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Population Ecology of the Vicuna (Vicugna, vicugna) at the Salinas y Aguada Blanca National Reserve, Arequipa, Peru: Bas...


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POPULATION ECOLOGY OF THE VICUA ( Vicugna vicugna ) AT THE SALINAS Y AGUADA BLANCA NATIONAL RESERVE, AREQUIPA, PERU: BASELINE DATA FOR SUSTAINABLE MANAGEMENT By JENNIFER E. DAVIES A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF ARTS UNIVERSITY OF FLORIDA 2003

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Copyright 2003 by Jennifer E. Davies

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For Murphy

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ACKNOWLEDGMENTS I would like to thank Dr. Lauren Chapman for all the knowledge and guidance she brought in the completion of this project. Her dedication to science, kindness, and integrity are truly inspiring. She definitely sets the standard in achievement both personally and professionally. I would also like to thank Dr. Charles Wood for all his help and support during my time in Florida, and for agreeing to serve on my committee. I would also like to thank Dr. Emilio Bruna for his support, encouragement, and knowledge, and for graciously agreeing to serve on my committee. In addition, I would like to thank all those at CONATURA and Dr. Catherine Sahley for providing assistance at the field site in Arequipa, use of their database, and for various crucial aspects of the project. The field research may not have been possible without the invaluable friendship and support of Vajk Lucas and Mariela Caceras (and Maya) in Arequipa. They shared their home to an unexpectedly long-term visitor, who would come home covered in dust and smelling of alpacas. I am honored to have been a part of their lives. Comments and knowledge offered by Colin Chapman and Ron Sarno were also invaluable and added greatly to the thesis. I also greatly appreciate the support offered by those in the Tropical Conservation and Development program throughout my years of study, namely Dr. Marianne Schmink and Hannah Covert. Technical support (and overall support) was provided by Joe Savastano. Financial support for this research was provided by the Department of Latin American Studies, the Tropical Conservation and Development program at the University of Florida, the Wildlife Conservation iv

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Society, and CONATURA. Finally, I would like to thank my family and friends for their encouragement and patience during the completion of this project. v

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TABLE OF CONTENTS page ACKNOWLEDGMENTS.................................................................................................iv LIST OF TABLES...........................................................................................................viii LIST OF FIGURES...........................................................................................................ix ABSTRACT.........................................................................................................................x CHAPTER 1 INTRODUCTION........................................................................................................1 General Ecology...........................................................................................................1 Objectives.....................................................................................................................4 2 STUDY SITE DESCRIPTION.....................................................................................6 3 METHODS.................................................................................................................10 Study Design...............................................................................................................10 Data Analyses.............................................................................................................12 Group Characteristics..........................................................................................12 Distribution and Density......................................................................................12 Density.........................................................................................................12 Density and distribution relative to the laguna.............................................13 Group size distribution relative to the laguna..............................................13 Responses to Changes in Water Distribution......................................................14 Spatial Patterns and Inter-Group Movement.......................................................15 4 RESULTS...................................................................................................................16 General Ecology.........................................................................................................16 Group Size and Population Composition............................................................16 Group Size and Composition by Group Type.....................................................17 Spatial Relationships and Intergroup Movement.......................................................18 Density and Distribution......................................................................................18 Density and distribution relative to the laguna.............................................18 Group size distribution relative to the laguna..............................................19 vi

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Group composition relative to distance from laguna...................................22 Responses to Changes in Water Distribution......................................................24 Average distance from laguna as a function of changes in water distribution..............................................................................................24 Density and distribution as a function of changes in water distribution......29 Spatial Patterns and Inter-Group Movement.......................................................29 5 DISCUSSION.............................................................................................................37 General Ecology.........................................................................................................37 Group Size and Type Composition.....................................................................37 Mating Strategies of the Vicua..........................................................................38 Group Composition Within Family Groups........................................................39 Group Size and Density.......................................................................................40 Spatial Relationships and Intergroup Movement.......................................................44 Distribution Patterns............................................................................................44 Seasonal Changes in Distribution and Density...................................................46 Spatial Patterns and Inter-Group Movement.......................................................49 Mean age of identified males found at laguna in 2002................................50 Site fidelity and dispersal.............................................................................51 Distance moved from family groups............................................................52 Description of shifts in group composition..................................................52 6 IMPLICATIONS FOR CONSERVATION...............................................................54 7 CONCLUSIONS........................................................................................................59 LIST OF REFERENCES...................................................................................................61 BIOGRAPHICAL SKETCH.............................................................................................65 vii

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LIST OF TABLES Table page 4-1.Composition of identified vicua (Vicugna vicugna)..................................................33 4-2.Identified male groups of vicua (Vicugna vicugna)..................................................33 4-3.Life history characteristics of tagged male vicua (Vicugna vicugna).......................36 4-4.Examples of movement within female vicua among family groups.........................36 5-1.Published denisty estimates of vicua (Vicugna vicugna)..........................................43 viii

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LIST OF FIGURES Figure page 2-1.Transect design for vicua (Vicugna vicugna) survey..................................................8 4-1.Percentage of group types in the vicua (Vicugna vicugna) population.....................17 4-2.Vicua (Vicugna vicugna) group density ...................................................................20 4-3.Distribution of vicua (Vicugna vicugna) group types around the laguna..................21 4-4.Size of vicua (Vicugna vicugna) groups in relation to the laguna.............................23 4-5.The number of females in family groups of vicua (Vicugna vicugna)......................25 4-6.The number of young vicua (Vicugna vicugna) in relation to distance.....................26 4-7.The number of males in vicua (Vicugna vicugna) groups.........................................27 4-8.Average distance of vicua (Vicugna vicugna) groups...............................................28 4-9.Distribution of vicua (Vicugna vicugna) groups.......................................................30 4-10.Density (groups per sq km) of vicua (Vicugna vicugna).........................................31 4-11.Percentage of individual vicua (Vicugna vicugna) group types..............................32 4-12.Spatial patterns of followed vicua (Vicugna vicugna) groups.................................35 6-1.Historic range of the vicua (Vicugna vicugna) in South America.............................55 ix

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Abstract of Thesis Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Arts POPULATION ECOLOGY OF THE VICUA (Vicugna vicugna) IN THE SALINAS Y AGUADA BLANCA NATIONAL RESERVE, AREQUIPA, PERU: BASELINE DATA FOR SUSTAINABLE MANAGEMENT By Jennifer E. Davies December 2003 Chair: Dr. Lauren J. Chapman Major Department: Latin American Studies The installation of corrals to facilitate the capture and protection of the vicua (Vicugna vicugna) has been encouraged by government agencies in Peru to meet sustainable development objectives in rural Andean communities. Corrals may have an impact on vicua populations and their habitats by altering the density of the vicua. However, little is known about the home range, movement patterns, or habitat requirements of free-ranging vicua, which precludes an accurate estimate of the size a corral should be to sustainably manage these wild camelids. The objective of the thesis is to provide quantative data on aspects of the population ecology of the vicua that will have application to corral management initiatives. Repeat samples of a 38-km transect and group follows were used to estimate four major ecological traits of vicua groups in the Salinas y Aguada Blanca National Reserve in Arequipa, Peru. These included 1) group characteristics, 2) movement patterns, 3) responses to declining water availability in the major laguna for vicua in the reserve, and 4) spatial patterns and inter-group x

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movement. Analyses of these four ecological traits were used to derive patterns of habitat use, the relationship to water resources, and density variation across space and time. Family groups made up 67% of all groups within the population, had an average group size of 5.6 individuals, and had a group composition of 1 male, 3.7 females, and 1.6 young. Overall density of vicua in the study area was 3.7 vicua km2. Water distribution and availability had a major impact on vicua movement, but no detectable effect on group and population composition. Group density was highest < 4 km from the laguna, suggesting that the water source was a key feature in the vicuas habitat. Movement patterns indicated differential habitat use among group types around the laguna. Large family groups and bachelor herds were more frequently found in proximity to the laguna, while smaller groups (non-defined groups and solitary males) were distributed more evenly. Family groups showed a low degree of spatial overlap around the laguna suggesting that the family groups in the reserve are territorial and maintain their boundaries while at the laguna. xi

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CHAPTER 1 INTRODUCTION General Ecology The vicua (Vicugna vicugna) is a New World camelid that inhabits the high, dry puna, or altiplano zones in the Andean mountains of Peru, Chile, Argentina, and Bolivia between 3,700 4,900 m (Koford, 1957; Franklin, 1983; Vila, 1992). This species is one of three wild ungulates occurring in these arid habitats. Others include the guanaco (Lama guanicoe) and the guemal (Hippocamelus antisiensis). It is widely reported that the vicua is a dietary specialist and therefore restricted to these elevations (Franklin, 1978; Lucherini, 1996). However, some authors suggest that since the arrival of humans and their livestock during the Incan period, the vicua has been displaced to higher elevations (Hofmann et al., 1983). The vicua is sedentary and spends most of its time foraging and ruminating on the hard bunch grasses of the puna (Bosch and Svendsen, 1987; Franklin, 1983; Lucherini, 1996; Menard, 1982; Renaudeau dArc et al., 2000; Vila and Cassini, 1993). Habitat use appears restricted, as vicuas have been found to avoid open rocky areas and only use areas that support preferred forage (Franklin, 1983; Renaudeau dArc et al., 2000). This vegetation preferred by vicuas includes species often associated with the scarce ground water supply found in the arid puna environment. The vicua requires a frequent supply of water not obtained by either oxidative metabolism or from the food they ingest (Bosch and Svendsen, 1987; Franklin, 1978, 1983; Menard, 1982; Vila and Roig, 1992; Vila and Cassini, 1993). Vicua have been reported to exhibit daily foraging and drinking patterns 1

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2 (Franklin, 1983; Renaudeau dArc et. al., 2000; Vila and Cassini, 1993) in response to the drastic fluctuations in temperature, low precipitation, and medium to poor forage quality. In their study of seasonal activity patterns Vila and Cassini (1993) found a daily peak in the use of water sources at noon. This pattern in activity was more defined during the dry season (Vila and Cassini, 1993), suggesting a strong relationship to the distribution of water. However, a fine-grained analysis on water requirements and effects of water distribution on vicua spatial patterns has not been undertaken. The social organization of the vicua is based on family groups and male groups (Franklin, 1983; Vila and Cassini, 1993). Several studies propose that male vicuas in family groups are territorial (Bosch and Svendsen, 1987; Franklin 1978, 1983; Koford, 1957; Vila and Roig, 1992), and there is evidence for variation in group type and size with general habitat quality. However, direct links between group type and water resources are not understood well. A more complete picture of vicua population ecology is critical given increasing pressures on their populations, and new intensive game management initiatives. Vicua fiber is well suited for the drastic extremes of the Andean climate and is considered to be the finest in the world. The fiber is 13-14 microns in diameter, which is finer than cashmere (15-19 microns in diameter). This makes vicua a target species for rural development strategies, because the fiber is viewed as such a lucrative, renewable natural resource. In fact, vicua fiber has been hailed as the gold of the Andes, a viable way to stimulate the economies of impoverished indigenous communities in the area (Cattan and Glade, 1989; Rabinovich et al., 1985; Sahley et al., 2001).

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3 The main threat to vicua populations is the poaching for hides and their fiber (Lichtenstein et al., 2002). This contributed to their rapid decline to an estimated 10,000 individual vicuna in the 1960s (CONACS, 1997). After being listed under Appendix I of the CITES convention since the 1970s, the vicua was reclassified to the Appendix II species list in 1995 (Wheeler and Hoces, 1997). Recent increases in the number of vicua are assumed to reflect protectionist practices centered on creation of national parks and sustainable use of wild vicua populations to harvest fiber (Cajal, 1991; CONACS, 1997). Three parks were established in Peru to facilitate protection of wild camelids: 1) Salinas y Aguada Blanca National Reserve (approximately 366,000 ha), located just north of Arequipa, 2) Huascaran National park in Ancash (28,000 ha) and 3) Pampas Galeras National Reserve (75,000 ha). A vicua sheared is a vicua saved became the slogan of stakeholders to encourage sustainable use of the vicua. Stakeholders claim that conservation objectives can be met with the protection of vicua by park guards from poachers, and the sale of sheared vicua fiber as an economic alternative through community initiatives. These measures have indeed been successful in bolstering vicua populations in a limited number of communities such as Pampa Galeras, but do not reflect an increase in distribution along the vicuas entire range in the Andes. In addition to the development of protected areas, new management schemes that intensively manage dense vicua herds in structured corrals have been initiated. The corrals facilitate capture and protection of vicua; and their utilization has been both encouraged by the government and adopted by numerous rural communities. These corrals are typically 500-1,000 ha in size and may contain 250-1,000 vicuas

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4 (Lichtenstein et al., 2002). The installation of corrals may have a detrimental impact on the vicua populations and the xeric habitat they inhabit by anthropogenically altering their natural density. Corrals that limit movement may have an impact on the ungulate population by: minimizing genetic dispersal, increasing the rate and incidence of disease, promoting the breakdown of family units, decreasing breeding or infant survival, and/or altering plant communities. However, we understand very little about the home range size, movement patterns, or territory requirements of the vicua. This precludes an accurate estimate of the size a corral should be to sustainably manage these wild camelids. Such data on habitat requirements are crucial to the establishment of effective management programs (Hixon, 1980; Renaudeau dArc et al., 2000). Objectives The goal of this study was to quantify key features of the population ecology of the vicua and to consider implications of these characteristics for corral initiatives. Field work was conducted in the Central Andes within the Salinas y Aguada Blanca National Reserve (Arequipa, Peru) from April 25th to July 12th of 2002. I assessed group composition, density, and movement patterns of the vicua, as well as the aggregative relationship to the distribution of water. My specific objectives were as follows. First, I quantified group composition characteristics to provide baseline data for comparison with free-ranging and corralled vicuna. These included: representation of age/sex classes within groups, sex ratio, and group size. Second, I used a repeat transect design to quantify general movement patterns and spatial distribution in a 20 km2 area that included representative habitat types and a major water source (a laguna). Data on spatial relationships were combined with group composition data to analyze movement patterns

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5 for group types relative to the laguna. Third, I explored the response to changes in water distribution by evaluating relationships between spatial distribution patterns and changes in the size of the major water source (index of dryness). Finally, I examined movement patterns of individually identified vicuna groups and quantified aspects of group stability. This permitted an evaluation of site fidelity among tagged males around the laguna, distance traveled over time, and group stability.

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CHAPTER 2 STUDY SITE DESCRIPTION The study was conducted between April 25 and July 12 of 2002 within the Salinas y Aguada Blanca National Reserve in the department of Arequipa, Peru (154505 and 161915 south and 705120 and 71340 west). The reserve was created in 1979 and contains 366,936 ha of dry puna characterized by high mountain plains, volcanoes, hillsides, and cliffs, with elevation between 3,400 to over 6,000 m. The daily temperature in the reserve averages between 2 to 8 C (absolute minima reaching C), and dramatic diel temperature fluctuations are characteristic (Caviedes and Knapp, 1995; Pulgar, 1996). Six Holdridge life zones are represented in the reserve. These include: 1) subtropical desert mountains, 2) subtropical sub alpine mattoral desert, 3) subtropical sub alpine humid paramo, 4) subtropical alpine humid tundra, 5) subtropical alpine very humid tundra, and 6) subtropical nival. Also characteristic of this region is the presence of snow at the highest elevations (typically the volcanoes) throughout the year. This is an important source of water for both the vegetation and wildlife. Total annual precipitation in this area fluctuates between 200 mm and 1,000 mm indicating a high degree of interannual variability in water availability. The April to July period of this study covered the end of the wet season, an extremely dry period, and an unexpected wet period associated with a July snowfall. Plants found in the study site display adaptations to dry puna, such as slow growth rates, small plant size, a bunched distribution, and a shallow root system. The common 6

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7 plants in the reserve include bunched grasses such as the common ichu grass (Stipa ichu), several kinds of cactus such as ulluyma (Opuntia floccosa), the huajoro (Opuntia lagopus), and the pajuro (Opuntina ignescens). This region also supports several species of scrub brush. These include the arbustos enanos such as the shauli, and the arbustos de Culli, (Colli, queua, and quinal) (Polylepis Racemosa) a tree-like plant that is heavily exploited for wood. The puma (Puma concolor), vicua (Vicugna vicugna), and guanaco (Lama guanicoe) are the only large non-domesticated mammals in Salinas y Aguada Blanca National Reserve. However, no signs of puma were found in the focal study area. The only mammalian predator assumed to be in the reserve is the Andean fox (Dusicyon culpaeus). In addition to the large wild mammals found in this region of the reserve, domestic animals inhabit regions near as well as in the reserve. Alpaca, sheep, goats, and horses are all competitors of the wild species. The domestic livestock are kept spatially separated from the vicua on the mojedales (wetlands). In other areas of the vicuas native range, these wetland areas are considered important habitat for wildlife, including the vicua (Franklin, 1983; Renaudeau dArc et. al., 2000; Rundel and Palma, 2000). However, in this reserve, only the domestic animals have access to these water and vegetative resources. The major water source for the vicua in the reserve was the laguna. No other water source was found in the reserve that was used by vicua (Davies, pers. obs.). Due to its location within the reserve, land use by the community of Tambo Caahuas also has a significant impact on wildlife populations. An important feature in this landscape is the road that runs through both the community and the reserve (Figure

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8 2-1). This is a major thoroughfare between the cities of Arequipa (the second largest city in the country) and Juliaca, a city with significant commerce. There are also several tourist attractions and active mines in this area that attract many large trucks and additional traffic. As it crosses through the reserve adjacent to the major water source, this road potentially contributes point-source pollution and road-kill mortalities that could have long lasting, detrimental impacts on the wildlife populations. # 80816 SNEW Paved Highway Transect LagunaSumbayChachani VolcanoMisti Volcanokilometers Study siteTambo Caahuas Figure2-1. Transect design for vicua (Vicugna vicugna) survey at the Salinas y Aguada Blanca National Reserve, Peru, during April-July 2002. Patterns of land use are indicated as well as the main roads that pass through the reserve. Current sustainable management strategies in Tambo Caahuas involve the live capture, shear and release (called the Chaccu) of free-ranging vicuas approximately once every 2 yr. CONATURA, an Arequipa-based non-governmental organization assists this community with logistical support, training, and organization. During the

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9 time when these vicuas are captured and sheared, biologists from CONATURA tag individuals with cow tags and take measurements on age, sex, condition, and length and diameter of the fiber. A total of 71 vicuas were tagged between 1997 and 2000. In addition, this NGO conducts scientific field studies and has established a monitoring program to help provide data for management decisions.

