductive season, at the scale of geomorphological units as potential feeding areas, would be characterized by: (1) selective habitat use based on vegetation features, preferring grazing areas with high availability of preferred plant species, and low prevalence of avoided or uneaten plant species; (2) constrained use of good grazing areas according to relief features probably related to predation risk, preferring areas with gentle terrain and low hiding cover; and (3) relative weakening of selective habitat use, with loss of avoidance of some poor and risky habitats when the guanaco abundance increases in the area. A complementary study on habitat use by guanacos in Northern Patagonia during winter would be relevant, considering the assumption that wildlife dynamics are usually constrained by winter survival (Morrison et al., 1998). Seasonal shifts in habitat use would be expected with winter conditions and the behavioral flexibility of guanacos. Winter is characterized by scarce food availability, harsh weather conditions, great aggregations of social groups, and absence of reproductive activities (Puig, 1995; Puig & Videla, 1995). Heterogeneous landscapes and unpredictable weather affect food availability in arid environments. Flexible movements of large herbivores, with free access to the scarce resources, are regarded as essential for the survival of ungulates in these environments (Scoones, 1995). Buttolph & Coppock (2001) warned that reducing the access to essential resources with fences could compromise the viability of camelids in Puna environments. Likewise, in Northern Patagonia the access to geomorphological units by guanacos, and therefore the selection among potential feeding areas, would be disrupted by fences. Construction of fences should be discouraged in La Payunia Reserve, especially fences parallel to piedmonts or basaltic scarps, in order to guarantee an optimal and complete habitat use by guanacos within this protected area.

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SAR data. In Proceedings of the 3rd Earth Observation Quarterly Symposium; Florence, European Space Agency. Florence, Italy. p. 1420. Franklin W. 1983. Contrasting socio-ecologies of South Americas wild camelids: the vicuna and the guanaco. In: Eisenberg JF, Kleiman D, editors. Recent advances in the study of mammalian behavior. American Society of Mammalogy. Lawrence, Kansas, USA. p. 573629, (American Society of Mammalogy Special Publication; 7). Gonzalez Daz EF. 1972. Descripcion geologica de la Hoja 30-d, Payun Matru, Mendoza. Buenos Aires: Direccion Nacional de Geologa y Minera. 97 p. (Carta Geologico-Economica de Republica Argentina Boletn; 130). Hair JF, Anderson RE, Tatham RL, Black WC. 1999. Multivariate data analysis. 5th ed, London: Prentice Hall. 832. Kie JG. 1999. Optimal foraging and risk of predation: effects on behavior and social structure in ungulates. J Mammal. 80:11141129. Kie JG, Bowyer RT. 1999. Sexual segregation in white-tailed deer: density-dependent changes in use of space, habitat selection, and dietary niche. J Mammal. 80:10041020. Kotliar NB, Wiens JA. 1990. Multiple scales of patchiness and patch structure: a hierarchical framework for the study of heterogeneity. Oikos. 59:253260. Martnez E, Dalmasso A. 1993. Flora y vegetacion. In Diseno del plan de manejo de Reserva La Payunia (Mendoza, Argentina). Multequina. 2:1922. McShea WJ, Aung M, Poszig D, Wemmer C, Monfort S. 2001. Forage, habitat use, and sexual segregation by a tropical deer (Cervus eldi thamin) in a dipterocarp forest. J Mammal. 82:848857. Morrison ML, Marcot BG, Mannan RW. 1998. Wildlifehabitat relationship. Concepts and applications. 2nd ed, Madison: University of Wisconsin Press. 458 p. Ortega IM, Franklin WL. 1995. Social organization, distribution and movements of a migratory guanaco population in the Chilean Patagonia. Rev Chil Hist Nat. 68:489500. Puig S. 1986. Ecologa poblacional del guanaco (Lama guanicoe, Camelidae, Artiodactyla) en la Reserva Provincial de La Payunia (Mendoza) [PhD dissertation]. Buenos Aires: Universidad de Buenos Aires. 532 p. Puig S. 1995. Uso de los recursos ambientales por el guanaco. In: Puig S, editor. Tecnicas para el manejo del guanaco. Gland (Switzrland), IUCN, p. 110126. Puig S, Videla F. 1995. Comportamiento y organizacion social del guanaco. In: Puig S, editor. Tecnicas para el manejo del guanaco. Gland (Switzrland), IUCN, p. 97118. Puig S, Videla F, Cona M. 1997. Diet and abundance of the guanaco (Lama guanicoe Muller 1776) in four habitats of northern Patagonia, Argentina. J Arid Environ. 36:343357. Puig S, Videla F, Monge S, Roig V. 1996. Seasonal variations in guanaco diet (Lama guanicoe Muller 1776) and in food availability in Northern Patagonia, Argentina. J Arid Environ. 34:215224. Quintana RD, Puig S, Rosati V, Arias SM. 2002. Seleccion de dieta por mamferos herbvoros silvestres a escala de habitat de alimentacion. In: Cid MS, Bonino N, Cassini M, Anchorena J, Pelliza Sbriller A, Arriaga M, editors. Seleccion de dieta por grandes herbvoros mamferos: procesos y escalas. Buenos Aires: Museo Argentino de Ciencias Naturales. p. 5977. Raedeke K. 1982. Habitat use by guanacos (Lama guanicoe) and sheep on common range, Tierra del Fuego, Chile. Turrialba. 32:309314.