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CHAPTER 3 METHODS Study Design Repeat samples of a 38-km transect and group follows were used to estimate four major ecological traits of vicua groups. These included 1) group and population composition, 2) density and distribution, 3) the response to changes in water distribution, and 4) spatial patterns and intergroup movement. Analyses of these four ecological traits were used to quantify spatial and temporal patterns of habitat use, dispersal, relationship to water resources, and density variation.In addition, a 3-yr CONATURA database was used to reference individual histories on the vicuas observed in the field survey. The database comes from a long-term monitoring project and is comprsied of 15 surveys between 1999 and 2001. Data collected by CONATURA include location, group size and composition, and identification of some tagged individuals. One large 38-km transect was established that covered a representative area of the reserve including the major water source (the laguna), a large pampa or intermontane plain, and regions near the highway. This transect was surveyed 13 times over the 4-month study period. For each group or individual vicua sighted, the following information was recorded: group type (i.e., bachelor herds, family groups, non-defined groups and solitary males), group composition (# males, # females, # young), the identity and gender of marked individuals, group activity, time of day, and GPS location recorded in UTM coordinates with a Garmin 12XL GPS unit (KS, USA). Analyses of movement patterns, distribution, density, and dispersal were based on obtaining exact positions of 10

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11 vicua family groups, bachelor herds, and solitary individuals. This was done using a variation of the distance method (Buckland et al., 1993). A Garmin 12xl GPS (Global Positioning Unit) in UTM was used from the transect to obtain coordinates of the vicua groups. The center of the group was taken as the location of the group sighted, and the bearing and distance were also recorded from this point. These data were then transformed using the equation (d)*Cos()= x and (d)*Sin () = y. x was then used with the equation: x = x1 + x2, and y = y1 + y2, and the new UTM coordinate was found for the exact position. Specific information on marked individuals and behavioral information were obtained with a Bushnell Spacemaster spotting scope. To determine family group composition and to identify marked individuals, I also recorded tag numbers of marked individuals (sexes can be differentiated because males have tags on left, while females have it on the right). Differentiation of yearlings and adult vicua is difficult, and this information was not collected. From the transect study, distinct group types, [family groups, bachelor herds, solitary males, non-defined groups (groups of unknown composition)] were identified and followed to assess movement patterns among different social associations. The study was conducted in two periods. The first sampling period ran from April 15 May 23. During this period the laguna was full, and the data collection focused on the group census. The second period ran from June 24 July 12, during which time the laguna and the surrounding plains became increasingly dry until a snowfall on July 10. The size of the laguna was mapped immediately after the rainy season when the laguna was full (April 25), and after a long dry period (June 28). The laguna was approximately 50 times larger in April than in June. In the first sampling period, transects were used to

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12 identify groups and document their composition. In the second period, I used the same transect design, but also followed known family groups to provide data on movement and site fidelity. Data Analyses Group Characteristics Group sizes observed during the wet and dry sampling periods were compared with a t-test and then combined to produce one transect data set. From the combined transect data, group size was averaged to obtain mean group size for all group types. To determine the sex ratio within the vicua population at the Salinas y Aguada Blanca National Reserve, averages of the number of males and females were calculated from the combined transect data. A one-way analysis of variance (ANOVA) was used to test for a difference in mean group size among bachelor herds, family groups, and non-defined groups. The Scheffe aposteriori test was used to detect differences in mean group size between any two group categories. Group size values were log transformed to normalize the data and stabilize the variance. Regressions were used to test for relationships among group composition characteristics such as group size, number of females, and number of young. Distribution and Density Density Vicua density was calculated by counting the number of vicua within a sighting distance of 500 m from the transect, taking into account sampling effort, and then dividing by the effective area sampled. Effective strip width (500 m) was defined by measuring the perpendicular distance from the transect to each vicua group observation. A sighting distance was determined from a frequency histogram of the distance a vicua

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13 group was observed (Chapman et al., 2000). The frequency of vicua sightings were consistently higher at distances less then 500 m. All vicua sightings greater then 500 m were taken out of the overall density estimate. This sighting distance of 500 m corresponds with figures used in previous studies that estimates vicua density (C. Sahley, unpublished data). The total numbers of vicuas in groups selected using this criterion were averaged to include in the density estimate. To determine the area effectively sampled, the sighting distance was multiplied by the transect length. Portions where the transect overlapped were taken out of the measurement. The density of vicuas was then calculated as the number of individual vicuas sighted divided by the effective area sampled. Group density was simply calculated with the aid of the map generated with Arc View, and calculated as the number of groups within certain distances from the laguna, corrected for sampling effort. Density and distribution relative to the laguna Arc View was used to create a series of maps describing the geographic locations of vicua groups in relation to the laguna in the transect survey. A series of 20 concentric rings (1 km apart) around the lagunas center were superimposed on the map of the transect data. These maps were used to calculate the distribution of group size, group composition, and group type in relation to distance from the laguna. Group density was calculated as the number of groups within a certain distance from the laguna corrected for sampling effort. Linear regression was used to detect the relationship between density (dependent variable) and distance from the laguna (independent variable). Group size distribution relative to the laguna Group size was extracted from the transect data and mapped with the aid of Arc View to find patterns of distribution around the laguna. Vicua groups were arbitrarily

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14 categorized into three size classes, small groups (1-5), medium groups (6-15), and large groups (16-40). A chi-square test of independence was used to detect differences between group sizes close (<4 km) and far (>4 km) from the laguna. The number of males, females and the number of young in groups were quantified in relation to their distance from the laguna. Regression analysis was used to detect relationships between the number of males, females, and young (dependent variable) to distance from the center of the laguna (independent variable). Arc View was also used to map this data for a visual display of distribution in relation to the laguna. Responses to Changes in Water Distribution The study was conducted in two periods (April 15-May 23 and June 24-July 12), therefore, a t-test was used to detect differences in group size between the wet and dry sampling periods. The distances of vicua groups from the laguna were categorized into monthly averages to reflect changes in distribution in relation to changing water distribution. An Arc View map of the trasect data was superimposed on a map of the study area. The distances vicua groups were observed from the laguna was recorded for each month, correcting for differences in sampling effort. Temporal changes in density and distribution were also calculated by quantifying variation between wet and dry periods. A series of Arc View-generated maps of the transect data with 1 km concentric circles around the center of the laguna was used to quantify variation in density and distribution of vicua groups relative to the center of the laguna in the two major sampling periods. A chi-square test for independence was used to detect differences in the distribution of vicua groups and group type around the laguna between the wet and dry periods.

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15 Spatial Patterns and Inter-Group Movement To estimate spatial characteristics among family groups, locations of identified dominant males were taken from the transect data and the group follows as these groups moved to the laguna to drink. These observations were then mapped with the aid of Arc View to show group movement patterns over time. In addition, the life histories of tagged males that I observed were extracted from the CONATURA database to estimatle site fidelity and to detect patterns in movement and group characteristics. These individually identifiable groups were followed based on opportunistic observations to obtain several geographic positions for 2-3 hr during the morning (sunrise to noon) and in the afternoon (noon to nightfall). These groups were found and followed within an approximate 5-km radius from the laguna. Finally, tagged females observed in indentified family groups were extracted from the CONATURA database to explore the validity of group stability as assumed in the literature.

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CHAPTER 4 RESULTS General Ecology Group Size and Population Composition Family group size (t=-1.295, P=0.197, mean wet=6.07, n=136, SE+/-0.24; mean dry=6.6, n=79, SE+/-0.31) and the number of females (t=0.108, P=0.914, mean wet=3.7, n=136, SE+/-0.16; mean dry=3.6, n=79), did not differ between the two major sampling periods. However, there were more young seen in the first period (mean=0.98, n=191) than the second period (mean=1.22, n=149, t-test, t=-2.845, P=0.005). Since young only comprised 22% of the population, and other characteristics did not differ between periods, I combined sampling periods to provide an overall evaluation of group characteristics (n=215groups). Family groups comprised 67% of the total groups sighted, and bachelor herds and solitary males represented 12% (6% each) of the sightings. Non-defined groups (indistinguishable groups of vicua) comprised 21% of the observed group types (Figure 4-1). Sex ratio The sex ratio for adult vicuas was 1.5 females for every 1 male. For family groups, this ratio increased to 3.7 females for every 1 male. This ratio is based on adults in identifiable group types, without consideration of age class. 16

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17 0.00%10.00%20.00%30.00%40.00%50.00%60.00%70.00%family groupsbachelorherdssolitarymalesnon definedgroups Figure 4-1. Percentage of group types in the vicua (Vicugna vicugna) population of the Salinas y Aguada Blanca National Reserve, Peru. Family groups were the groups most frequently found in the survey and were comprised of a single male and a variable number of females and young. Non-defined groups were comprised of unidentified individuals with no young. Bachelor herds were groups made up of several identified males and no females or young. Solitary males comprised of a single male. Total group sightings = 317. Group Size and Composition by Group Type Family group size averaged 5.6 (antilog mean, n=215) and ranged from 2 to 13. Bachelor herd size was larger than that of family and non-defined groups (ANOVA F= 102.98, p<0.001, Scheffe p<0.001), averaging 15.9 (n=18) and ranging from 8 to 40. Non-defined groups were classified as indistinguishable groupings of vicua whose group size averaged 2.9 (n=65) and ranged from 2 to 13. There were an average of 3.7 females and 1.6 young per family group. Family group composition thus averaged 1: 3.7: 1.6 (male: female: young). Thirty-two percent

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18 of all family groups had four or more females. For observed males, 39.2% were in family groups; 57.4 % were in bachelor herds, and 3.5 % were solitary. Both the number of young and the number of females were correlated with group size (young: r = 0.84, p<0.001, n=215; females: r = 0.93, p<0.001, n=215). The number of young was also positively correlated with the number of females (r = 0.60, p<0.001, n=215). However, this relationship was not as strong as the relationship between the number of young and group size, suggesting a greater amount of variation in the number of young per family group. In addition, for males there was only one male in every family group despite the variation in family group size. Spatial Relationships and Intergroup Movement Density and Distribution Density and distribution relative to the laguna At the Salinas y Aguada Blanca National Reserve vicua density averaged 3.7 vicuas/ km2 over the period of the investigation. Vicua density increased closer to the laguna (r = -0.50, p<0.009; Figure 4-2). Forty-eight percent of all vicua groups in a 20 km radius were found within 2 km from the center of the laguna. For data combined across the 4-month study period, all group types were present at the laguna. However, habitat use of group types differed between the area close to the laguna (within a 4-km radius) and areas more distant (4 to 20 km). The proportion of group types was not independent of distance (2=15.16, p<0.005). Family groups dominated in the first 4 km from the laguna, comprising 71% of all group types. Non-defined groups made up 20%, followed by bachelor herds (6%) and solitary males (3%, Figure 4-3). Four to 20 km from the laguna, family groups made up 57% of all group types (a 14% decline), while the percentage of non-defined groups increased to 28%.

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19 There were no bachelor herds more then 4 km from the laguna, and solitary males comprised 15% of the groups 4 to 20 km from the laguna. Non-defined groups and solitary males were more evenly distributed throughout the study area than bachelor herds and family groups (2=9.07, p<0.005). Seventy percent of all family groups and 90% of all bachelor herds were found within 4-km of the laguna. Only 56% of all non-defined groups and 27% of solitary males were found in this distance class. In comparison, 4 to 20 km from the laguna, only 10% of the bachelor herds were observed, and only 30% of the family groups. While 44% of all non-defined groups and 73% of all solitary males were observed in farther regions from the laguna. When translated to group densities, patterns were similar. Group size distribution relative to the laguna Vicua groups were arbitrarily divided into three size classes: small groups (1-5 individuals), medium groups (6-15 individuals), and large groups (16-40 individuals). Small groups accounted for 54% of the whole vicua study population, while medium and large groups comprised 44% and 2% of the groups, respectively. To evaluate group size relative to the laguna, I quantified the frequency of group size types as a function of distance from the laguna using 1-km increments (Figure 4-4). There was a tendency for larger groups to be more frequent closer to the laguna than in the drier regions (2=5.1, 0.1
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20 00.511.522.531234567891011121314151617181920Distance from laguna (km)Groups per sq km Figure 4-2. Vicua (Vicugna vicugna) group density as a function of increasing distance from a laguna in the Salinas y Aguada Blanca National Reserve, Peru. Forty-eight percent of all vicua groups were found 2 km or less from the center of the laguna.

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21 Bachelor Herds Family Groups Non-defined Groups Solitary Males Figure 4-3. Distribution of vicua (Vicugna vicugna) group types around the laguna at the Salinas y Aguada Blanca National Reserve, Peru. Data were collected over 13 sampling dates between April 25 and July 12, 2002 by observing groups from a 38-km transect.

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22 To evaluate group size relative to the laguna, I quantified the frequency of group size types as a function of distance from the laguna using 1-km increments (Figure 4-4). There was a tendency for larger groups to be more frequent closer to the laguna than in the drier regions (2=5.1, 0.14 km; 2 = 16.37, p<0.005). All large family groups were found closer than 4 km from the center of the laguna. For medium sized groups, 65% were found within 4 km from the laguna, while 58% of small family groups were found in this distance class. Group composition relative to distance from laguna The number of females and the number of young in groups were quantified in relation to their distance from the laguna. The number of females decreased with increasing distance

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23 Sumbay Laguna Tambo Caaguas Transect Road Group Size 1-5 6-15 16-40 Figure 4-4. Size of vicua (Vicugna vicugna) groups in relation to the laguna. Vicua groups were categorized into three size classes: small (1-5), medium (6-15), and large (16-40). There was a tendency for large groups to be found closer to the laguna than in the drier regions of the study site. Group size data were collected through the repeat census (number of group sightings=317) of a 38-km transect between April 25 and July 12, 2002.

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24 from the laguna (r = -0.64, p<0.001), as did the number of young (r = -0.70, p<0.001, Figures 4-5,4-6,4-7). Only 19% of all family groups had more then the average number of young. Large grouping of males (bachelor herds) were almost exclusively found near the laguna (Figure 4-7), and there was a strong relationship between the size of bachelor herds and water distribution (r = -0.60, p<0.001). Responses to Changes in Water Distribution Average distance from laguna as a function of changes in water distribution The distances from the laguna that vicua groups were observed, were categorized into monthly averages to reflect changes in distribution in relation to changing water distribution. In April and May, the average distance of vicua from the laguna was 4072 m, (SD+/-1158.19) and 4127 m, (SD+/-4544.65) respectively. However, average distance declined to 2568 m, (SD+/-549.88) in June, which coincided with much lower water levels in the laguna. In July, the average distance increased to 3996 m, (SD+/-2119.72) (Figure 6a), possibly due to the unexpected snowfall in July, that increased the extent of water around the plains and mountains. This analysis included the farthest regions sampled, which may contain vicua groups using water sources other then the laguna. To control for this potential bias, I repeated the analysis removing groups more than 12 km from the laguna. The pattern was similar, although averages were slightly lower (Figure 4-8).

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25 Figure 4-5. The number of females in family groups of vicua (Vicugna vicugna) as a function of distance from the laguna in the Salinas y Aguada Blanca National Reserve, Peru. Family groups were divided into two categories: those with less then the average number of females (0-3) and those with more then the average (4-10). Data points represent observations from repeat sampling of a 38-km transect on 13 occasions between April 25 and July 12, 2002.

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26 N umber of young 0-2 3-10 Distance to Laguna Figure 4-6. The number of young vicua (Vicugna vicugna) in relation to distance from the laguna at Salinas y Aguada Blanca National Reserve, Peru. Family groups that had less than average number of young (0-2) and those that had more then average (3-10) were compared. Data points represent observations from repeat sampling of a 38-km transect on 13 occasions between April 25 and July 12, 2002.

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27 N umber of Males 0-1 2-40 Distance to La g una Figure 4-7. The number of males in vicua (Vicugna vicugna) groups in relation to distance from the laguna at Salinas y Aguada Blanca National Reserve, Peru. Because family groups consist of only one male, most groups (67%) had only one male. Vicua groups with more then one male were bachelor herds. Data points represent observations from repeat sampling of a 38-km transect on 13 occasions between April 25 and July 12, 2002.

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050010001500200025003000350040004500AprilMayJuneJuly 050010001500200025003000350040004500AprilMayJuneJulyMonthDistance to laguna (m)(a)(b) 28 Figure 4-8. (a) Average distance of vicua (Vicugna vicugna) groups from the laguna at Salinas y Aguada Blanca National Reserve, Peru, and (b) average distance of groups to the laguna not including distances greater than 10 km. Average distances were based on repeat sampling of a 38-km transect on 13 occasions between April and July, 2002.

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29 Density and distribution as a function of changes in water distribution A higher density of vicua groups was observed with delcining distances from the laguna (Figure 4-9). When the laguna was full, 65% (n=82) of all vicua groups were found within the first 4 km from the center of the laguna. In contrast, when the laguna was dry, 83% (n= 102) of all vicua groups were found within this 4-km radius (Figure 4-10). Family groups and bachelor herds seem to have been most affected by decreasing water availability. The distribution within group types was quantified and divided into those <4 km and >4 km from the laguna when full and when almost dry (Figure 4-11). Seventy-five percent of all bachelor herds were found less then 4 km to the laguna when full, and 100% when the laguna was almost dry. Sixty percent of all family groups were present <4 km when the laguna was full and 75% were present when almost dry (Figure 4-11). These differences in group distribution were marginally significant (family groups: 2= 2.85, 0.1

4 km from the laguna was independent of season suggesting a more even distribution that did not change with season (non-defined: 2 = 0.3, 0.9
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30 (a) Wet season (b) Dry season Solitary Male Family Group Non-defined Groups Bachelor Herds Figure 4-9. (a) Distribution of vicua (Vicugna vicugna) groups with respect to the laguna in the Salinas y Aguada Blanca National Reserve, Peru (a) after the wet season (April 25 to May 23, 2002) when the laguna was full and (b) during a drier period (June 25 to July 12, 2002) when the laguna was reduced to 2% of the wet season area.

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31 01234561234567891011121314151617181920Area of ringGroup density per sq km Full Almost dr y Figure 4-10. Density (groups per sq km) of vicua (Vicugna vicugna) groups around the laguna at the Salinas y Aguada Blanca National Reserve when the laguna was full and when almost dry.

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32 0102030405060708090100123456 bh fg ndf solLaguna fullLaguna almost dry<4 km>4 km>4 km<4 kmPercentage of groups Figure 4-11. Percentage of individual vicua (Vicugna vicugna) group types found <4 km and 4-20 km from the laguna when it was full (April 25 to May 23, 2002) and when almost dry (June 25 to July 12, 2002) at the Salinas y Aguada Blanca National Reserve, Peru.

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33 Table 4-1. Composition of identified vicua (Vicugna vicugna) family groups in the Salinas y Aguada Blanca National Reserve, Peru (April to July 2002). Original identification and tagging were done by CONATURA. Group composition Tagged individuals in family group Males Females 1 male, 7 females, 6 young n=14 Blue #140 Orange #18 Orange/Al #16 Orange/Al #15 1 male, 6 females, 4 young n=11 Blue/Al #142 Orange/Al #20 Purple #103 Yellow tag 1 male, 5 females, 2 young n=8 Orange #6 Orange #9 Green #151 Green #156 Yellow tag 1 male, 3 female, 2 young n=6 Blue #142 Blue #144 Green #153 Orange #11 1 male, 1 female, 0 young n=2 Orange #4 -Table 4-2. Identified male groups of vicua (Vicugna vicugna) in the Salinas y Aguada Blanca National Reserve, Peru (April to July 2002). Original identification and tagging were done by CONATURA. Group type Group size Identified tagged individuals Solitary male n = 1 Blue #127 Solitary male n = 1 Blue #146 Bachelor herd variable White #52 White # 54 White #55 White #58 White #59 White #60 White #61 White #64

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34 that were followed (Figure 4-12) and a high degree of site fidelity on behalf of the identified male over the duration of the study. However, there was distributional overlap with other non-identified groups. The mean age of males from each group type was extracted from the CONATURA database on the identified males observed during the 2002 field survey. Solitary males averaged 11.5 yr; while young males (<3 yr) were found in bachelor herds, and males in family groups averaged 9.5 yr. The latter males made up the effective breeding population, as access to females is limited to those in a males defended family group (Koford, 1957; Vila and Cassini, 1994). The histories of the tagged males followed in this study were also extracted from the CONATURA database to establish trends in age, group type, and group size as well as site fidelity to the laguna (Table 4-3). Identified vicuas were repeatedly found at the laguna for up to 5 yr, indicating a high degree of site fidelity to the laguna. Within the social context of establishing corrals, data on the distances that vicuas move is critical. Distances moved by the followed groups were compared between the morning and evening. In the morning, vicuas moved an average distance of 713 m hr-1 (SE+/-162.70), while they only moved 508 m hr-1 (SE+/45.67) in the afternoon. Tagged females observed in identified family groups were extracted from the CONATURA database to explore the validity of group stability as assumed in the literature. Our study indicates that females vary in strategies. Some exhibited a high degree of mobility among family groups, and others remained with the same male over a long period. For example, tagged female orange #17 was observed to change family groups four times within 2 yr (Table 4-4). She was observed with male blue/al #142,

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35 switched to male blue #146, and then returned to male blue/al #142. In the same month, she switched again to blue #143 and to a family group with no Male # 146 Male # 2 Male # 143 Male # 6 Male # 140 Male # 142 Bachelor Herd Laguna Figure 4-12. Spatial patterns of followed vicua (Vicugna vicugna) groups around the laguna at Salinas y Aguada Blanca National Reserve, Peru. A low degree of spatial overlap was observed between family groups in regions <4 km from the laguna.

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36 Table4-3. Life history characteristics of tagged male vicua (Vicugna vicugna) in the Salinas y Aguada Blanca National Reserve, Peru in 2002. Characteristics extracted from the CONATURA database. Individual Year tagged Longevity at laguna # years observed Estimated age Group size Group type Blue 127 1997 5 2 12 1 Solitary Blue 146 1997 5 4 11 1 Soitary Blue 140 2000 2 2 13 14 Family group Blue 142 2000 2 2 10 11 Family group Blue 143 2000 2 2 9 8 Family group Orange 4 1999 3 2 9 2 Family group Orange 6 1999 3 4 8 6 Family group White 52 2000 2 2 2 3 to 28 Bachelor herd White 54 2000 2 2 2 3 to 28 Bachelor herd White 55 2000 2 2 2 3 to 28 Bachelor herd White 58 2000 2 2 2 3 to 28 Bachelor herd White 59 2000 2 2 2 3 to 28 Bachelor herd White 60 2000 2 2 2 3 to 28 Bachelor herd White 61 2000 2 2 2 3 to 28 Bachelor herd White 64 2000 2 2 2 3 to 28 Bachelor herd Table4-4. Examples of movement within female vicua (Vicugna vicugna) among family groups in Salinas y Aguada Blanca National Reserve, Peru from 1999 to 2002. Female Orange #9 remained with the same group over a 4-yr period. Female Orange #17 switched among four groups. Dates tagged males and females observed from 1999-2001 extracted from the CONATURA database. Female Date Male Total group size Orange #9 21-Dec-99 Orange #6 2 20-Apr-00 Orange #6 3 22-Jun-00 Orange #6 7 13-Dec-00 Orange #6 6 23-May-02 Orange #6 6 26-Jun-02 Orange #6 8 Orange #17 24-Nov-00 Blue/Al #142 2 13-Dec-00 Blue #146 2 30-Mar-01 Blue #146 2 11-Apr-02 Blue/Al #142 11 15-Apr-02 Blue #143 7 25-Apr-02 Blue #143 8 24-May-02 No Marks 4 other individuals marked the next month. In contrast, tagged female orange #9 was consistently observed with the same male (orange #6) over 3 yr.

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CHAPTER 5 DISCUSSION General Ecology The vicua of Salinas y Aguada Blanca were dominated by family groups that move daily and seasonally, but show minimal spatial overlap. Water seems to be a fundamental environmental character influencing movement patterns and seasonal changes in local density. These characteristics are very important when considering the potential of current management initiatives. They also provide basic descriptors of the ecology of vicua and are useful in establishing interdemic patterns of variation. The main objective of the study was to offer baseline ecological data on population parameters of free-ranging vicua, and compare this data with other studies on vicua and ungulates. The population characteristics presented in this study included: 1) group size and group type composition within the population, 2) density and distribution, 3) movement patterns in response to water distribution, and 4) spatial patterns and intergroup movement. Group Size and Type Composition I used four categories to describe basic social units in the vicua of Salinas y Aguada Blanca. These included: family groups, bachelor herds, solitary males, and non-defined groups. These categories were created based on the different roles males play within the population. Males in family groups are potentially the only males that are breeding; thus family groups make up the effective breeding population. In Salinas y Aguada Blanca, 67% of all vicua groups were in family groups, while only 12% were 37

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38 male groups, indicating that approximately two-thirds of the males in this population maintain a group of females and are breeding. These trends in group type composition support an earlier study on the socioecology of vicuas. In his long-term study of vicuas in the Pampa Galeras reserve, Franklin (1983) described vicua population structure in relation to behavior and spatial distribution. He recognized three different vicua group categories: family groups, male groups, and solo males. In this study, Franklin found that 75% of all males were in territorial groups while only 24% were in other male groups. Franklin also suggested that male vicua in family groups are territorial and maintain boundary-oriented territories. The limited overlap in range use documented in my study also suggest that over half of the males in this vicua population may be territorial. These results on group type composition in vicua populations coincide with the general population characteristics of ungulates: within the social distribution of ungulates, generally 2/3 of the adult males within a population are territorial (Owen-Smith, 1977). Mating Strategies of the Vicua The life history strategies of ungulates evolved within the context of density dependence and carrying capacity. As a result, strategies common to most ungulates include the asymmetry of reproductive success between the sexes, a dimorphism in body size, and a skewed sex ratio towards females (McCullough, 1999). A skewed sex ratio of 1.5 females: 1 male was found in the vicua population at the Salinas y Aguada Blanca National Reserve. Similarly, the vicua in Pampa Galeras exhibited a similarly skewed sex ratio of 1.2 females to 1 male (Franklin, 1983). However, in both these studies, it is uncertain how this sex ratio changes with age. In many mammals, the sex ratio will be

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39 skewed towards males at birth, then shift towards females in older age groups (Smith, 1996). Earlier studies report resource-defense polygyny as the mating system of the vicua (Bosch and Svendsen, 1987; Franklin, 1983; Vila, 1992). The structure of the family group includes one adult dominant male, a number of females, and that years young. Our study detected no relationship between the number of males and group size, providing evidence that the population in Salinas y Aguada Blanca also exist in polygynous herds. Group Composition Within Family Groups Data collected during the field study suggest that vicua populations adopt a rigid family group structure independent of environmental gradients and varying management strategies. The mean group composition at the Salinas y Aguada Blanca National Reserve (1 male, 3.7 females, and 1.6 young) falls within the range reported in earlier studies. In Abrapampa Argentina, the mean group size was 1 male, 3-4 females, and 2-3 young (Vila, 1995), while a study done by the same author in 1992 in Catamarca, Argentina reported a mean group composition to be 1 male, 3.6 females, and 1.9 young. Both of these sites have dry and harsh climates, similar to the Salinas y Aguada Blanca National Reserve. In the Lauca National Park in Las Cuevas, Chile, the mean group composition was 1 male, 3.1 females, and 1.6 young. In Pampa Galeras, a region with a high density of vicua (67.9 vicua/km2) and with more sources of water then my field site, the ratio was 1 male, 3 females, and 2 young (Franklin, 1983). The mean group size among vicuas in corralled habitat in the humid puna where water is not such a limiting resource was also 1 male, 3.4 females, and 1.7 young.

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40 Group Size and Density The average group size of family groups (antilog mean=5.6) in the Salinas y Aguada Blanca National Reserve is similar to the averages reported from other studies. In Pampa Galeras, the average group size was 6, (Franklin, 1983), and Vila and Roig (1992) in the Laguna Blanca National Reserve, Argentina, found the average group size to be 6 to7. Group size of bachelor herds ranged from 8 to 40 individuals in our study. The dynamic nature of bachelor herd group size seems to be characteristic of vicua as Franklin (1983) reported a range of 2 to 155 individuals and Vila and Roig, (1992) reported a range of 2 to 22 individual bachelors. There was a strong positive relationship between the number of females and the number of young and total group size. However, the relationship between the number of young in a group and the number of females is not as strong. This suggests that not every female is giving birth every year. These females may be newly-dispersed young females, or older, non-reproducing females. It could potentially be disadvantageous for the herd as a whole to support too many young simultaneously due to the increasing energy requirements of lactating females and the need to be alert more frequently (Vila and Cassini, 1994). Other alternatives to explain this relationship include the mortality of young and the inability of the female to conceive that year. No published data exist on fertility among free-ranging vicua, however, my data suggest that population growth rate and harvest models should not only incorporate a time lag, but also not assume one young per female a year as a parameter (Cattan and Glade, 1989). In general, there is a positive relationship between group size and an open habitat structure (i.e., no cover) among ungulates. Large group size among ungulates is believed to be a strategy to avoid predators and a way to optimize foraging behavior (Kie, 1999).

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41 Increasing the number of females in a family group implies a requirement for increased vigilance on behalf of the dominant male. However, in an analysis of vicua time budgets, Bosch and Svendsen (1987) reported no difference in the amount of time a male defends females between large and small family groups. In fact, the more females a male has in his harem, the more individuals there are to watch for predators/aggressors while the group feeds; thus increasing the amount of time each individual can spend foraging instead of being alert. Therefore, under the spatio-optic conditions high Andean mountain plains afford, the dominant male can increase the size of his family group by adding females with little to no increase in energetic costs in terms of vigilance. As the male vicua increases the number of individuals in his herd, the collective herd increases their overall grazing efficiency. In addition, the number of females in a group is a good predictor of the mating success among male vicua (Vila, 1995, Vila and Cassini, 1994). Thus a large group size may be optimal under these arid and open conditions. Not all vicua, however, occur in large groups. Over half (54%) of the vicua population in the Salinas y Aguada Blanca National Reserve occurred in groups with 1 to 5 individuals. Franklin (1983) suggested that a section of the vicua population that occurred more frequently in marginal habitats were temporary family groups with an average of 4 individuals (1 male: 3 females: 1 young). These marginal grazing territories were classified as feeding territories of hard bunch grasses with little to no water sources. This variation in group size in the vicuas of Salinas y Aguada Blanca National Reserve may reflect spatial variation in habitat quality and the dependency on water sources for large groups. Domestic livestock are given priority for the use of wetlands, leaving few permanent water sources, such as the laguna, for the vicua population. This may

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42 explain the tendency of large groups, especially family groups to be observed more frequently than small groups closer to the laguna, suggesting differential habitat use. Density estimates reported for vicua populations are highly variable among sites ranging from 2.3 vicua/km2 at San Guillermo in Argentina, to 67.9 vicua/km2 at Pampa Galeras, in Peru (Table 4). In our field study at Salinas y Aguada Blanca, the vicua population occurred at a density of 3.7 individuals/ km2, falling within the range reported for free-ranging vicua. The source of variation among vicua population estimates may reflect density increases in corralled populations, temporal variation due to poaching and other anthropogenic factors, natural decline, or differences in sampling techniques among sites. Other managed and non-managed ungulates also exist at very different densities. Thompsons gazelle (Gazella thomsoni), another species adapted to harsh environments exists locally at the Ngorongoro crater in Tanzania at 70 individuals/ km2, but more generally at 14 individuals/ km2 (McCullough, 1999). Ungulates inhabit a tremendous diversity of ecosystems, and therefore face different kinds of environmental constraints. However, they all demonstrate territorial behavior and exist in polygynous herds for at least part of the year. Although no direct comparisons can be made to this study, these data in conjunction with published estimates for polygynous ungulates suggest that there is a high degree of both interspecific and intraspecific variation in the density of these mammals. In light of current management initiatives in Peru, it is also important to consider the landscape and long-term population effects of maintaining vicua at artificially high densities. Ecosystem management initiatives have only recently explored some of the

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43 positive and negative feedback systems between herbivorous ungulates and vegetation (Weisberg et al., 2002). Organisms at very high densities have to deal with limited resources and decreases in overall population health. It is notable that densities in corralled vicua populations are approximately Table 5-1. Published denisty estimates of vicua (Vicugna vicugna) along its range in South America. Author Place Year Density Korford Peruvian Andes 1957 25 vicua km-2 Corraled vicua Brack Pampas Galeras 1980 67.9 vicua km-2 Cajal San Guillermo (Argentina) 1991 2.3 vicua km-2 Free-ranging vicua Villa Catamarca (Argentina) 1992 5.5 vicua km-2 Davies Arequipa 2002 3.7 vicua km-2 10 to 27 times greater then estimates from free-ranging populations (Table 5-1). The long-term effects of such intensive use of puna habitat by vicua remains largely unknown. McCullough (1999) states that vicua are long-lived, large bodied animals with relatively low reproductive rates and a high level of female investment in raising young. He also proposes that most ungulate populations, due to density-dependent responses, have a tendency to exist at densities close to carrying capacity. Populations close to carrying capacity are prone to explosions in growth rate and subsequent over-consumption of resources that may induce a possible population crash (Caughley and Sinclair, 1994; Smith, 1996).

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44 Spatial Relationships and Intergroup Movement Distribution Patterns Understanding mechanisms underlying spatial and temporal patterns of distribution is important in making both effective conservation and management decisions. Although the Andean puna landscape appears to be barren and homogenous, different types of habitat can be identified, and differential use in this habitat has been observed among vicua (Renaudeau dArc et al., 2000). The optimality of habitat use is based on the need to balance the intake of high quality food, growth, reproduction, and the need to avoid predation, disease, etc. In the vicua population of Salinas y Aguada Blanca, the laguna is an integral feature in the landscape and seems to have a major influence on movement and spatial patterns. Areas closer to the laguna may be of higher habitat quality based on water availability for both vicua and vegetation, and on the increased density of vicua near this water source. Large groups were more frequently located near the laguna. Both the number of females and young increased as distance to the laguna decreased. However, at Salinas y Aguada Blanca, large bachelor herds (mean size >19) were also found closer to the laguna, further supporting the important role of the laguna for this population. Other studies have reported bachelor herds to move greater distances, as they forage in marginal lands and avoid lands that territorial males occupy (Franklin, 1983). This relationship to water in vicua re-asserts what is known about the relationship between ungulates and xeric habitats. In dry environments, water is a major factor limiting movement patterns and distribution. In our study we assume that habitat quality increases as distance to laguna decreases. Working from this assumption, it could be

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45 argued that medium and large groupings of vicua (n > 6) tend to be found more frequently in areas close to the laguna, and possibly maintain a low degree of spatial overlap around the laguna because of the high habitat quality. Although there is more competition and possibly opportunity for predation, conflict, and invasion, there are also more vigilant individuals in habitat of higher quality. In addition, less time and energy is spent traveling to a water source if a territory is maintained nearby. Family groups were the dominant group type in regions near the laguna; and all large family groups (n > 6) were found less than 3 km from the center of the laguna. The analysis of the socioecology of the vicua in Pampa Galeras conducted by Franklin (1983) provides a useful comparison. The habitats are relatively comparable as both have similar annual precipitation (Pampas Galeras: 281 to 742 mm; Salinas y Aguada Blanca: 200 to 1,000 mm) and vegetation structure. In his study, Franklin found that the permanent territorial family group consisted of an average of 6 individuals with a mean group composition of 1 male, 3 females, and 2 young. These were groups that had well-established high-quality territories. High quality territories were those that contained water sources and vegetation associated with puna swampland such as quinsa, chiula, and mojadales. Smaller family groups with 3-4 individuals were often found in territories with no source of free water. This poor quality habitat was characterized by the overbearing presence of roqueo, piedras de loma and the ichu grass, vegetation that neither requires nor contains much water. Within the context of corral installation, the mechanisms behind differential habitat use among vicua groups are important to explore. Increasing density either through the use of management and husbandry techniques or by eliminating dispersal, despite

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46 population growth, could put greater pressure on these vicua groups and the puna ecosystem. Large groups could potentially be forced to reduce their feeding territories due to over crowding, especially directly around the laguna. This could in turn, reduce mean group size. Reducing mean group size may result in increased pressure on the remaining individuals in the family group to spend more time being vigilant and less time feeding. This process of overall group size reduction may already be happening in the Salinas y Aguada Blanca National Reserve as 54% of all vicua groups have only 1-5 individuals. These animals are already relegated to habitat patches that appear to be of poor quality and little to no water. The wetland areas (Bofedales, Mojadales) are dedicated to supporting the hundreds of alpaca and sheep from the community. Not only are the vicua prohibited access to these water and vegetative sources by the presence of livestock, but are also deterred by the paved road separating the regions where the vicua are and the wetland areas. Seasonal Changes in Distribution and Density The vicua must drink water regularly, even daily, to sustain their biological processes. However, these ungulates have evolved within this context of severe water limitation, and have adopted certain behaviors to cope with water restriction. Many of these adaptations may be behaviors that are expressed in movement patterns such as spatial distribution and group composition in relationship to water. For example, a consistent pattern of movement from the edges of the mountains in the morning to water sources in the afternoon has been observed in various habitats across the vicuas range (Vila, 1992; Franklin, 1983; Vila and Cassini, 1993). To further explore this relationship to water, the size of the laguna was used as a dryness index. Although these two sampling periods may not represent two different seasons at their extremes (extreme wet

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47 vs. extreme dry), there was a definate decrease in the water available. The perimeter of the laguna was measured in April and decreased to 2% of its size when it was measured again at the end of June. In the Salinas y Aguada Blanca National Reserve, we detected a marked decrease in the average distance of vicua from the center of the laguna between the end of the rainy season (May) and the onset of the dry season (June). The vicua were approximately 1.6 km closer to the laguna when it is almost dry. However, the transect included territories that were > 10 km from the laguna. Due to potential habitat fragmentation, the vicua groups on these territories could potentially be using a different water source. After these observations (distance > 10 km) were taken out of the analysis, vicua groups were found to be approximately 1.8 km closer to the laguna when it was dry then when it was full. The distance of vicua groups from the laguna is almost 2 km less at the onset of the dry season then at the end of the wet season. This marked decrease coincided temporally with a decrease in precipitation in the whole region, as expressed by a decrease in water distribution and water levels at the laguna. The decrease in the average distance of groups from the laguna was reflected in an increase in vicua density near the laguna in the dry season. A complementary relationship between vicuas and water distribution in the wet and dry season was also observed in a similar habitat, a high, dry Andean grassland of Argentina by Vila and Cassini, (1993). Their probability of finding individual vicuas near water remained constant throughout the dry season, but decreased in the afternoon during the wet season. In our study, the average distance of vicua groups from the center of the laguna decreased by 729.53 m in July. This was most likely due to the snow fall that occurred

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48 throughout the month of July. Although the snow would melt by the afternoon, it served to distribute water along the entire pampa and surrounding foothills. Vicua cannot depend solely on the water from snowfall; however, the increased availability of water gives both the vegetation a boost, and helps vicua with their intake of water as it collects on the plants they graze on. The results of this study suggest that movement behaviors in response to changing water distribution is a key element for survival in vicua. Larger family groups and bachelor herds occupy the habitat immediately around the laguna more frequently then smaller groups. I also detected differential habitat use among vicua group types between the wet and dry seasons. How different group types of vicua use the patches of habitat directly around the laguna offers some insight to adaptive behaviors and population dynamics in free-ranging vicua. For example, in general, over half of the family groups were found <2 km from the center of the laguna, and 100% of the bachelor herds were found in this distance class as well. However, non-defined groups (antilog mean =2.9), were observed to be more evenly distributed along the study area. This suggests that these groups could be occupying dry, marginal habitat. This decrease in representation of non-defined groups (both during the dry and wet season) in habitat directly around the laguna further supports Franklins notion that these are small temporary family groups occupying marginal areas. When the laguna was full, family groups were observed to dominate within the first 1-km from the center of the laguna. This was followed by the bachelor herds, the solitary males, and lastly, the non-defined groups. When the laguna was almost dry, there was an increase in the number of bachelor herds within this first 1-km radius. Although it has been proposed that bachelor herds move greater distances, there was a definite trend in

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49 movement towards the laguna. Most bachelor herds in this study were considered large (17.5 individuals) and were observed a distance of 3 km or less from the center of the laguna. This suggests that group size may limit how far vicua groups can travel and forage from the laguna. Franklin (1983) also reported seasonal changes in vicua density. However, he attributed these fluctuations to expulsion of young during certain seasons. In Pampa Galeras, the young from family groups are expelled between October and March, decreasing family group density. From April to May, there is an observed increase in density, corresponding to the birthing season. From June to August in Pampa Galeras, there is another unexplained decrease in density. Although our field study only covers a fraction of the time data was collected on vicua density in Pampa Galeras, it could offer a potential explanation to the observed decrease in density as vicua groups are observed to limit their movements around water holes. Spatial Patterns and Inter-Group Movement Identified family groups were followed to gain insight into movement patterns at the group and individual level. Earlier studies e.g., (Franklin, 1983) estimated home range and relative territory size, however, these studies did not follow individually marked individuals over long periods of time. Our study showed that vicua groups exhibit a low degree of spatial overlap around the laguna. The first formal study on vicuas conducted by Koford (1957) in the Peruvian Andes suggested that male vicua were highly territorial and would frequently engage in fights and chases with intruding males. Territoriality is a common characteristic among ungulates and leads ultimately to differential mating and reproductive success among members within a population (Owen-Smith, 1977). In Pampa Galeras, Franklin (1983)

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50 also proposed territoriality as a characteristic of the vicua; he observed males from family herds frequently participating in antagonistic interactions with males from other groups. He classified two different habitat types as vicua territories. One was a sleeping territory on flattened ridges, and the other was a feeding territory on the lower slopes and flatlands with higher vegetative biomass. He found this territoriality to be based on its regular defense and on the maintenance of boundaries on behalf of the dominant male in the family group. In addition, Franklin found that resident neighbors respected these boundaries, while outside vicua groups did not. The nature of this territoriality, found both from Franklins study and from the data collected at the Salinas y Aguada Blanca National Reserve, appears to be boundaryoriented (Owen-Smith, 1977). In our study, the low degree of spatial overlap between family groups, and to a lesser degree, between solitary males and bachelor herds, suggests that vicua groups are maintaining their daytime feeding territories at the laguna. Mean age of identified males found at laguna in 2002 There are no published data describing age structure within vicua populations. The age a male establishes residency on a territory and the length of time that male retains a territory remains largely unknown (Bosch and Svendsen, 1987). At the time of tagging during the chaccu, CONATURA estimates an individuals age by the wear on their teeth. Therefore, I was able to compile preliminary data on relationships among age classes, population structure, and movement. Our results indicate a specific age-class structure within the vicua population, with the bachelor herds being the youngest, males in family groups in a middle range, and solitary males in the oldest age class. This finding is a departure from the observations at Pampa Galeras. Franklin (1983) suggested that solitary males are sexually mature attempting to establish territories. However, age

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51 estimates found in this population suggest that solitary males may be in a stage of senescence, not in dispersal and establishment. In addition, solitary males were also found at farther distances from the laguna and in more marginal habitat patches than other group types when water was scarce. Site fidelity and dispersal Not all tagged individuals were observed at the laguna during the course of the study in 2002. This may reflect observation bias, death or, dispersal. We recorded only one case of dispersal during our study. A solitary male, (Blue 127; age=12) was observed approximately 17 km from the laguna where it was originally tagged. Although one example of dispersal is not enough to make any conclusions about movements in free-ranging vicua, it does suggest that the older, solitary males may have larger home ranges or dispersal distances than other males found in family groups or bachelor herds. The clustering behavior of young males around the laguna appears to be counter-intuitive. Earlier studies (Koford, 1957; Franklin, 1983) suggest that these young males disperse long distances to establish their own territories and family groups, thereby encouraging genetic shuffling. The distance traveled by bachelor groups and their spatial distribution suggest that they are potentially optimizing their opportunities to obtain females at higher densities found around the laguna, while simultaneously being limited in movement due to the scarcity of water. Our data suggest that median-aged males maintain territories for several years. Although not every tagged male was observed during the study, there is evidence that these males have been coming to the laguna consistently for several years and exhibit a high degree of site tenacity.