Acknowledgments We thank R. Quintana for his enriching comments on the manuscript, A. Scolaro for his inestimable assistance in multivariate analyses, R. Gonzalez del Solar and G. Videla for field assistance, M. I. Rosi for laboratory assistance, N. Horak for his assistance in the English translation, and A. Rocher, owner of Ea. El Peralito, for the kind hospitality. Financial support was provided by CONICET (PID 3-387200). References
Altmann J. 1973. Observational study of behavior: sampling methods. Behaviour. XLIX:227267. Bailey BJR. 1980. Large sample simultaneous confidence intervals for the multinomial probabilities based on transformation of the cell frequencies. Technometrics. 22:583589. Bailey DW, Gross JE, Laca EA, Rittenhouse LR, Coughenour MB, Swift DM, Sims PL. 1996. Mechanisms that result in large herbivore grazing distribution patterns. J Range Manag. 49:386400. Baldi R, Abon SD, Elston DA. 2001. Guanacos and sheep: evidence for continuing competition in arid Patagonia. Oecologia. 129:561570. Barnes TG, Heitschmidt RK, Varner LW. 1991. Wildlife. In: Heitschmidt RK, Stuth JW, editors. Grazing management: an ecological perspective. Portland (OR): Timber Press. p. 179190. Barten NL, Bowyer RT, Jenkins KJ. 2001. Habitat use by female caribou: tradeoffs associated with parturition. J Wildl Manag. 65:7792. Bello J, Gallina S, Equihua M. 2001. Characterization and habitat preferences by while-tailed deer in Mexico. J Range Manag. 54:537545. Belovsky GE, Schmitz OJ. 1994. Plant defenses and optimal foraging by mammalian herbivores. J Mammal. 75:816832. Berger J. 1991. Pregnancy incentives, predation constraints and habitat shifts: experimental and field evidence for wild bighorn sheep. Anim Behav. 41:6177. Buttolph LP, Coppock DL. 2001. Intensified alpaca production leads to privatization of key grazing resources in Bolivia. Rangelands. 23:1013. Cabrera AL, Willink A. 1980. Biogeografa de America Latina. 2nd ed, Washington: Organization of American States. p. 1122, (Biological Series of the Organization of American States; 13). Cajal JL. 1989. Uso del habitat por vicunas y guanacos en la Reserva San Guillermo, Argentina. Vida Silvestre Neotrop. 2:2131. Cajal JL, Lopez NE. 1987. The puma as a predator of wild camelids in the San Guillermo Reserve, San Juan, Argentina. Rev Chil Hist Nat. 60:8791. Cherry S. 1996. A comparison of confidence interval methods for habitat use-availability studies. J Wildl Manag. 60:653658. Colwell RK, Futuyma DJ. 1971. On the measurement of niche breadth and overlap. Ecology. 52:567576. Consejo Federal de Inversiones. 1977. Proyecto de desarrollo ganadero del oeste de La Pampa. Volume I. Buenos Aires: Consejo Federal de Inversiones. 253 p. Daget P, Poissonet J. 1971. Une methode danalyse phytologique des prairies. Criteres dapplication. Ann Agron. 22:541. ` del Valle HF, de Lamo DA, Gagliardini DA. 1997. Environmental affinity of the guanaco (Lama guanicoe Muller, Camelidae) in two selected areas of Central Patagonia supported by ERS-1

Studies on Neotropical Fauna and Environment


Sarno RJ, Clark WR, Bank MS, Prexl WS, Behl MJ, Johnson WE, Franklin WL. 1999. Juvenile guanaco survival: management and conservation implications. J Appl Ecol. 36:937945. Scogland T, Grovan B. 1991. The effects of human disturbance on the activity of wild reindeer in different physical condition. Rangifer. 9:1119. Scoones I. 1995. Exploiting heterogeneity: habitat use by cattle in dryland Zimbabwe. J Arid Environ. 29:221237. Senft RL, Coughenour MB, Bailey DW, Rittenhouse LR, Sala OE, Swift DM. 1987. Large herbivore foraging and ecological hierarchies. BioScience. 37:789799. Stuth JW. 1991. Foraging behavior. In: Heitschmidt RK, Stuth JW, editors. Grazing management: an ecological perspective. Portland (OR): Timber Press. p. 6584.

Torres H. 1992. South American camelids: an action plan for their conservation. Gland (Switzerland): IUCN Species Survival Commission. 26 p. Underwood R. 1982. Vigilance behaviour in grazing African antelopes. Behaviour. 79:81107. Videla F, Puig S. 1993. Fauna de vertebrados. In Diseno del plan de manejo de Reserva La Payunia (Mendoza, Argentina). Multequina. 2:2327. Wheeler JC. 1991. Origen, evolucion y status actual. In: Fernandez Baca S, editor. Avances y perspectivas del conocimiento de los camelidos sudamericanos. FAO, p. 1148. Zar JH. 1984. Biostatistical analysis. 2nd ed, Upper Saddle River (NJ): Prentice-Hall. 718 p.

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