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52 Distance moved from family groups How much space a vicua group needs to fulfill its daily requirements is a central question within the context of the installation of corrals. The size of corrals will vary dramatically with habitat quality, density, water resources, and food supply. To begin to answer this question, vicua family groups were followed, and their movements were documented. The results suggest that vicua do not move very far, and that the corrals would not limit their movement greatly. However, observations were taken only in regions at or adjacent to the laguna, and did not represent movement across the vicuas entire home range. In addition, differential movement patterns were observed as vicua groups moved greater distances in the morning then in the afternoon. Vicua movement was high in the morning while the groups were heading toward the water source as compared to the afternoon when they move back to their sleeping areas. These results correspond well with the study of Vila and Roig (1992) that focused on daily movements and behavior of family groups. Their study found that vicuas moved greater distances in the morning towards the water source and grazed in the afternoon. Description of shifts in group composition In this study males tended to maintain spatially discrete ranges around the laguna, that remained relatively constant for the 4 months of observation. I hypothesize that males establish territories, and females will occasionally switch family groups. In this way, there is the potential for genetic shuffling between resident vicuas in this population, while maintaining a stable spatial distribution over time. Other studies have assumed that groups are basically stable, comprising of the same individuals over time (Franklin, 1983; Vila, 1995). Females in my study were found to adopt different mating

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53 strategies. Some were sighted over several years with the same identified male, and other females switch between different males at different rates. Female vicuas exhibit iteroparous reproduction, giving live birth to one young at a time, repeatedly during her lifespan. Young ruminants depend completely on their mothers milk for the required amino acids, vitamins and the inoculation of rumen bacteria to develop ruminant fermentation (van Soest, 1994). Thus, the female vicua has different energy requirements and constraints in maximizing reproductive success then males (Kie, 1999; Vila, 1995). The number of family groups a female belongs to in her lifespan may not affect her fitness. What matters more for her reproductive success is the habitat quality of the dominant male in her family group, i.e., the quality, quantity, and heterogeneity of the vegetation for embryo development, lactation, and the continual growth of her young (Robinson et al., 1999).

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CHAPTER 6 IMPLICATIONS FOR CONSERVATION An increase in the numbers of vicua has been reported over the last 30 years in Peru. There were reportedly only 10,000 vicuas left in 1964 (CONACS, 1997); as a result of intensive management practices, the vicua population in Peru reportedly increased to 150,000 individuals by 2000 (Lichtenstein et al., 2002). Despite the increase in numbers, the distribution of the vicua has been dramatically reduced mainly to the three national reserves in Pampa Galeras, Huaraz, and Salinas y Aguada Blanca. Vicua no longer exist along the entire historic range in Peru (Figure 6-1). It is unknown if the increase in overall vicua numbers is due to strict protectionist measures, or to local dynamics in a few communities that have dedicated themselves to promoting the growth rate among vicua to facilitate rational use of vicua fiber as a sustainable management strategy. Thus, increases in numbers may potentially only reflect a geographically limited increase in population density. This centralized increase in vicua numbers is exacerbated by new management initiatives that involve the installation of corrals. This may potentially change the distribution of vicuas and alter important ecological characteristics of their population ecology. In addition, it invariably segregates potentially inter-breeding vicua populations, encouraging homozygosity. It is important to understand the mechanisms of this population growth among vicua, because changes in social behavior associated with changes in density may change patterns of habitat use, and may negatively alter the puna ecosystem. 54

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55 Study site Figure 6-1. Historic range of the vicua (Vicugna vicugna) in South American Andes. Figure adapted from The Camelid (W. Ross Cockrill, 1979). Information on behavior patterns of free-ranging vicua is scarce. Much of the data on their habitat requirements are from studies that focused solely on biomass, husbandry, and management (Wilson, 1994). Much of the current data on vicua ecology comes from highly managed vicuas in corrals and does not directly answer any of the questions surrounding the issue of limiting movement in regard to group composition, movement patterns, spacing patterns, and territoriality. Like other density-dependent ungulates, the vicuas may exhibit population oscillations. Therefore great precaution must be taken in determining corral placement, size, and the number of vicuas to maintain in those corrals. If the population falls below a viable level, there could be detrimental consequences for the vicua population as a result of decreased genetic heterogeneity. For example, populations with low genetic diversity are at risk of lowered immunity to the spread of disease throughout the entire

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56 population, and more vulnerable to stochastic abiotic factors (Caro, 1998). On the other hand, if the population grows rapidly, the puna ecosystem could be at risk of over-grazing, homogenous vegetative composition, soil erosion, and desertification. Currently, the focus in conservation and sustainable development programs seems to be in increasing the population of vicua. Much effort has gone into determining how many vicua can subsist given forage production, given that their use of fibrous feeds is extremely efficient because of their highly specialized mode of fore-gut digestion (Cajal, 1991; Cueto et al., 1985). However, there are no studies that estimate how long the puna ecosystem can maintain a high density of vicua before nutrient levels in the soil decline, vegetative structure is degraded, or hydrological regimes altered. The ability of vicua to have access to water and forage of sufficient quality and quantity is essential in maintaining a viable vicua population. The data suggest that vicua in the Salinas y Aguada Blanca National Reserve respond behaviorally to harsh conditions by moving in response to resource availibility. Limiting vicua movement through the implementation of corrals may have negative impacts for both the health of the vicuas and their habitats. The nutritional quality and quantity of biomass in puna ecosystems has been reported to decline from the end of the wet season through the dry winter (Pfister et al., 1989). This impacts vicua and their movement significantly as the typical functional response in ruminants is to increase the time spent foraging to compensate for the lowered nutritional value. My results show decreased movement centered around the laguna during the dry season. This limitation in movement around the laguna may be a result of an increased water requirement. Or, there may be more vegetation of higher quality around the water

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57 source available, decreasing the energy needed to satisfy water and nutritional requirements. There are other constraints operating in conjunction with the decrease of water availability and changes in nutritional quantity and quality. As temperatures decrease during the dry winter, there is a higher cost of thermoregulation (Vila and Cassini, 1993). Movement in search of forage among vicua also results in heat production and lowered energy retention, especially when both the quantity and quality of forage is reduced (Murray, 1991). Limiting movement through the installation of corrals during the dry season may not have a significant impact on the social structure of the vicua, or on natural movement patterns. However, overgrazing in these areas near the laguna could have significant long-term consequences on the quality of forage for subsequent years. In addition, the dry season is also marked by frequent snowfall, as observed in our study from July 10 to July 12. At this time, vicua groups were found father from the water source, and most identified family groups were not found at the laguna at all. The increased distribution of water possibly allowed for greater movement, subsequently releasing local vegetation around the laguna from foraging pressure. The installation of corrals would obviously limit a functional response among vicua to availability to resources. Another potential consequence of corral installation in the Salinas y Aguada Blanca National Reserve is related to land use and the geographic position of certain key landscape features. The area where vicua are found is bisected by the paved road running from Arequipa to the Colca and Juliaca-Cuzco and presents a potential source of

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58 vicua mortality. There were several instances during the study where vicua groups were observed crossing this highway in search of water in the reserve. Placing a corral in this reserve would exclude other vicua groups traveling in search of water and forcing them to find other sources, either in regions near domestic livestock, or in farther regions. This could potentially decrease the amount of vicuas available for shearing during the roundup. The primary aim of the rational use of the vicua as a sustainable management program is two-fold. First, it claims to protect the vicua, while secondarily providing a legal, alternate source of income for impoverished Andean communities. A major threat to the vicua is anthropogenic in nature as Andean community members claim that there is competition for feed resources between naturally occurring vicua and their domestic livestock (Urquieta et al., 1994). The implementation of corrals has been hailed as a viable compromise that will alleviate this competition, provide protection for wild vicuas and facilitate the capture process for shearing. However, these corrals may serve to alter the puna ecosystem and the social and foraging behavior of the vicua that may lead to negative consequences.

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CHAPTER 7 CONCLUSIONS Baseline data on the population characteristics of free-ranging vicua populations are critical to the evaluation of effective conservation and management practices and to our understanding of the importance of local habitat features to population density and distribution patterns. Group characteristics of the vicua of the Salinas y Aguada Blanca National Reserve are comparable to studies of other free-ranging vicua, with densities notably lower than corralled populations. The changes in distribution and density over the 4-month study suggest that the location of a permanent water source may be an integral factor contributing to patterns of distribution and movement over the seasonal cycle. The major conclusions drawn from this study are summarized below. 1. In the Salinas y Aguada Blanca National Reserve, the mean family group composition is 1 male, 3.7 females, and 1.6 young. These figures fall within the range of other studies that include both corralled and free-ranging populations, suggesting that the vicua have a rigid family group structure within different populations that occur across ecological gradients and management programs. 2. In my study average density was 3.7 vicuas/km2. This falls within the range of studies of free-ranging vicua and is much lower than corral estimates. 3. Water distribution and availability seem to have a major effect on vicua movement. Groups were found to be 2 km closer to the laguna in drier conditions. This may have management implications, as water is a major limiting factor in the puna. Although domestic livestock and vicua herds have not been found to compete for forage, they could be competing for water resources. 4. Overall, large groups and large family groups are found closer to the laguna. This indicates that large groups are limited in their movements by water availability, and that large groups tend to establish territories in higher quality habitats near the laguna. 59

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60 5. The data on the Salinas y Aguada Blanca vicua in combination with other studies on ungulates, suggest that large family groups in open habitats is an optimal strategy for feeding and reproducing, as smaller vicua groups are more frequently in marginal habitats. 6. There may be a rigid population structure and family group structure within vicua populations found in different habitats. Several group characteristics such as group type distribution in the population, group size, and group composition remained stable despite changing water distribution. 7. There was a low degree of spatial overlap around laguna. This suggests that the dominant males in family groups are maintaining boundaries in relation to other family groups when traveling to the laguna. This also provides further evidence for territoriality among males in family groups. 8. Individual vicuas and groups were found to exhibit differential movement patterns throughout the day. Vicuas move greater distances in the morning, presumably to reach the laguna by midday. The rate of movement decreases in the afternoon. These differential movement patterns are important when estimating how far a vicua will move, and subsequently how large their territory requirements are in the context of corral initiatives. 9. Our study provides evidence that some females will stay with the same male in a family group for an extended period of time, while others will switch annually among family groups. 10. With the use of age estimations provided by the CONATURA database, an age class structure was found in the vicua in the Salinas y Aguada Blanca National Reserve. Bachelor herds were found to be young males that are non-territorial dispersers. Other studies report that solitary males are in process of establishing their territories, however, this study indicates that solitary males are older and in a stage of senescence. Although there was overlap, the mean age of solitary males was higher then those in family groups.

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61 LIST OF REFERENCES Bosch, P. C. and G. Svendsen. 1987. Behavior of male and fema le vicuna as it related to reproductive effort. Journal of Mammalogy 62: 425-429. Brack, A. 1980. Situacin actual de la poblac in de vicuas en Pampa Galeras y zonas aledaas y recomendaciones para su ma nejo. Proyecto Especial Utilizacin Racional de la Vicua. Ministerio de Agricultura y Alimentacin, Lima. Buckland, S.T, D. R. Anderson, K. P. Burnham, and J. L. Laake. 1993. Distance Sampling. Chapman and Hall London. Cajal, J. 1991. An integrated approach to the management of wild camelids in Argentina. In: Latin American Mammology. History, Biodiversity and Conservation. M. A. Mares and D. J. Schmidly (eds.). University of Oklahoma Press, Norman. Caro, T. (ed). 1998. Behavioral Ecology a nd Conservation Biology. Oxford University Press, New York Cattan, P and A. Glade. 1989. Management of the Vicugna vicugna in Chile: Use of a matrix model to assess harvest rates. Biological Cons ervation 49 131-140. Caughley, G. and A. R. E. Sinclair. 1994. Wildlife Ecology and Management Blackwell Science Oxford. Caviedes, C. and G. Knapp. 1995. South Am erica. Prentice Hall, Englewood Cliffs, New Jersey. Chapman, C. A., S. R. Balcomb, T. R. Gilesp ie, J. P. Skorupa, and T. T. Struhsaker. 2000. Long-term effects of logging on African primate communities: A 28-year comparison from Kibale National Park, Uganda. Conservation Biology 14: 207217. Cockril, W. R. 1979. The Camelid. An All Purpose Animal. Volume I. Proceedings of the Khartoum Workshop on Camels. Scan dinavian Institute of African Studies, Uppsala. Consejo Nacional de Camelidos Sudamericanos (CONACS) website maintained by Universidad Agraria de la Molina. 1997. Antonio Tovar conacs@amauta.rcp.net.pe. (9-11-01).

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62 Convention for the Conservation and Management of the Vicuna. Website maintained by Center for International Earth Science In formation Network. h ttp://sedac.ciesin.org. (9-11-01) Cueto, L. J, C. F. Ponce, E. Cardich, and M. Rios. 1985. Management of vicua: Its contribution to rural development in the High Andes of Peru. FAO Conservation Guide No. 11 Franklin, W. L. 1978. Socioecology of the vicuna. Dissertation, Ut ah State University. Franklin, W. L. 1983. Contrasting socioecol ogies of South Americas wild camelids: The vicua and the guanaco. In: Advances in the Study of Mammalian Behavior. J.F. Eisenberg and Devra G. Kleiman (e ds). Special publication No. 7. The American Society of Mammol ogists, Stillwater, OK. Hixon, M. A. 1980. Food production and competitor density as the determinants of feeding territory size. Am erican Naturalist 115: 510-530. Hoffman, R. K., K. Otte, C. F. Ponce, and M. A. Rios. 1983. El Manejo de la Vicua Silvestre. Tomo I. Eschborn. Kie, J. G. 1999. Optimal foraging behavior a nd risk of predation: effects on behavior and social structure in ungulates. Journal of Mammology 80: 1114-1129. Koford, C. B. 1957. The vicua and th e puna. Ecological M onographs 27: 153-219. Lichtenstein, G., F. Oribe, M. Grieg-Gran, and S. Mazzucchelli. 2002. Manejo comunitario de vicuas en Per. Estudio de caso del manejo comunitario de vida silvestre PIE Series No. 2 Lucherini, M. 1996. Group size, spatial segregation and ac tivity of wild sympatric vicuas Vicugna vicugna and guanacos Lama guanicoe Small Ruminant Research 20:193-198. McCullough, D. R. 1999. Density dependence and life-history strate gies of ungulates. Journal of Mammology 80: 1130-1146. Menard, N. 1982. Some aspects of the socioecology of the vicua ( Lama vicugna ) Molina. Terre et la vie-revue decologie appl iquee 36:15-35. Murray, M. G. 1991. Maximizing energy reten tion in grazing ruminants. Journal of Animal Ecology 60: 1029-1045. Owen-Smith, N. 1977. On territoriality in ungul ates and an evolutionary model. The Quarterly Review of Biology 52: 1-38.

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63 Pfister, J. A., F. San Martin, L. Rosales, D. V. Sissson, E. Flores, and F. C. Bryant. 1989. Grazing behaviour of llamas, alpacas and sheep in the Andes of Peru. Applied Animal Behavioral Sciences 23: 237-246. Pulgar Vidal, J. 1996. Geografia del Per u. Las ocho regiones naturales. Dcima edicin. Peisa. Lima, Per. Rabinovich, J. E., M. Hernandez, and J. L. Cajal. 1985. A simulation model for the management of vicua populations Ecological Modeling 30: 275-295. Renaudeau dArc, N., M. H. Cassini, and B. L. Vila, 2000. Habitat use by vicuas Vicugna vicugna in the Laguna Blanca Reserve (Catam arca, Argentina). Journal of Arid Environments 46: 107-115. Robinson, J. J, K.D. Sinclair, R. D. Randel, and A. R. Sykes. 1999. Nutritional management of the female ruminant: mechanistic approaches and predictive models. In: Nutritional Ecology of He rbivores. Proceedings of the Vth International Symposium on the Nutrition of Herbivores. H-J. G. Jung and G. C. Fahey, Jr. (eds). American Society of Animal Science, Savoy, Illinois. Rundel, P.W. and B. Palma. 2000. Pres erving the unique puna ecosystems of the Andean Alitplano: A desc riptive account of Lauca Nati onal Park, Chile. Mountain Research and Development 20: 262-271. Sahley, C., J. T.Vargas, and J. Sanchez Va ldivia. 2001. Community ownership and live shearing of vicuas in Peru: Evalua ting management strategies and their sustainability. Asociacion para la inves tigacion y conservacion de la Naturaleza, CONATURA. Smith, R. L. 1996. Ecology and Field Biology. (5th ed.). Harper Collins, New York Urquieta, B, R. Cepeda, J.E. Caceras, L. A. Raggi, and J.R. Rojas. 1994. Seasonal variation in some reproductive parameters of male vicuna in the High Andes of northern Chile. Journal of Arid Environments 26: 79-87. van Soest, P. J. 1994. Nutritional Ecology of the Ruminant. (2nd ed.). Comstock Publishing Associates, a division of Corn ell Universtiy Press, Ithaca and London. Vila, B. L. 1992. Mother-offspr ing relationship in the vicuna, Vicugna vicugna (Mammalia: Camelidae). Ethology 92: 293-300. Vila, B. L. 1995. Spacing patterns within grou ps in vicuas, in relation to sex and behavior. Studies on Neotropical Fauna and Environment 30:45-51. Vila, B. L and M. H. Cassini. 1993. Summer a nd autumn activity patterns in the vicua. Studies on Neotropical Fauna and Environment 28: 251-258.

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64 Vila, B. L and M. Cassini. 1994. Time a llocation during the reproductive season in vicuas. Ethology 97: 226-235. Vila, B.L. and V.G. Roig. 1992. Diurnal movements, family groups and alertness of vicua ( Vicugna vicugna ), during the late dry season in Laguna Blanca Reserve (Catamarca, Argentina). Small Ruminant Research 7: 289-297. Weisberg, P. J., N. T. Hobbs, J. E. Ellis and M. B. Coughenour. 2002. An ecosystem approach to population management of ungulates. Journal of Environmental Management 65: 181-197. Wheeler, J. and D. Hoces. 1997. Community participation, sustainable use and vicuna conservation in Peru. Journal of Mount ain Research and Development 17: 283287. Wilson, T. R. 1994. Towards committed and collaborative camelid research. Journal of Arid Environments 26: 95-103.

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65 BIOGRAPHICAL SKETCH Jennifer Elena Davies was born on May 4, 1975, in San Diego, California. She is the only child of Thomas M. Davies Jr. and Eloisa Davies Monzn. She earned her degree of Bachelor of Arts in Nature and Culture, with a minor in Spanish from the University of California, Davis, in 1999. Ther e she translated her respect and love for the natural world to academic study and research in human-animal interactions. To continue her commitment to interdisciplinary research in the natural sciences she moved to Gainesville to enroll in graduate studies in the Center for Latin American Studies/ Tropical Conservation and Development in the fall of 2001.


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Permanent Link: http://ufdc.ufl.edu/UFE0002502/00001

Material Information

Title: Population Ecology of the Vicuna (Vicugna, vicugna) at the Salinas y Aguada Blanca National Reserve, Arequipa, Peru: Baseline Data for Sustainable Management
Physical Description: Mixed Material
Language: English
Creator: Davies, Jennifer E.
Publication Date: 2003
Copyright Date: 2003

Record Information

Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
System ID: UFE0002502:00001

Permanent Link: http://ufdc.ufl.edu/UFE0002502/00001

Material Information

Title: Population Ecology of the Vicuna (Vicugna, vicugna) at the Salinas y Aguada Blanca National Reserve, Arequipa, Peru: Baseline Data for Sustainable Management
Physical Description: Mixed Material
Language: English
Creator: Davies, Jennifer E.
Publication Date: 2003
Copyright Date: 2003

Record Information

Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
System ID: UFE0002502:00001


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POPULATION ECOLOGY OF THE VICUrjA (Vlcugna vlcugna) AT THE SALINAS
Y AGUADA BLANCA NATIONAL RESERVE, AREQUIPA, PERU: BASELINE
DATA FOR SUSTAINABLE MANAGEMENT






















By


JENNIFER E. DAVIES


A THESIS PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REOUIREMENTS FOR THE DEGREE OF
MASTER OF ARTS


UNIVERSITY OF FLORIDA


2003







































Copyright 2003

by

Jennifer E. Davies





































For Murphy

















ACKNOWLEDGMENTS

I would like to thank Dr. Lauren Chapman for all the knowledge and guidance she

brought in the completion of this project. Her dedication to science, kindness, and

integrity are truly inspiring. She definitely sets the standard in achievement both

personally and professionally. I would also like to thank Dr. Charles Wood for all his

help and support during my time in Florida, and for agreeing to serve on my committee.

I would also like to thank Dr. Emilio Bruna for his support, encouragement, and

knowledge, and for graciously agreeing to serve on my committee.

In addition, I would like to thank all those at CONATURA and Dr. Catherine

Sahley for providing assistance at the field site in Arequipa, use of their database, and for

various crucial aspects of the project. The field research may not have been possible

without the invaluable friendship and support of Vajk Lucas and Mariela Caceras land

Maya) in Arequipa. They shared their home to an unexpectedly long-term visitor, who

would come home covered in dust and smelling of alpacas. I am honored to have been a

part of their lives. Comments and knowledge offered by Colin Chapman and Ron Same

were also invaluable and added greatly to the thesis. I also greatly appreciate the support

offered by those in the Tropical Conservation and Development program throughout my

years of study, namely Dr. Marianne Schmink and Hannah Covert. Technical support

land overall support) was provided by Joe Savastano. Financial support for this research

was provided by the Department of Latin American Studies, the Tropical Conservation

and Development program at the University of Florida, the Wildlife Conservation










Society, and CONATURA. Finally, I would like to thank my family and friends for their

encouragement and patience during the completion of this project.


















TABLE OF CONTENTS




ACKNOWLEDGMENTS ........................................ iv

LIST OF TABLES ................... ................... ................... .........

LIST OF FIGURES ........................................ ix

ABSTRACT................................

CHAPTER

1 INTRODUCTION ........._____ ........._____ ........._____ ..........


General Ecology ........................................
Objectives ........................................

2 STUDY SITE DESCRIPTION.............................

3 METHODS .........___ .........___ .........___ ..........


Study Design..................................
Data Analyses ................... ................... ................... ..........
Group Characteristics ................... ................... ................... ..........
Distribution and Density.................................
Density ........................................
Density and distribution relative to the laguna. ................... ................... ......13
Group size distribution relative to the laguna ................... ................... ........13
Responses to Changes in Water Distribution ................... ................... ................14
Spatial Patterns and Inter-Group Movement ............... ................... .................15

4 RESULTS .........___ .........___ .........___ ..........


General Ecology ................... ................... ................... ..........
Group Size and Population Composition ................... ................... ................... ...16
Group Size and Composition by Group Type ............. ............. ...............17
Spatial Relationships and Intergroup Movement ................... ................... .................18
Density and Distribution............................
Density and distribution relative to the laguna. ................... ................... ......18
Group size distribution relative to the laguna ................... ................... ........19











Group composition relative to distance from laguna ................... ..........
Responses to Changes in Water Distribution ................... ................... ................24
Average distance from laguna as a function of changes in water
distribution ................... ................... ................... ...........
Density and distribution as a function of changes in water distribution ......29
Spatial Patterns and Inter-Group Movement ................... ................... .................29

5 DISCUSSION..............................

General Ecology ........................................
Group Size and Type Composition ........................................
Mating Strategies of the Vicui~a ................... ................... ................... .......
Group Composition Within Family Groups ................... ................... .................. 39
Group Size and Density.................................
Spatial Relationships and Intergroup Movement ........................................
Distribution Patterns ................... ................... ................... ..........
Seasonal Changes in Distribution and Density ........................................
Spatial Patterns and Inter-Group Movement ................... ................... .................49
Mean age of identified males found at laguna in 2002 ................. ...............50
Site fidelity and dispersal ........................................
Distance moved from family groups .............. .............. ............. ...52
Description of shifts in group composition ................... ............... ............52

6 IMPLICATIONS FOR CONSERVATION ........................................

7 CONCLUSIONS ........................................

LIST OF REFERENCES .........____ .........____ .........____ .........

BIOGRAPHICAL SKETCH ........................................

















LIST OF TABLES


Table

4-1.Composition of identified vicui~a (Vlcugna vlcugna)................................

4-2.Identified male groups ofvicui~a (Vlcugna vlcugna) ........................................

4-3.Life history characteristics of tagged male vicui~a (Vlcugna vlcugna) .......................36

4-4. Examples of movement within female vicui~a among family groups. ................. .....36

5-1.Published denisty estimates of vicui~a (Vlcugna vlcugna) ................. ................. ......43

















LIST OF FIGURES


Firure

2-1.Transect design for vicui~a (Vlcugna vlcugna) survey. ........................................

4-1.Percentage of group types in the vicui~a (Vlcugna vlcugna) population ................... ..17

4-2.Vicui~a (Vlcugna vlcugna) group density ........................................

4-3.Distribution of vicui~a (Vlcugna vlcugna) group types around the laguna ..................2 1

4-4. Size of vicui~a (Vlcugna vlcugna) groups in relation to the laguna ................... ..........23

4-5.The number of females in family groups of vicui~a (Vlcugna vlcugna) ................... ...25

4-6. The number of young vicui~a (Vlcugna vlcugna) in relation to distance. ................... .26

4-7. The number of males in vicui~a (Vlcugna vlcugna) groups ................... ................... ...27

4-8. Average distance of vicui~a (Vlcugna vlcugna) groups ................... ................... .........28

4-9.Distribution ofvicui~a (Vlcugna vlcugna) groups ........................................

4-1O.Density (groups per sq km) of vicui~a (Vlcugna vlcugna) ................... ................... ...31

4-11.Percentage of individual vicui~a (Vlcugna vlcugna) group types ................... ...........32

4-12.Spatial patterns of followed vicui~a (Vlcugna vlcugna) groups.................................3

6-1.Historic range of the vicui~a (Vlcugna vlcugna) in South America.............................55

















Abstract of Thesis Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Master of Arts

POPULATION ECOLOGY OF THE VICUrjA (Vlcugna vlcugna) IN THE SALINAS Y
AGUADA BLANCA NATIONAL RESERVE, AREQUIPA, PERU: BASELINE DATA
FOR SUSTAINABLE MANAGEMENT

By

Jennifer E. Davies

December 2003

Chair: Dr. Lauren J. Chapman
Major Department: Latin American Studies

The installation of corrals to facilitate the capture and protection of the vicui~a

(Vlcugna vlcugna) has been encouraged by government agencies in Peru to meet

sustainable development objectives in rural Andean communities. Corrals may have an

impact on vicui~a populations and their habitats by altering the density of the vicui~a.

However, little is known about the home range, movement patterns, or habitat

requirements of free-ranging vicui~a, which precludes an accurate estimate of the size a

corral should be to sustainably manage these wild camelids. The objective of the thesis is

to provide quantative data on aspects of the population ecology of the vicui~a that will

have application to corral management initiatives. Repeat samples of a 38-km transect

and group follows were used to estimate four major ecological traits of vicui~a groups in

the Salinas y Aguada Blanca National Reserve in Arequipa, Peru. These included 1)

group characteristics, 2) movement patterns, 3) responses to declining water availability

in the major laguna for vicui~a in the reserve, and 4) spatial patterns and inter-group










movement. Analyses of these four ecological traits were used to derive patterns of

habitat use, the relationship to water resources, and density variation across space and

time. Family groups made up 67% of all groups within the population, had an average

group size of 5.6 individuals, and had a group composition of 1 male, 3.7 females, and

1.6 young. Overall density of vicui~a in the study area was 3.7 vicui~a km2. Water

distribution and availability had a major impact on vicui~a movement, but no detectable

effect on group and population composition. Group density was highest < 4 km from the

laguna, suggesting that the water source was a key feature in the vicui~a's habitat.

Movement patterns indicated differential habitat use among group types around the

laguna. Large family groups and bachelor herds were more frequently found in

proximity to the laguna, while smaller groups (non-defined groups and solitary males)

were distributed more evenly. Family groups showed a low degree of spatial overlap

around the laguna suggesting that the family groups in the reserve are territorial and

maintain their boundaries while at the laguna.
















CHAPTER 1
INTRODUCTION

General Ecology

The vicui~a (Vlcugna vlcugna) is a New World camelid that inhabits the high, dry

puna, or altiplano zones in the Andean mountains of Peru, Chile, Argentina, and Bolivia

between 3,700 4,900 m (Koford, 1957; Franklin, 1983; Vila, 1992). This species is one

of three wild ungulates occurring in these arid habitats. Others include the guanaco

(Lama guanlcoe) and the guemal (H~ppocamelus antlslensls). It is widely reported that

the vicui~a is a dietary specialist and therefore restricted to these elevations (Franklin,

1978; Lucherini, 1996). However, some authors suggest that since the arrival of humans

and their livestock during the Incan period, the vicui~a has been displaced to higher

elevations (Hofmann et al., 1983).

The vicui~a is sedentary and spends most of its time foraging and ruminating on the

hard bunch grasses of the puna (Bosch and Svendsen, 1987; Franklin, 1983; Lucherini,

1996; Menard, 1982; Renaudeau d'Arc et al., 2000; Vila and Cassini, 1993). Habitat use

appears restricted, as vicui~as have been found to avoid open rocky areas and only use

areas that support preferred forage (Franklin, 1983; Renaudeau d'Arc et al., 2000). This

vegetation preferred by vicui~as includes species often associated with the scarce ground

water supply found in the arid puna environment. The vicui~a requires a frequent supply

of water not obtained by either oxidative metabolism or from the food they ingest (Bosch

and Svendsen, 1987; Franklin, 1978, 1983; Menard, 1982; Vila and Roig, 1992; Vila and

Cassini, 1993). Vicui~a have been reported to exhibit daily foraging and drinking patterns










(Franklin, 1983; Renaudeau d'Arc et. al., 2000; Vila and Cassini, 1993) in response to the

drastic fluctuations in temperature, low precipitation, and medium to poor forage quality.

In their study of seasonal activity patterns Vila and Cassini (1993) found a daily peak in

the use of water sources at noon. This pattern in activity was more defined during the dry

season (Vila and Cassini, 1993), suggesting a strong relationship to the distribution of

water. However, a fine-grained analysis on water requirements and effects of water

distribution on vicui~a spatial patterns has not been undertaken.

The social organization of the vicui~a is based on family groups and male groups

(Franklin, 1983; Vila and Cassini, 1993). Several studies propose that male vicui~as in

family groups are territorial (Bosch and Svendsen, 1987; Franklin 1978, 1983; Koford,

1957; Vila and Roig, 1992), and there is evidence for variation in group type and size

with general habitat quality. However, direct links between group type and water

resources are not understood well.

A more complete picture of vicui~a population ecology is critical given increasing

pressures on their populations, and new intensive game management initiatives. Vicui~a

fiber is well suited for the drastic extremes of the Andean climate and is considered to be

the finest in the world. The fiber is 13-14 microns in diameter, which is finer than

cashmere (15-19 microns in diameter). This makes vicui~a a target species for rural

development strategies, because the fiber is viewed as such a lucrative, renewable natural

resource. In fact, vicui~a fiber has been hailed as the "gold of the Andes," a viable way to

stimulate the economies of impoverished indigenous communities in the area (Cattan and

Glade, 1989; Rabinovich et al., 1985; Sahley et al., 2001).










The main threat to vicui~a populations is the poaching for hides and their fiber

(Lichtenstein et al., 2002). This contributed to their rapid decline to an estimated 10,000

individual vicuna in the 1960's (CONACS, 1997). After being listed under Appendix I

of the CITES convention since the 1970's, the vicui~a was reclassified to the Appendix II

species list in 1995 (Wheeler and Hoces, 1997). Recent increases in the number of

vicui~a are assumed to reflect protectionist practices centered on creation of national

parks and sustainable use of wild vicui~a populations to harvest fiber (Cajal, 1991;

CONACS, 1997). Three parks were established in Peru to facilitate protection of wild

camelids: 1) Salinas y Aguada Blanca National Reserve (approximately 366,000 ha),

located just north of Arequipa, 2) Huascaran National park in Ancash (28,000 ha) and 3)

Pampas Galeras National Reserve (75,000 ha). "A vicui~a sheared is a vicui~a saved"

became the slogan of stakeholders to encourage sustainable use of the vicui~a.

Stakeholders claim that conservation objectives can be met with the protection of vicui~a

by park guards from poachers, and the sale of sheared vicui~a fiber as an economic

alternative through community initiatives. These measures have indeed been successful

in bolstering vicui~a populations in a limited number of communities such as Pampa

Galeras, but do not reflect an increase in distribution along the vicui~a's entire range in

the Andes.

In addition to the development of protected areas, new management schemes that

intensively manage dense vicui~a herds in structured corrals have been initiated. The

corrals facilitate capture and protection of vicui~a; and their utilization has been both

encouraged by the government and adopted by numerous rural communities. These

corrals are typically 500-1,000 ha in size and may contain 250-1,000 vicui~as










(Lichtenstein et al., 2002). The installation of corrals may have a detrimental impact on

the vicui~a populations and the xeric habitat they inhabit by anthropogenically altering

their natural density.

Corrals that limit movement may have an impact on the ungulate population by:

minimizing genetic dispersal, increasing the rate and incidence of disease, promoting the

breakdown of family units, decreasing breeding or infant survival, and/or altering plant

communities. However, we understand very little about the home range size, movement

patterns, or territory requirements of the vicui~a. This precludes an accurate estimate of

the size a corral should be to sustainably manage these wild camelids. Such data on

habitat requirements are crucial to the establishment of effective management programs

(Hixon, 1980; Renaudeau d'Arc et al., 2000).

Objectives

The goal of this study was to quantify key features of the population ecology of the

vicui~a and to consider implications of these characteristics for corral initiatives. Field

work was conducted in the Central Andes within the Salinas y Aguada Blanca National

Reserve (Arequipa, Peru) from April 25'" to July 12" of 2002. I assessed group

composition, density, and movement patterns of the vicui~a, as well as the aggregative

relationship to the distribution of water. My specific objectives were as follows. First, I

quantified group composition characteristics to provide baseline data for comparison with

~ee-ranging and corralled vicuna. These included: representation of age/sex classes

within groups, sex ratio, and group size. Second, I used a repeat transect design to

quantify general movement patterns and spatial distribution in a 20 km2 area that

included representative habitat types and a major water source (a laguna). Data on spatial

relationships were combined with group composition data to analyze movement patterns










for group types relative to the laguna. Third, I explored the response to changes in water

distribution by evaluating relationships between spatial distribution patterns and changes

in the size of the major water source tinder of dryness). Finally, I examined movement

patterns of individually identified vicuna groups and quantified aspects of group stability.

This permitted an evaluation of site fidelity among tagged males around the laguna,

distance traveled over time, and group stability.
















CHAPTER 2
STUDY SITE DESCRIPTION

The study was conducted between April 25 and July 12 of 2002 within the Salinas

y Aguada Blanca National Reserve in the department of Arequipa, Peru (15"45'05" and

16"19'15 south and 70"51'20" and 71"34'0" west). The reserve was created in 1979 and

contains 366,936 ha of dry puna characterized by high mountain plains, volcanoes,

hillsides, and cliffs, with elevation between 3,400 to over 6,000 m. The daily

temperature in the reserve averages between 2 to 8" C (absolute minima reaching 18"

C), and dramatic diel temperature fluctuations are characteristic (Caviedes and Knapp,

1995; Pulgar, 1996).

Six Holdridge life zones are represented in the reserve. These include: 1)

subtropical desert mountains, 2) subtropical sub alpine mattoral desert, 3) subtropical

sub alpine humid paramo, 4) subtropical alpine humid tundra, 5) subtropical alpine very

humid tundra, and 6) subtropical nival. Also characteristic of this region is the presence

of snow at the highest elevations (typically the volcanoes) throughout the year. This is an

important source of water for both the vegetation and wildlife. Total annual precipitation

in this area fluctuates between 200 mm and 1,000 mm indicating a high degree of

interannual variability in water availability. The April to July period of this study

covered the end of the wet season, an extremely dry period, and an unexpected wet

period associated with a July snowfall.

Plants found in the study site display adaptations to dry puna, such as slow growth

rates, small plant size, a bunched distribution, and a shallow root system. The common










plants in the reserve include bunched grasses such as the common "ichu" grass (Stlpa

Ichu), several kinds of cactus such as "ulluyma" (OpuntlaJloccosa), the "huajoro"

(Opuntla lagopusl, and the "pajuro" (Opuntlna ignescens). This region also supports

several species of scrub brush. These include the arbustos enanos such as the "shauli",

and the arbustos de Culli, (Colli, quei~ua, and quinal) (Polylepls Racemosa) a tree-like

plant that is heavily exploited for wood.

The puma (Puma concolorl, vicui~a (Vlcugna vlcugna), and guanaco (Lama

guanlcoe) are the only large non-domesticated mammals in Salinas y Aguada Blanca

National Reserve. However, no signs of puma were found in the focal study area. The

only mammalian predator assumed to be in the reserve is the Andean fox jDuslcyon

culpaeusl. In addition to the large wild mammals found in this region of the reserve,

domestic animals inhabit regions near as well as in the reserve. Alpaca, sheep, goats, and

horses are all competitors of the wild species.

The domestic livestock are kept spatially separated from the vicui~a on the

mojedales (wetlands). In other areas of the vicui~a's native range, these wetland areas are

considered important habitat for wildlife, including the vicui~a(Franklin, 1983;

Renaudeau d'Arc et. al., 2000; Rundel and Palma, 2000). However, in this reserve, only

the domestic animals have access to these water and vegetative resources. The major

water source for the vicui~a in the reserve was the laguna. No other water source was

found in the reserve that was used by vicui~a (Davies, pers. obs.).

Due to its location within the reserve, land use by the community of Tambo

Cai~ahuas also has a significant impact on wildlife populations. An important feature in

this landscape is the road that runs through both the community and the reserve (Figure









2-1) Tlus is a malor thoroughfare between the cities of Areqinpa (the second largest city

mn the country) and Juhlaca, a city with sigmuficant commerce There are also several

tounst attractions and active nunes m tlus area that attract many large trucks and

additional traffic As it crosses through the resenre adjacent to the malor water source,

tlus road potentially contnbutes pomnt-source polhition and road-loll mortahties that could

have long lastmng, detnmental impacts on the wlldhfe populations





iM i---- Sumbay
Study site




Tambo
-- Cafiahuas
Laguna


S / -- Transect

Chachanl Volcano -- Paved Highway

Misti Volcano

a 0 aa ilometers




Figure2-1 Transect design for vacufia (Vlcugna vicugna) sunrey at the Salmas y Aguada
Blanca National Reserve, Peru, durmg Apnl-July 2002 Patterns of land use
are mdicated as well as the mamn roads that pass through the reserve

Current sustainable management strategies m Tambo Cafialulas mvolve the hive

capture, shear and release (called the Chaccu) of fr-ee-rangmng vcufias approximately

once every 2 yr CONATURA, an Areqinpa-based non-governmental organization

assists tlus communuty with logistical support, traimung, and organization Dunng the










time when these vicui~as are captured and sheared, biologists from CONATURA tag

individuals with cow tags and take measurements on age, sex, condition, and length and

diameter of the fiber. A total of 71 vicui~as were tagged between 1997 and 2000. In

addition, this NGO conducts scientific field studies and has established a monitoring

program to help provide data for management decisions.
















CHAPTER 3
METHODS

Study Design

Repeat samples of a 38-km transect and group follows were used to estimate four

major ecologicaltraits of vicui~a groups. These included 1) group and population

composition, 2) density and distribution, 3) the response to changes in water distribution,

and 4) spatial patterns and intergroup movement. Analyses of these four ecological traits

were used to quantify spatial and temporal patterns of habitat use, dispersal, relationship

to water resources, and density variation.In addition, a 3-yr CONATURA database was

used to reference individual histories on the vicui~as observed in the field survey. The

database comes from a long-term monitoring project and is comprsied of 15 surveys

between 1999 and 2001. Data collected by CONATURA include location, group size

and composition, and identification of some tagged individuals.

One large 38-km transect was established that covered a representative area of the

reserve including the major water source (the laguna), a large pampa or intermontane

plain, and regions near the highway. This transect was surveyed 13 times over the 4-

month study period. For each group or individual vicui~a sighted, the following

information was recorded: group type (i.e., bachelor herds, family groups, non-defined

groups and solitary males), group composition (# males, # females, # young), the identity

and gender of marked individuals, group activity, time of day, and GPS location recorded

in UTM coordinates with a Garmin 12XL GPS unit (KS, USA). Analyses of movement

patterns, distribution, density, and dispersal were based on obtaining exact positions of










vicui~a family groups, bachelor herds, and solitary individuals. This was done using a

variation of the distance method (Buckland et al., 1993). A Garmin 12x1 GPS (Global

Positioning Unit) in UTM was used from the transect to obtain coordinates of the vicui~a

groups. The center of the group was taken as the location of the group sighted, and the

bearing and distance were also recorded from this point. These data were then

transformed using the equation (d)*Cos(B)= a x and (d)*Sin (8) = ay. a x was then used

with the equation: ax = xl + xz~ and ay = yl + yz, and the new UTM coordinate was

found for the exact position. Specific information on marked individuals and behavioral

information were obtained with a Bushnell Spacemaster spotting scope. To determine

family group composition and to identify marked individuals, I also recorded tag

numbers of marked individuals (sexes can be differentiated because males have tags on

left, while females have it on the right). Differentiation of yearlings and adult vicui~a is

difficult, and this information was not collected. From the transect study, distinct group

types, [family groups, bachelor herds, solitary males, non-defined groups (groups of

unknown composition)] were identified and followed to assess movement patterns among

different social associations.

The study was conducted in two periods. The first sampling period ran from April

15 May 23. During this period the laguna was full, and the data collection focused on

the group census. The second period ran from June 24 July 12, during which time the

laguna and the surrounding plains became increasingly dry until a snowfall on July 10.

The size of the laguna was mapped immediately after the rainy season when the laguna

was full (April 25), and after a long dry period (June 28). The laguna was approximately

50 times larger in April than in June. In the first sampling period, transects were used to










identify groups and document their composition. In the second period, I used the same

transect design, but also followed known family groups to provide data on movement and

site fidelity.

Data Analyses

Group Characteristics

Group sizes observed during the wet and dry sampling periods were compared with

a t-test and then combined to produce one transect data set. From the combined transect

data, group size was averaged to obtain mean group size for all group types. To

determine the sex ratio within the vicui~a population at the Salinas y Aguada Blanca

National Reserve, averages of the number of males and females were calculated from the

combined transect data. A one-way analysis of variance (ANOVA) was used to test for a

difference in mean group size among bachelor herds, family groups, and non-defined

groups. The Scheffe apostenon test was used to detect differences in mean group size

between any two group categories. Group size values were log transformed to normalize

the data and stabilize the variance. Regressions were used to test for relationships among

group composition characteristics such as group size, number offemales, and number of

young.

Distribution and Density

Density

Vicui~a density was calculated by counting the number of vicui~a within a sighting

distance of 500 m from the transect, taking into account sampling effort, and then

dividing by the effective area sampled. Effective strip width (500 m) was defined by

measuring the perpendicular distance from the transect to each vicui~a group observation.

A sighting distance was determined from a frequency histogram of the distance a vicui~a










group was observed (Chapman et al., 2000). The frequency of vicui~a sightings were

consistently higher at distances less then 500 m. All vicui~a sightings greater then 500 m

were taken out of the overall density estimate. This sighting distance of 500 m

corresponds with figures used in previous studies that estimates vicui~a density (C.

Sahley, unpublished data). The total numbers of vicui~as in groups selected using this

criterion were averaged to include in the density estimate. To determine the area

effectively sampled, the sighting distance was multiplied by the transect length. Portions

where the transect overlapped were taken out of the measurement. The density of

vicui~as was then calculated as the number of individual vicui~as sighted divided by the

effective area sampled. Group density was simply calculated with the aid of the map

generated with Are View, and calculated as the number of groups within certain distances

~om the laguna, corrected for sampling effort.

Density and distribution relative to the laguna

Are View was used to create a series of maps describing the geographic locations

of vicui~a groups in relation to the laguna in the transect survey. A series of 20 concentric

rings (1 km apart) around the laguna's center were superimposed on the map of the

transect data. These maps were used to calculate the distribution of group size, group

composition, and group type in relation to distance from the laguna. Group density was

calculated as the number of groups within a certain distance from the laguna corrected for

sampling effort. Linear regression was used to detect the relationship between density

(dependent variable) and distance from the laguna (independent variable).

Group size distribution relative to the laguna

Group size was extracted from the transect data and mapped with the aid of Are

View to find patterns of distribution around the laguna. Vicui~a groups were arbitrarily










categorized into three size classes, small groups (1-5), medium groups (6-15), and large

groups (16-40). A chi-square test of independence was used to detect differences

between group sizes close (<4 km) and far (>4 km) from the laguna.

The number of males, females and the number of young in groups were quantified

in relation to their distance from the laguna. Regression analysis was used to detect

relationships between the number of males, females, and young (dependent variable) to

distance from the center of the laguna (independent variable). Are View was also used to

map this data for a visual display of distribution in relation to the laguna.

Responses to Changes in Water Distribution

The study was conducted in two periods (April 15-May 23 and June 24-July 12),

therefore, a t-test was used to detect differences in group size between the wet and dry

sampling periods. The distances of vicui~a groups from the laguna were categorized into

monthly averages to reflect changes in distribution in relation to changing water

distribution. An Are View map of the trasect data was superimposed on a map of the

study area. The distances vicui~a groups were observed from the laguna was recorded for

each month, correcting for differences in sampling effort.

Temporal changes in density and distribution were also calculated by quantifying

variation between wet and dry periods. A series of Are View-generated maps of the

transect data with 1 km concentric circles around the center of the laguna was used to

quantify variation in density and distribution of vicui~a groups relative to the center of the

laguna in the two major sampling periods. A chi-square test for independence was used

to detect differences in the distribution of vicui~a groups and group type around the

laguna between the wet and dry periods.










Spatial Patterns and Inter-Group Movement

To estimate spatial characteristics among family groups, locations of identified

dominant males were taken from the transect data and the group follows as these groups

moved to the laguna to drink. These observations were then mapped with the aid of Are

View to show group movement patterns over time. In addition, the life histories of

tagged males that I observed were extracted from the CONATURA database to estimated

site fidelity and to detect patterns in movement and group characteristics. These

individually identifiable groups were followed based on opportunistic observations to

obtain several geographic positions for 2-3 hr during the morning (sunrise to noon) and in

the afternoon (noon to nightfall). These groups were found and followed within an

approximate 5-km radius from the laguna. Finally, tagged females observed in

indentified family groups were extracted from the CONATURA database to explore the

validity of group stability as assumed in the literature.
















CHAPTER 4
RESULTS

General Ecology

Group Size and Population Composition

Family group size (t=-1.295, P=0.197, mean wet=6.07, n=136, SE+/-0.24; mean

dry=6.6, n=79, SE+/-0.31) and the number of females (t=0.108, P=0.914, mean wet=3.7,

n=136, SE+/-0.16; mean dry=3.6, n=79), did not differ between the two major sampling

periods. However, there were more young seen in the first period (mean=0.98, n=191)

than the second period (mean=1.22, n=149, t-test, t=-2. 845, P=0.005). Since young only

comprised 22% of the population, and other characteristics did not differ between

periods, I combined sampling periods to provide an overall evaluation of group

characteristics (n=21Sgroups).

Family groups comprised 67% of the total groups sighted, and bachelor herds and

solitary males represented 12% (6% each) of the sightings. Non-defined groups

(indistinguishable groups ofvicui~a) comprised 21% of the observed group types (Figure

4- 1).

Sex ratio

The sex ratio for adult vicui~as was 1.5 females for every 1 male. For family

groups, this ratio increased to 3.7 females for every 1 male. This ratio is based on adults

in identifiable group types, without consideration of age class.















70.00%


60.00%


50.00%


40.00%


30.00%


20.00%


10.00%


0.00%

fanily groups bachelor solitary non defined
herds r~Elles groups



Figure 4-1. Percentage of group types in the vicui~a (Vlcugna vlcugna) population of the
Salinas y Aguada Blanca National Reserve, Peru. Family groups were the
groups most frequently found in the survey and were comprised of a single
male and a variable number of females and young. Non-defined groups were
comprised of unidentified individuals with no young. Bachelor herds were
groups made up of several identified males and no females or young. Solitary
males comprised of a single male. Total group sightings = 317.

Group Size and Composition by Group Type

Family group size averaged 5.6 (antilog mean, n=215) and ranged from 2 to 13.


Bachelor herd size was larger than that of family and non-defined groups (ANOVA F=


102.98, p<0.001, Scheffe p<0.001), averaging 15.9 (n=18) and ranging from 8 to 40.


Non-defined groups were classified as indistinguishable groupings ofvicui~a whose


group size averaged 2.9 (n=65) and ranged from 2 to 13.


There were an average of 3.7 females and 1.6 young per family group. Family


group composition thus averaged i. 3.7. 1.6 (male: female: young). Thirty-two percent










of all family groups had four or more females. For observed males, 39.2% were in

family groups; 57.4 % were in bachelor herds, and 3.5 % were solitary.

Both the number of young and the number of females were correlated with group

size(young: r = 0.84, p<0.001, n=215; females: r = 0.93, p<0.001, n=215). The number

of young was also positively correlated with the number of females (r = 0. 60, p<0.001,

n=215). However, this relationship was not as strong as the relationship between the

number of young and group size, suggesting a greater amount of variation in the number

of young per family group. In addition, for males there was only one male in every

family group despite the variation in family group size.

Spatial Relationships and Intergroup Movement

Density and Distribution

Density and distribution relative to the laguna

At the Salinas y Aguada Blanca National Reserve vicui~a density averaged 3.7

vicui~asl km2 over the period of the investigation. Vicui~a density increased closer to the

laguna (r = -0.50, p<0.009; Figure 4-2). Forty-eight percent of all vicui~a groups in a 20

km radius were found within 2 km from the center of the laguna.

For data combined across the 4-month study period, all group types were present at

the laguna. However, habitat use of group types differed between the area close to the

laguna (within a 4-km radius) and areas more distant (4 to 20 km). The proportion of

group types was not independent of distance (12=15.16, p<0.005). Family groups

dominated in the first 4 km from the laguna, comprising 71% of all group types. Non-

defined groups made up 20%, followed by bachelor herds (6%) and solitary males (3%,

Figure 4-3). Four to 20 km from the laguna, family groups made up 57% of all group

types (a 14% decline), while the percentage of non-defined groups increased to 28%.










There were no bachelor herds more then 4 km from the laguna, and solitary males

comprised 15% of the groups 4 to 20 km from the laguna.

Non-defined groups and solitary males were more evenly distributed throughout

the study area than bachelor herds and family groups (X2=9.07, p<0.005). Seventy

percent of all family groups and 90% of all bachelor herds were found within 4-km of the

laguna. Only 56% of all non-defined groups and 27% of solitary males were found in

this distance class. In comparison, 4 to 20 km from the laguna, only 10% of the bachelor

herds were observed, and only 30% of the family groups. While 44% of all non-defined

groups and 73% of all solitary males were observed in farther regions from the laguna.

When translated to group densities, patterns were similar.

Group size distribution relative to the laguna

Vicui~a groups were arbitrarily divided into three size classes: small groups (1-5

individuals), medium groups (6-15 individuals), and large groups (16-40 individuals).

Small groups accounted for 54% of the whole vicui~a study population, while medium

and large groups comprised 44% and 2% of the groups, respectively.

To evaluate group size relative to the laguna, I quantified the frequency of group

size types as a function of distance from the laguna using l-km increments (Figure 4-4).

There was a tendency for larger groups to be more frequent closer to the laguna than in

the drier regions (XZ=51, O l
with 16 or more individuals (large groups which are exclusively bachelor herds) were

found within 4 km from the center of the laguna (n=5). For medium-sized groups (6-15

individuals, mostly family groups), 69% were found within 4 km of the laguna (n=94).

And, 59 % of small groups were found within 4 km from the center of the laguna (n=l 1 5)

















3 25

5~ 2

a 1.5
vl
a
1 1

0.5


1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20

Distance from laguna (km)







Figure 4-2. Vicui~a (Vlcugna vlcugna) group density as a function of increasing distance
~om a laguna in the Salinas y Aguada Blanca National Reserve, Peru. Forty-
eight percent of all vicui~a groups were found 2 km or less from the center of
the laguna.

































~~ ~~~~ -~

Figure 4-3 Dlstnbutlon ofvlcui~a (Vlcugna vlcugna) group types around the laguna at
the Salmas y Aguada Blanca National Reserve, Peru Data were collected
over 13 samplmg dates between Apnl 25 and July 12, 2002 by observmg
groups from a 38-km transect


Bachelor Herds

Family Groups


Non-defined Groups


Solitary Males










To evaluate group size relative to the laguna, I quantified the frequency of group size

types as a function of distance from the laguna using l-km increments (Figure 4-4).

There was a tendency for larger groups to be more frequent closer to the laguna than in

the drier regions (XZ=51, O l
with 16 or more individuals (large groups which are exclusively bachelor herds) were

found within 4 km from the center of the laguna (n=5). For medium-sized groups (6-15

individuals, mostly family groups), 69% were found within 4 km of the laguna (n=94).

And, 59 % of small groups were found within 4 km from the center of the laguna

(n=115).

Family group size as a function of distance from the laguna was also quantified.

Small family groups (1-5 individuals) made up 32% of all family groups, medium family

groups (6-10 individuals) made up 47% of all family groups, and large family groups

(11-15 individuals) represented 21% of all family groups. Larger groups were more

frequently observed closer to the laguna (< 4 km vs. >4 km; XZ = 16.37, p<0.005). All

large family groups were found closer than 4 km from the center of the laguna. For

medium sized groups, 65% were found within 4 km from the laguna, while 58% of small

family groups were found in this distance class.

Group composition relative to distance from laguna

The number of females and the number of young in groups were quantified in

relation to their distance from the laguna. The number of females decreased with

increasing distance













-- Sumbay






















-d

18 Kllometerr


--- Transect


Group Size


Tambo
Cai~aguas


Laguna


Figure 4-4 Size ofvlcui~a (Vlcugna vlcugna) groups m relation to the laguna Vlcui~a
groups were categorized into three size classes small(l-5), medium (6-15),
and large (16-40) Tnere was a tendency for large groups to be found closer to
the laguna than m the drier regions of the study site Group size data were
collected through the repeat census (number of group slghtmgs=317) of a 38-
km transect between Apnl 25 and July 12, 2002










from the laguna (r = -0.64, p<0.001), as did the number of young (r = -0.70, p<0.001,

Figures 4-5,4-6,4-7).

Only 19% of all family groups had more then the average number of young. Large

grouping of males (bachelor herds) were almost exclusively found near the laguna

(Figure 4-7), and there was a strong relationship between the size of bachelor herds and

water distribution (r = -0.60, p<0.001).

Responses to Changes in Water Distribution

Average distance from laguna as a function of changes in water distribution

The distances from the laguna that vicui~a groups were observed, were categorized

into monthly averages to reflect changes in distribution in relation to changing water

distribution. In April and May, the average distance of vicui~a from the laguna was 4072

m, (SD+/-1158.19) and 4127 m, (SD+/-4544.65) respectively. However, average

distance declined to 2568 m, (SD~-549.88) in June, which coincided with much lower

water levels in the laguna. In July, the average distance increased to 3996 m, (SD+/-

2119.72) (Figure 6a), possibly due to the unexpected snowfall in July, that increased the

extent of water around the plains and mountains.

This analysis included the farthest regions sampled, which may contain vicui~a

groups using water sources other then the laguna. To control for this potential bias, I

repeated the analysis removing groups more than 12 km from the laguna. The pattern

was similar, although averages were slightly lower (Figure 4-8).
















Final data set for a
0-3
4-10
Distance to Center
~1K
I c. 1 O 2K
[L7] 4K
O 8K
O 12K
16K
22K
I No Data





,, ",, ~

Figure 4-5. The number of females in family groups ofvicui~a (V~cugna mcugna) as a
function of distance from the laguna in the Salinas y Aguada Blanca National
Resenre, Peru. Family groups were divided into two categories: those with
less then the average number of females (0-3) and those with more then the
average (4-10). Data points represent obsenrations from repeat sampling of a
38-km transect on 13 occasions between April 25 and July 12, 2002.





Number ofyoung
0 o-2

3-10


Distance to Laguna

~1K
72K
~ 4K
78K
~ 12K
I 16K
122K
I No Data




~ ~


I
C
C
C
C
I
I
I


I


Figure 4-6 The number of young mcui~a (Vlcugna vlcugna) m relation to distance from
the laguna at Salmas y Aguada Blanca National Reserve, Peru Family groups
that had less than average number of young (0-2) and those that had more then
average (3-10) were compared Data pomts represent observations from
repeat samplmg of a 38-km transect on 13 occasions between April 25 and
July 12, 2002















Number of Males

0 0-i

2-40

Distance to Laguna
~ 1K
~-~r O 2K
O 4K
O 8K
~ 12K
I 1SK
122K
I No Data




.~~....., ~



Figure 4-7 The number of males in vlcui~a (Vlcugna vlcugna) groups m relation to
distance from the laguna at Salmas y Aguada Blanca National Reserve, Peru
Because family groups consist of only one male, most groups (67%) had only
one male Vlcui~a groups with more then one male were bachelor herds Data
points represent observations from repeat samplmg of a 38-km transect on 13
occasions between April 25 and July 12, 2002

















(a)
3100




2000
h
E """

Cd
E 100
3
t~a april M sy Juur July

,o ,,,,
Q) (b) ,,,,
O I I N
E 3soo -I 1 00
,,,,
.s
,,,,
zooo
isoo
iooo
soo


April M ay June July

M onth


Figure 4-8. (a) Average distance of vicui~a (Vlcugna vlcugna) groups from the laguna at Salinas y Aguada Blanca National Reserve,
Peru, and (b) average distance of groups to the laguna not including distances greater than 10 km. Average distances were
based on repeat sampling of a 38-km transect on 13 occasions between April and July, 2002.










Density and distribution as a function of changes in water distribution

A higher density of vicui~a groups was observed with delcining distances from the

laguna (Figure 4-9). When the laguna was full, 65% (n=82) of all vicui~a groups were

found within the first 4 km from the center of the laguna. In contrast, when the laguna

was dry, 83% (n= 102) of all vicui~a groups were found within this 4-km radius (Figure

4-10).

Family groups and bachelor herds seem to have been most affected by decreasing

water availability. The distribution within group types was quantified and divided into

those <4 km and >4 km from the laguna when full and when almost dry (Figure 4-11).

Seventy-five percent of all bachelor herds were found less then 4 km to the laguna when

full, and 100% when the laguna was almost dry. Sixty percent of all family groups were

present <4 km when the laguna was full and 75% were present when almost dry (Figure

4-11). These differences in group distribution were marginally significant (family

groups: XZ= 2.85, 0.1


For non-defined groups and solitary males, the proportion of groups <4 km and >4

km from the laguna was independent of season suggesting a more even distribution that

did not change with season (non-defined: XZ = 0.3, 0.9
0.5
when the laguna was full, and 56% and 33%, respectively were found in this distance

class when the laguna was almost dry (Figure 4-11).

Spatial Patterns and Inter-Group Movement

To examine spatial distribution patterns within the population, identified family

groups were followed (Table 4-1,4-2). There was a low degree of spatial overlap among

family groups










(a') Wet season*


(b) Dry season


OSoltary Male

gFarmly Group

Non-defined Groups

Bachelor Herds


:*"" Hilmee


Figure 4-9 (a') Dlstributon of vaicuia (Vicugna vicugna) groups wilth respect to the
laguna in the Sahnas y Aguada Blanca Nallonal Resenre, Peru (a) after the
wet season (Apn1 25 to May 23, 2002) when the laguna was full and (b)
dunng a dner penod (June 25 to July 12, 2002) when the laguna was reduced
to 2% of the wet season area















rn 5

~ Full
a

~h 3 1 ~ Almost dry

2
a
a



1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20


Area of ring



Figure 4-10. Density (groups per sq km) of vicui~a (Vlcugna vlcugna) groups around the
laguna at the Salinas y Aguada Blanca National Reserve when the laguna was
full and when almost dry.













































Figure 4-11. Percentage of individual vicui~a (Vlcugna vlcugna) group types found <4
km and 4-20 km from the laguna when it was full (April 25 to May 23, 2002)
and when almost dry (June 25 to July 12, 2002) at the Salinas y Aguada
Blanca National Reserve, Peru.


bh

fg
O ndf

Osol


1 2 3 4 5 6
m >4 km <4 km >4


Laguna almost dry


km


Laguna full


<4 k











Table 4-1. Composition of identified vicui~a (Vlcugna vlcugna) family groups in the
Salinas y Aguada Blanca National Reserve, Peru (April to July 2002).
Original identification and tagging were done by CONATURA.
Group composition Tagged individuals in family group
Males Females
1 male, 7 females, 6 young Blue #140 Orange #18
n=14 Orange/Al#16
Orange/Al #15
1 male, 6 females, 4 young Blue/Al #142 Orange/Al #20
n=ll Purple #103
Yellow tag
1 male, 5 females, 2 young Orange #6 Orange #9
n=8 Green#151
Green #156
Yellow tag
1 male, 3 female, 2 young Blue #142 Blue #144
n=6 Green #153
Orange #11
1 male, 1 female, Oyoung Orange #4
n=2


Table 4-2. Identified male groups of vicui~a (Vlcugna vlcugna) in the Salinas y Aguada
Blanca National Reserve, Peru (April to July 2002). Original identification
and tagging were done by CONATURA.
Group type Group size Identified tagged individuals
Solitary male n= 1 Blue #127
Solitary male n= 1 Blue #146
Bachelor herd variable White #52
White# 54
White #55
White #58
White #59
White #60
White #61
White #64










that were followed (Figure 4-12) and a high degree of site fidelity on behalf of the

identified male over the duration of the study. However, there was distributional overlap

with other non-identified groups.

The mean age of males from each group type was extracted from the CONATURA

database on the identified males observed during the 2002 field survey. Solitary males

averaged 11.5 yr; while young males (<3 yr) were found in bachelor herds, and males in

family groups averaged 9.5 yr. The latter males made up the effective breeding

population, as access to females is limited to those in a male's defended family group

(Koford, 1957; Vila and Cassini, 1994). The histories of the tagged males followed in

this study were also extracted from the CONATURA database to establish trends in age,

group type, and group size as well as site fidelity to the laguna (Table 4-3). Identified

vicui~as were repeatedly found at the laguna for up to 5 yr, indicating a high degree of site

fidelity to the laguna.

Within the social context of establishing corrals, data on the distances that vicui~as

move is critical. Distances moved by the followed groups were compared between the

morning and evening. In the morning, vicui~as moved an average distance of 713 m hr~'

(SE+/-162. 70), while they only moved 508 m hi' (SE+/- 45.67) in the afternoon.

Tagged females observed in identified family groups were extracted from the

CONATURA database to explore the validity of group stability as assumed in the

literature. Our study indicates that females vary in strategies. Some exhibited a high

degree of mobility among family groups, and others remained with the same male over a

long period. For example, tagged female orange #17 was observed to change family

groups four times within 2 yr (Table 4-4). She was observed with male blue/al #142,








switched to male blue #146, and then returned to male blue/al #142 In the same month,

she switched agam to blue #143 and to a family group with no


Laguna

~ _t


-~


Rgure 4-12 Spatial patterns of followed vlcui~a (Vlcugna vlcugna) groups around the
laguna at Salmas y Aguada Blanca National Reserve, Peru A low degree of
spatial overlap was observed between family groups in regions <4 km from
the laguna


Male # 146

Male # 2

Male # 143

Male # 6

OMale#140

Male # 142

Bachelor Herd


d











Table4-3. Life history characteristics of tagged male vicui~a (Vlcugna vlcugna) in the
Salinas y Aguada Blanca National Reserve, Peru in 2002. Characteristics
extracted from the CONATURA database.
Individual Year Longevity at # years Estimated Group Group type
tagged laguna observed age size
Blue 127 1997 5 2 12 1 Solitary
Blue 146 1997 5 4 11 1 Soitary
Blue 140 2000 2 2 13 14 Family group
Blue 142 2000 2 2 10 11 Family group
Blue 143 2000 2 2 9 8 Family group
Orange 4 1999 3 2 9 2 Family group
Orange 6 1999 3 4 8 6 Family group
White 52 2000 2 2 2 3 to 28 Bachelor herd
White 54 2000 2 2 2 3 to 28 Bachelor herd
White 55 2000 2 2 2 3 to 28 Bachelor herd
White 58 2000 2 2 2 3 to 28 Bachelor herd
White 59 2000 2 2 2 3 to 28 Bachelor herd
White 60 2000 2 2 2 3 to 28 Bachelor herd
White 61 2000 2 2 2 3 to 28 Bachelor herd
White 64 2000 2 2 2 3 to 28 Bachelor herd


Table4-4. Examples of movement within female vicui~a (Vlcugna vlcugna) among
family groups in Salinas y Aguada Blanca National Reserve, Peru from 1999
to 2002. Female Orange #9 remained with the same group over a 4-yr period.
Female Orange #17 switched among four groups. Dates tagged males and
females observed from 1999-2001 extracted from the CONATURA database.
Female Date Male Total group size
Orange #9 21-Dec-99 Orange #6 2
20-Apr-00 Orange #6 3
22-Jun-00 Orange #6 7
13-Dec-00 Orange #6 6
23-May-02 Orange #6 6
26-Jun-02 Orange #6 8
Orange #17 24-Nov-00 Blue/A1#142 2
13-Dec-00 Blue #146 2
30-Mar-O 1 Blue #146 2
1 i-Apr-02 Blue/A1#142 11
15-Apr-02 Blue #143 7
25-Apr-02 Blue #143 8
24-May-02 No Marks 4


other individuals marked the next month. In contrast, tagged female orange #9 was


consistently observed with the same male (orange #6) over 3 yr.
















CHAPTER 5
DISCUSSION

General Ecology

The vicui~a of Salinas y Aguada Blanca were dominated by family groups that

move daily and seasonally, but show minimal spatial overlap. Water seems to be a

fundamental environmental character influencing movement patterns and seasonal

changes in local density. These characteristics are very important when considering the

potential of current management initiatives. They also provide basic descriptors of the

ecology ofvicui~a and are useful in establishing interdemic patterns ofvariation.

The main objective of the study was to offer baseline ecological data on population

parameters of free-ranging vicui~a, and compare this data with other studies on vicui~a

and ungulates. The population characteristics presented in this study included: 1) group

size and group type composition within the population, 2) density and distribution, 3)

movement patterns in response to water distribution, and 4) spatial patterns and

intergroup movement.

Group Size and Type Composition

I used four categories to describe basic social units in the vicui~a of Salinas y

Aguada Blanca. These included: family groups, bachelor herds, solitary males, and non-

defined groups. These categories were created based on the different roles males play

within the population. Males in family groups are potentially the only males that are

breeding; thus family groups make up the effective breeding population. In Salinas y

Aguada Blanca, 67% of all vicui~a groups were in family groups, while only 12% were










male groups, indicating that approximately two-thirds of the males in this population

maintain a group of females and are breeding.

These trends in group type composition support an earlier study on the

socioecology of vicui~as. In his long-term study of vicui~as in the Pampa Galeras reserve,

Franklin (1983) described vicui~a population structure in relation to behavior and spatial

distribution. He recognized three different vicui~a group categories: family groups, male

groups, and solo males. In this study, Franklin found that 75% of all males were in

territorial groups while only 24% were in other male groups. Franklin also suggested that

male vicui~a in family groups are territorial and maintain boundary-oriented territories.

The limited overlap in range use documented in my study also suggest that over half of

the males in this vicui~a population may be territorial. These results on group type

composition in vicui~a populations coincide with the general population characteristics of

ungulates: within the social distribution of ungulates, generally 2/3 of the adult males

within a population are territorial (Owen-Smith, 1977).

Mating Strategies of the Vic~a

The life history strategies of ungulates evolved within the context of density

dependence and carrying capacity. As a result, strategies common to most ungulates

include the asymmetry of reproductive success between the sexes, a dimorphism in body

size, and a skewed sex ratio towards females (McCullough, 1999). A skewed sex ratio of

1.5 females: 1 male was found in the vicui~a population at the Salinas y Aguada Blanca

National Reserve. Similarly, the vicui~a in Pampa Galeras exhibited a similarly skewed

sex ratio of 1.2 females to 1 male (Franklin, 1983). However, in both these studies, it is

uncertain how this sex ratio changes with age. In many mammals, the sex ratio will be










skewed towards males at birth, then shift towards females in older age groups (Smith,

1996).

Earlier studies report resource-defense polygyny as the mating system of the vicui~a

(Bosch and Svendsen, 1987; Franklin, 1983; Vila, 1992). The structure of the family

group includes one adult dominant male, a number of females, and that year's young.

Our study detected no relationship between the number of males and group size,

providing evidence that the population in Salinas y Aguada Blanca also exist in

polygynous herds.

Group Composition Within Family Groups

Data collected during the field study suggest that vicui~a populations adopt a rigid

family group structure independent of environmental gradients and varying management

strategies. The mean group composition at the Salinas y Aguada Blanca National

Reserve (1 male, 3.7 females, and 1.6 young) falls within the range reported in earlier

studies. In Abrapampa Argentina, the mean group size was 1 male, 3-4 females, and 2-3

young (Vila, 1995), while a study done by the same author in 1992 in Catamarca,

Argentina reported a mean group composition to be 1 male, 3.6 females, and 1.9 young

Both of these sites have dry and harsh climates, similar to the Salinas y Aguada Blanca

National Reserve. In the Lauca National Park in Las Cuevas, Chile, the mean group

composition was 1 male, 3.1 females, and 1.6 young. In Pampa Galeras, a region with a

high density of vicui~a (67. 9 vicui~alkm2) and with more sources of water then my field

site, the ratio was 1 male, 3 females, and 2 young(Franklin, 1983). The mean group size

among vicui~as in corralled habitat in the humid puna where water is not such a limiting

resource was also 1 male, 3.4 females, and 1.7 young.










Group Size and Density

The average group size of family groups (antilog mean=5.6) in the Salinas y

Aguada Blanca National Reserve is similar to the averages reported from other studies.

In Pampa Galeras, the average group size was 6, (Franklin, 1983), and Vila and Roig

(1992) in the Laguna Blanca National Reserve, Argentina, found the average group size

to be 6 to7. Group size of bachelor herds ranged from 8 to 40 individuals in our study.

The dynamic nature of bachelor herd group size seems to be characteristic of vicui~a as

Franklin (1983) reported a range of 2 to 155 individuals and Vila and Roig, (1992)

reported a range of 2 to 22 individual bachelors.

There was a strong positive relationship between the number of females and the

number of young and total group size. However, the relationship between the number of

young in a group and the number of females is not as strong. This suggests that not every

female is giving birth every year. These females may be newly-dispersed young females,

or older, non-reproducing females. It could potentially be disadvantageous for the herd

as a whole to support too many young simultaneously due to the increasing energy

requirements of lactating females and the need to be alert more frequently (Vila and

Cassini, 1994). Other alternatives to explain this relationship include the mortality of

young and the inability of the female to conceive that year. No published data exist on

fertility among free-ranging vicui~a, however, my data suggest that population growth

rate and harvest models should not only incorporate a time lag, but also not assume one

young per female a year as a parameter (Cattan and Glade, 1989).

In general, there is a positive relationship between group size and an open habitat

structure (i.e., no cover) among ungulates. Large group size among ungulates is believed

to be a strategy to avoid predators and a way to optimize foraging behavior(Kie, 1999).










Increasing the number of females in a family group implies a requirement for increased

vigilance on behalf of the dominant male. However, in an analysis of vicui~a time

budgets, Bosch and Svendsen (1987) reported no difference in the amount of time a male

defends females between large and small family groups. In fact, the more females a male

has in his harem, the more individuals there are to watch for predators/aggressors while

the group feeds; thus increasing the amount of time each individual can spend foraging

instead of being alert. Therefore, under the spatio-optic conditions high Andean

mountain plains afford, the dominant male can increase the size of his family group by

adding females with little to no increase in energetic costs in terms of vigilance. As the

male vicui~a increases the number of individuals in his herd, the collective herd increases

their overall grazing efficiency. In addition, the number of females in a group is a good

predictor of the mating success among male vicui~a(Vila, 1995, Vila and Cassini, 1994).

Thus a large group size may be optimal under these arid and open conditions.

Not all vicui~a, however, occur in large groups. Over half (54%) of the vicui~a

population in the Salinas y Aguada Blanca National Reserve occurred in groups with 1 to

5 individuals. Franklin (1983) suggested that a section of the vicui~a population that

occurred more frequently in marginal habitats were temporary family groups with an

average of 4 individuals (1 male: 3 females: 1 young). These marginal grazing territories

were classified as feeding territories of hard bunch grasses with little to no water sources.

This variation in group size in the vicui~as of Salinas y Aguada Blanca National Reserve

may reflect spatial variation in habitat quality and the dependency on water sources for

large groups. Domestic livestock are given priority for the use of wetlands, leaving few

permanent water sources, such as the laguna, for the vicui~a population. This may










explain the tendency of large groups, especially family groups to be observed more

frequently than small groups closer to the laguna, suggesting differential habitat use.

Density estimates reported for vicui~a populations are highly variable among sites

ranging from 2.3 vicui~alkm2 at San Guillermo in Argentina, to 67.9 vicui~alkm2 at Pampa

Galeras, in Peru (Table 4). In our field study at Salinas y Aguada Blanca, the vicui~a

population occurred at a density of 3.7 individuals/ km2, falling within the range reported

for free-ranging vicui~a. The source of variation among vicui~a population estimates may

reflect density increases in corralled populations, temporal variation due to poaching and

other anthropogenic factors, natural decline, or differences in sampling techniques among

sites.

Other managed and non-managed ungulates also exist at very different densities.

Thompson's gazelle (Gazella thomsonl), another species adapted to harsh environments

exists locally at the Ngorongoro crater in Tanzania at 70 individuals/ km2, but more

generally at 14 individuals/ km2(McCullough, 1999). Ungulates inhabit a tremendous

diversity of ecosystems, and therefore face different kinds of environmental constraints.

However, they all demonstrate territorial behavior and exist in polygynous herds for at

least part of the year. Although no direct comparisons can be made to this study, these

data in conjunction with published estimates for polygynous ungulates suggest that there

is a high degree of both interspecific and intraspecific variation in the density of these

mammals.

In light of current management initiatives in Peru, it is also important to consider

the landscape and long-term population effects of maintaining vicui~a at artificially high

densities. Ecosystem management initiatives have only recently explored some of the










positive and negative feedback systems between herbivorous ungulates and vegetation

(Weisberg et al., 2002). Organisms at very high densities have to deal with limited

resources and decreases in overall population health. It is notable that densities in

corralled vicui~a populations are approximately

Table 5-1. Published denisty estimates ofvicui~a (Vlcugna vlcugna) along its range in
South America.
Author Place Year Density
-z
Korford Peruvlan Andes 1957 25 vlcuna km Corraled vlcuna

Brack Pampas Galeras 1980 67.9 vicui~a km~2
-z
Cajal San Guillermo 1991 2.3 vlcuna km Free-ranging vlcuna
(Argentina)
Villa Catamarca 1992 5.5 vicui~a km~2
(Argentina)
Davies Arequipa 2002 3.7 vicui~a km~2


10 to 27 times greater then estimates from free-ranging populations (Table 5-1). The

long-term effects of such intensive use of puna habitat by vicui~a remains largely

unknown.


McCullough (1999) states that vicui~a are long-lived, large bodied animals with

relatively low reproductive rates and a high level of female investment in raising young.

He also proposes that most ungulate populations, due to density-dependent responses,

have a tendency to exist at densities close to carrying capacity. Populations close to

carrying capacity are prone to explosions in growth rate and subsequent over-

consumption of resources that may induce a possible population crash (Caughley and

Sinclair, 1994; Smith, 1996).










Spatial Relationships and Intergroup Movement

Distribution Patterns

Understanding mechanisms underlying spatial and temporal patterns of distribution

is important in making both effective conservation and management decisions. Although

the Andean puna landscape appears to be barren and homogenous, different types of

habitat can be identified, and differential use in this habitat has been observed among

vicui~a (Renaudeau d'Arc et al., 2000). The optimality of habitat use is based on the need

to balance the intake of high quality food, growth, reproduction, and the need to avoid

predation, disease, etc.

In the vicui~a population of Salinas y Aguada Blanca, the laguna is an integral

feature in the landscape and seems to have a major influence on movement and spatial

patterns. Areas closer to the laguna may be of higher habitat quality based on water

availability for both vicui~a and vegetation, and on the increased density of vicui~a near

this water source.

Large groups were more frequently located near the laguna. Both the number of

females and young increased as distance to the laguna decreased. However, at Salinas y

Aguada Blanca, large bachelor herds (mean size >19) were also found closer to the

laguna, further supporting the important role of the laguna for this population. Other

studies have reported bachelor herds to move greater distances, as they forage in marginal

lands and avoid lands that territorial males occupy (Franklin, 1983).

This relationship to water in vicui~a re-asserts what is known about the relationship

between ungulates and xeric habitats. In dry environments, water is a major factor

limiting movement patterns and distribution. In our study we assume that habitat quality

increases as distance to laguna decreases. Working from this assumption, it could be










argued that medium and large groupings of vicui~a (n > 6) tend to be found more

frequently in areas close to the laguna, and possibly maintain a low degree of spatial

overlap around the laguna because of the high habitat quality. Although there is more

competition and possibly opportunity for predation, conflict, and invasion, there are also

more vigilant individuals in habitat of higher quality. In addition, less time and energy is

spent traveling to a water source if a territory is maintained nearby. Family groups were

the dominant group type in regions near the laguna; and all large family groups (n > 6)

were found less than 3 km from the center of the laguna.

The analysis of the socioecology of the vicui~a in Pampa Galeras conducted by

Franklin (1983) provides a useful comparison. The habitats are relatively comparable as

both have similar annual precipitation (Pampas Galeras: 281 to 742 mm; Salinas y

Aguada Blanca: 200 to 1,000 mm) and vegetation structure. In his study, Franklin found

that the permanent territorial family group consisted of an average of 6 individuals with a

mean group composition of 1 male, 3 females, and 2 young. These were groups that had

well-established high-quality territories. High quality territories were those that

contained water sources and vegetation associated with puna swampland such as quinsa,

chiula, and mojadales. Smaller family groups with 3-4 individuals were often found in

territories with no source of free water. This poor quality habitat was characterized by

the overbearing presence of roquei~o, piedras de loma and the ichu grass, vegetation that

neither requires nor contains much water.

Within the context of corral installation, the mechanisms behind differential habitat

use among vicui~a groups are important to explore. Increasing density either through the

use of management and husbandry techniques or by eliminating dispersal, despite










population growth, could put greater pressure on these vicui~a groups and the puna

ecosystem. Large groups could potentially be forced to reduce their feeding territories

due to over crowding, especially directly around the laguna. This could in turn, reduce

mean group size. Reducing mean group size may result in increased pressure on the

remaining individuals in the family group to spend more time being vigilant and less time

feeding. This process of overall group size reduction may already be happening in the

Salinas y Aguada Blanca National Reserve as 54% of all vicui~a groups have only 1-5

individuals. These animals are already relegated to habitat patches that appear to be of

poor quality and little to no water. The wetland areas (Bofedales, Mojadales) are

dedicated to supporting the hundreds of alpaca and sheep from the community. Not only

are the vicui~a prohibited access to these water and vegetative sources by the presence of

livestock, but are also deterred by the paved road separating the regions where the vicui~a

are and the wetland areas.

Seasonal Changes in Distribution and Density

The vicui~a must drink water regularly, even daily, to sustain their biological

processes. However, these ungulates have evolved within this context of severe water

limitation, and have adopted certain behaviors to cope with water restriction. Many of

these adaptations may be behaviors that are expressed in movement patterns such as

spatial distribution and group composition in relationship to water. For example, a

consistent pattern of movement from the edges of the mountains in the morning to water

sources in the afternoon has been observed in various habitats across the vicui~a's range

(Vila, 1992; Franklin, 1983; Vila and Cassini, 1993). To further explore this relationship

to water, the size of the laguna was used as a dryness index. Although these two

sampling periods may not represent two different seasons at their extremes (extreme wet










vs. extreme dry), there was a definate decrease in the water available. The perimeter of

the laguna was measured in April and decreased to 2% of its size when it was measured

again at the end of June.

In the Salinas y Aguada Blanca National Reserve, we detected a marked decrease

in the average distance of vicui~a from the center of the laguna between the end of the

rainy season (May) and the onset of the dry season (June). The vicui~a were

approximately 1.6 km closer to the laguna when it is almost dry. However, the transect

included territories that were > 10 km from the laguna. Due to potential habitat

~agmentation, the vicui~a groups on these territories could potentially be using a different

water source. After these observations (distance > 10 km) were taken out of the analysis,

vicui~a groups were found to be approximately 1.8 km closer to the laguna when it was

dry then when it was full. The distance of vicui~a groups from the laguna is almost 2 km

less at the onset of the dry season then at the end of the wet season. This marked

decrease coincided temporally with a decrease in precipitation in the whole region, as

expressed by a decrease in water distribution and water levels at the laguna. The

decrease in the average distance of groups from the laguna was reflected in an increase in

vicui~a density near the laguna in the dry season. A complementary relationship between

vicui~as and water distribution in the wet and dry season was also observed in a similar

habitat, a high, dry Andean grassland of Argentina by Vila and Cassini, (1993). Their

probability of finding individual vicui~as near water remained constant throughout the dry

season, but decreased in the afternoon during the wet season.

In our study, the average distance of vicui~a groups from the center of the laguna

decreased by 729.53 m in July. This was most likely due to the snow fall that occurred










throughout the month of July. Although the snow would melt by the afternoon, it served

to distribute water along the entire pampa and surrounding foothills. Vicui~a cannot

depend solely on the water from snowfall; however, the increased availability of water

gives both the vegetation a boost, and helps vicui~a with their intake of water as it collects

on the plants they graze on. The results of this study suggest that movement behaviors in

response to changing water distribution is a key element for survival in vicui~a.

Larger family groups and bachelor herds occupy the habitat immediately around

the laguna more frequently then smaller groups. I also detected differential habitat use

among vicui~a group types between the wet and dry seasons. How different group types

of vicui~a use the patches of habitat directly around the laguna offers some insight to

adaptive behaviors and population dynamics in free-ranging vicui~a. For example, in

general, over half of the family groups were found <2 km from the center of the laguna,

and 100% of the bachelor herds were found in this distance class as well. However, non-

defined groups (antilog mean -2.9), were observed to be more evenly distributed along

the study area. This suggests that these groups could be occupying dry, marginal habitat.

This decrease in representation of non-defined groups (both during the dry and wet

season) in habitat directly around the laguna further supports Franklin's notion that these

are small temporary family groups occupying marginal areas.

When the laguna was full, family groups were observed to dominate within the first

l-km from the center of the laguna. This was followed by the bachelor herds, the solitary

males, and lastly, the non-defined groups. When the laguna was almost dry, there was an

increase in the number of bachelor herds within this first l-km radius. Although it has

been proposed that bachelor herds move greater distances, there was a definite trend in










movement towards the laguna. Most bachelor herds in this study were considered large

(17.5 individuals) and were observed a distance of 3 km or less from the center of the

laguna. This suggests that group size may limit how far vicui~a groups can travel and

forage from the laguna.

Franklin (1983) also reported seasonal changes in vicui~a density. However, he

attributed these fluctuations to expulsion of young during certain seasons. In Pampa

Galeras, the young from family groups are expelled between October and March,

decreasing family group density. From April to May, there is an observed increase in

density, corresponding to the birthing season. From June to August in Pampa Galeras,

there is another unexplained decrease in density. Although our field study only covers a

fraction of the time data was collected on vicui~a density in Pampa Galeras, it could offer

a potential explanation to the observed decrease in density as vicui~a groups are observed

to limit their movements around water holes.

Spatial Patterns and Inter-Group Movement

Identified family groups were followed to gain insight into movement patterns at

the group and individual level. Earlier studies e.g., (Franklin, 1983) estimated home

range and relative territory size, however, these studies did not follow individually

marked individuals over long periods of time. Our study showed that vicui~a groups

exhibit a low degree of spatial overlap around the laguna.

The first formal study on vicui~as conducted by Koford (1957) in the Peruvian

Andes suggested that male vicui~a were highly territorial and would frequently engage in

tights and chases with intruding males. Territoriality is a common characteristic among

ungulates and leads ultimately to differential mating and reproductive success among

members within a population (Owen-Smith, 1977). In Pampa Galeras, Franklin (1983)










also proposed territoriality as a characteristic of the vicui~a; he observed males from

family herds frequently participating in antagonistic interactions with males from other

groups. He classified two different habitat types as vicui~a territories. One was a

sleeping territory on flattened ridges, and the other was a feeding territory on the lower

slopes and flatlands with higher vegetative biomass. He found this territoriality to be

based on its regular defense and on the maintenance of boundaries on behalf of the

dominant male in the family group. In addition, Franklin found that resident neighbors

respected these boundaries, while outside vicui~a groups did not. The nature of this

territoriality, found both from Franklin's study and from the data collected at the Salinas

y Aguada Blanca National Reserve, appears to be "boundary- oriented" (Owen-Smith,

1977). In our study, the low degree of spatial overlap between family groups, and to a

lesser degree, between solitary males and bachelor herds, suggests that vicui~a groups are

maintaining their daytime feeding territories at the laguna.

Mean age of identified males found at laguna in 2002

There are no published data describing age structure within vicui~a populations.

The age a male establishes residency on a territory and the length of time that male

retains a territory remains largely unknown (Bosch and Svendsen, 1987). At the time of

tagging during the chaccu, CONATURA estimates an individual's age by the wear on

their teeth. Therefore, I was able to compile preliminary data on relationships among age

classes, population structure, and movement. Our results indicate a specific age-class

structure within the vicui~a population, with the bachelor herds being the youngest, males

in family groups in a middle range, and solitary males in the oldest age class. This

tinding is a departure from the observations at Pampa Galeras. Franklin (1983) suggested

that solitary males are sexually mature attempting to establish territories. However, age










estimates found in this population suggest that solitary males may be in a stage of

senescence, not in dispersal and establishment. In addition, solitary males were also

found at farther distances from the laguna and in more marginal habitat patches than

other group types when water was scarce.

Site fidelity and dispersal

Not all tagged individuals were observed at the laguna during the course of the

study in 2002. This may reflect observation bias, death or, dispersal. We recorded only

one case of dispersal during our study. A solitary male, (Blue 127; age=12) was

observed approximately 17 km from the laguna where it was originally tagged. Although

one example of dispersal is not enough to make any conclusions about movements in

~ee-ranging vicui~a, it does suggest that the older, solitary males may have larger home

ranges or dispersal distances than other males found in family groups or bachelor herds.

The clustering behavior of young males around the laguna appears to be counter-

intuitive. Earlier studies (Koford, 1957; Franklin, 1983) suggest that these young males

disperse long distances to establish their own territories and family groups, thereby

encouraging genetic shuffling. The distance traveled by bachelor groups and their spatial

distribution suggest that they are potentially optimizing their opportunities to obtain

females at higher densities found around the laguna, while simultaneously being limited

in movement due to the scarcity of water.

Our data suggest that median-aged males maintain territories for several years.

Although not every tagged male was observed during the study, there is evidence that

these males have been coming to the laguna consistently for several years and exhibit a

high degree of site tenacity.










Distance moved from family groups

How much space a vicui~a group needs to fulfill its daily requirements is a central

question within the context of the installation of corrals. The size of corrals will vary

dramatically with habitat quality, density, water resources, and food supply. To begin to

answer this question, vicui~a family groups were followed, and their movements were

documented. The results suggest that vicui~a do not move very far, and that the corrals

would not limit their movement greatly. However, observations were taken only in

regions at or adjacent to the laguna, and did not represent movement across the vicui~as

entire home range.

In addition, differential movement patterns were observed as vicui~a groups moved

greater distances in the morning then in the afternoon. Vicui~a movement was high in the

morning while the groups were heading toward the water source as compared to the

afternoon when they move back to their sleeping areas. These results correspond well

with the study of Vila and Roig (1992) that focused on daily movements and behavior of

family groups. Their study found that vicui~as moved greater distances in the morning

towards the water source and grazed in the afternoon.

Description of shifts in group composition

In this study males tended to maintain spatially discrete ranges around the laguna,

that remained relatively constant for the 4 months of observation. I hypothesize that

males establish territories, and females will occasionally switch family groups. In this

way, there is the potential for genetic shuffling between resident vicui~as in this

population, while maintaining a stable spatial distribution over time. Other studies have

assumed that groups are basically stable, comprising of the same individuals over time

(Franklin, 1983; Vila, 1995). Females in my study were found to adopt different mating










strategies. Some were sighted over several years with the same identified male, and other

females switch between different males at different rates.

Female vicui~as exhibit iteroparous reproduction, giving live birth to one young at a

time, repeatedly during her lifespan. Young ruminants depend completely on their

mother's milk for the required amino acids, vitamins and the inoculation of rumen

bacteria to develop ruminant fermentation (van Soest, 1994). Thus, the female vicui~a

has different energy requirements and constraints in maximizing reproductive success

then males (Kie, 1999; Vila, 1995). The number of family groups a female belongs to in

her lifespan may not affect her fitness. What matters more for her reproductive success is

the habitat quality of the dominant male in her family group, i.e., the quality, quantity,

and heterogeneity of the vegetation for embryo development, lactation, and the continual

growth of her young (Robinson et al., 1999).
















CHAPTER 6
IMPLICATIONS FOR CONSERVATION

An increase in the numbers of vicui~a has been reported over the last 30 years in

Peru. There were reportedly only 10,000 vicui~as left in 1964 (CONACS, 1997); as a

result of intensive management practices, the vicui~a population in Peru reportedly

increased to 150,000 individuals by 2000 (Lichtenstein et al., 2002). Despite the increase

in numbers, the distribution of the vicui~a has been dramatically reduced mainly to the

three national reserves in Pampa Galeras, Huaraz, and Salinas y Aguada Blanca. Vicui~a

no longer exist along the entire historic range in Peru (Figure 6-1). It is unknown if the

increase in overall vicui~a numbers is due to strict protectionist measures, or to local

dynamics in a few communities that have dedicated themselves to promoting the growth

rate among vicui~a to facilitate "rational use" of vicui~a fiber as a sustainable management

strategy. Thus, increases in numbers may potentially only reflect a geographically

limited increase in population density.

This centralized increase in vicui~a numbers is exacerbated by new management

initiatives that involve the installation of corrals. This may potentially change the

distribution of vicui~as and alter important ecological characteristics of their population

ecology. In addition, it invariably segregates potentially inter-breeding vicui~a

populations, encouraging homozygosity. It is important to understand the mechanisms of

this population growth among vicui~a, because changes in social behavior associated with

changes in density may change patterns of habitat use, and may negatively alter the puna

ecosystem.















'~r (.

Study site II
.1'




"'D
"' ""




Figure 6-1. Historic range of the vicui~a (Vlcugna vlcugna) in South American Andes.
Figure adapted from The Camelid (W. Ross Cockrill, 1979).

Information on behavior patterns of free-ranging vicui~a is scarce. Much of the data

on their habitat requirements are from studies that focused solely on biomass, husbandry,

and management (Wilson, 1994). Much of the current data on vicui~a ecology comes

from highly managed vicui~as in corrals and does not directly answer any of the questions

surrounding the issue of limiting movement in regard to group composition, movement

patterns, spacing patterns, and territoriality.

Like other density-dependent ungulates, the vicui~as may exhibit population

oscillations. Therefore great precaution must be taken in determining corral placement,

size, and the number of vicui~as to maintain in those corrals. If the population falls below

a viable level, there could be detrimental consequences for the vicui~a population as a

result of decreased genetic heterogeneity. For example, populations with low genetic

diversity are at risk of lowered immunity to the spread of disease throughout the entire










population, and more vulnerable to stochastic abiotic factors (Caro, 1998). On the other

hand, if the population grows rapidly, the puna ecosystem could be at risk of over-

grazing, homogenous vegetative composition, soil erosion, and desertification.

Currently, the focus in conservation and sustainable development programs seems to be

in increasing the population of vicui~a. Much effort has gone into determining how many

vicui~a can subsist given forage production, given that their use of fibrous feeds is

extremely efficient because of their highly specialized mode of fore-gut digestion (Cajal,

1991; Cueto et al., 1985). However, there are no studies that estimate how long the puna

ecosystem can maintain a high density of vicui~a before nutrient levels in the soil decline,

vegetative structure is degraded, or hydrological regimes altered.

The ability of vicui~a to have access to water and forage of sufficient quality and

quantity is essential in maintaining a viable vicui~a population. The data suggest that

vicui~a in the Salinas y Aguada Blanca National Reserve respond behaviorally to harsh

conditions by moving in response to resource availibility. Limiting vicui~a movement

through the implementation of corrals may have negative impacts for both the health of

the vicui~as and their habitats. The nutritional quality and quantity of biomass in puna

ecosystems has been reported to decline from the end of the wet season through the dry

winter (Pfister et al., 1989). This impacts vicui~a and their movement significantly as the

typical functional response in ruminants is to increase the time spent foraging to

compensate for the lowered nutritional value.

My results show decreased movement centered around the laguna during the dry

season. This limitation in movement around the laguna may be a result of an increased

water requirement. Or, there may be more vegetation of higher quality around the water










source available, decreasing the energy needed to satisfy water and nutritional

requirements. There are other constraints operating in conjunction with the decrease of

water availability and changes in nutritional quantity and quality. As temperatures

decrease during the dry winter, there is a higher cost of thermoregulation (Vila and

Cassini, 1993). Movement in search of forage among vicui~a also results in heat

production and lowered energy retention, especially when both the quantity and quality

offorage is reduced(Murray, 1991).

Limiting movement through the installation of corrals during the dry season may

not have a significant impact on the social structure of the vicui~a, or on natural

movement patterns. However, overgrazing in these areas near the laguna could have

significant long-term consequences on the quality of forage for subsequent years. In

addition, the dry season is also marked by frequent snowfall, as observed in our study

from July 10 to July 12. At this time, vicui~a groups were found father from the water

source, and most identified family groups were not found at the laguna at all. The

increased distribution of water possibly allowed for greater movement, subsequently

releasing local vegetation around the laguna from foraging pressure. The installation of

corrals would obviously limit a functional response among vicui~a to availability to

resources.

Another potential consequence of corral installation in the Salinas y Aguada Blanca

National Reserve is related to land use and the geographic position of certain key

landscape features. The area where vicui~a are found is bisected by the paved road

running from Arequipa to the Colca and Juliaca-Cuzco and presents a potential source of










vicui~a mortality. There were several instances during the study where vicui~a groups

were observed crossing this highway in search of water in the reserve.

Placing a corral in this reserve would exclude other vicui~a groups traveling in

search of water and forcing them to find other sources, either in regions near domestic

livestock, or in farther regions. This could potentially decrease the amount of vicui~as

available for shearing during the roundup.

The primary aim of the rational use of the vicui~a as a sustainable management

program is two-fold. First, it claims to protect the vicui~a, while secondarily providing a

legal, alternate source of income for impoverished Andean communities. A major threat

to the vicui~a is anthropogenic in nature as Andean community members claim that there

is competition for feed resources between naturally occurring vicui~a and their domestic

livestock (Urquieta et al., 1994). The implementation of corrals has been hailed as a

viable compromise that will alleviate this competition, provide protection for wild

vicui~as and facilitate the capture process for shearing. However, these corrals may serve

to alter the puna ecosystem and the social and foraging behavior of the vicui~a that may

lead to negative consequences.
















CHAPTER 7
CONCLUSIONS

Baseline data on the population characteristics of free-ranging vicui~a populations

are critical to the evaluation of effective conservation and management practices and to

our understanding of the importance of local habitat features to population density and

distribution patterns. Group characteristics of the vicui~a of the Salinas y Aguada Blanca

National Reserve are comparable to studies of other free-ranging vicui~a, with densities

notably lower than corralled populations. The changes in distribution and density over

the 4-month study suggest that the location of a permanent water source may be an

integral factor contributing to patterns of distribution and movement over the seasonal

cycle. The major conclusions drawn from this study are summarized below.

1. In the Salinas y Aguada Blanca National Reserve, the mean family group
composition is 1 male, 3.7 females, and 1.6 young. These figures fall within the
range of other studies that include both corralled and free-ranging populations,
suggesting that the vicui~a have a rigid family group structure within different
populations that occur across ecological gradients and management programs.

2. In my study average density was 3.7 vicui~aslkm2. This falls within the range of
studies of free-ranging vicui~a and is much lower than corral estimates.

3. Water distribution and availability seem to have a major effect on vicui~a
movement. Groups were found to be 2 km closer to the laguna in drier conditions.
This may have management implications, as water is a major limiting factor in the
puna. Although domestic livestock and vicui~a herds have not been found to
compete for forage, they could be competing for water resources.

4. Overall, large groups and large family groups are found closer to the laguna. This
indicates that large groups are limited in their movements by water availability, and
that large groups tend to establish territories in higher quality habitats near the
laguna.










5. The data on the Salinas y Aguada Blanca vicui~a in combination with other studies
on ungulates, suggest that large family groups in open habitats is an optimal
strategy for feeding and reproducing, as smaller vicui~a groups are more frequently
in marginal habitats.

6. There may be a rigid population structure and family group structure within vicui~a
populations found in different habitats. Several group characteristics such as group
type distribution in the population, group size, and group composition remained
stable despite changing water distribution.

7. There was a low degree of spatial overlap around laguna. This suggests that the
dominant males in family groups are maintaining boundaries in relation to other
family groups when traveling to the laguna. This also provides further evidence for
territoriality among males in family groups.

8. Individual vicui~as and groups were found to exhibit differential movement patterns
throughout the day. Vicui~as move greater distances in the morning, presumably to
reach the laguna by midday. The rate of movement decreases in the afternoon.
These differential movement patterns are important when estimating how far a
vicui~a will move, and subsequently how large their territory requirements are in
the context of corral initiatives.

9. Our study provides evidence that some females will stay with the same male in a
family group for an extended period oftime, while others will switch annually
among family groups.

10. With the use of age estimations provided by the CONATURA database, an age
class structure was found in the vicui~a in the Salinas y Aguada Blanca National
Reserve. Bachelor herds were found to be young males that are non-territorial
dispersers. Other studies report that solitary males are in process of establishing
their territories, however, this study indicates that solitary males are older and in a
stage of senescence. Although there was overlap, the mean age of solitary males
was higher then those in family groups


















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BIOGRAPHICAL SKETCH

Jennifer Elena Davies was born on May 4, 1975, in San Diego, California. She is

the only child of Thomas M. Davies Jr. and Eloisa Davies Monzhn. She earned her

degree of Bachelor of Arts in Nature and Culture, with a minor in Spanish from the

University of California, Davis, in 1999. There she translated her respect and love for the

natural world to academic study and research in human-animal interactions. To continue

her commitment to interdisciplinary research in the natural sciences she moved to

Gainesville to enroll in graduate studies in the Center for Latin American Studies/

Tropical Conservation and Development in the fall of 2001.