NUTRITION OF GRAZING HIGH ALTITUDE RANGELAND IN SOUTHERN PERU by RICHARD JAMES REINER, B.S., M.S. A DISSERTATION IN AGRICULTURE Submitted to the Graduate Faculty of Texas Tech University in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY Approved

May, 1985 //^.^'-^ . y > .' ACKNOWLEDGMENTS

Funding for this project was provided by the U.S. Agency for International Development Small Ruminant Collaborative Research Support Program. I would like to extend my sincere thanks to my major professor and "compadre," Dr. Fred C. Bryant, without whose personal support and financial magic, this dissertation would not have been possible. I am especially grateful to committee members. Dr. Carlton Britton, Dr. Billie E. Dahl, Dr. B. Frank Craddock, and Dr. Rodney L. Preston, for their guidance and careful reviews of this manuscript. The tedious laboratory work would never have been compleated without the help of Susan Rikert, Brigid Holland, Gretchen Scott and Khang Nguy. identification was confirmed by Dr. Oscar Tovar of the Universidad de San Marcos, Lima, Peru. Esta disertacio'n no habria sido posible sin la ayuda de los Doctores Luis Carlos Fierro, Julio Sumar, Felipe San Martin, Arturo Rosales y del personal de las Estaciones Experimentales de La Raya y Chuquibambilla. Mi especial agradecimiento a los Ingenieros Ramiro Farfan, Timoteo Huisa y Godofredo Atamari por dirigir el trabajo de campo y por la paciencia que me tuvieron, debido al poco conocimiento que

11 tengo del Idioma Espanol. Muchas gracias por tratarme como un amigo y por hacer placentera mi estancia en Peru. On the night of July 22, 1983 the La Raya research station was attacked by terrorists armed with machine guns and explosives. Susan Rikert, myself and the La Raya staff were held at gun-point for a terrifying night while the station and our home were destroyed. It took very special people to work under conditions like these. Only because of your courage was this project a success.

"This dissertation is dedicated to Champa."

Ill CONTENTS

ACKNOWLEDGMENTS i i ABSTRACT vi LIST OF TABLES ^^ii LIST OF FIGURES x

CHAPTER I. INTRODUCTION AND OBJECTIVES 1 II. LITERATURE REVIEW 5 Reproduction 6 Animal Management and the Andean Environment 7 Diets 9 Alpaca Digestive Morphology and Physiology 11 Digestive Efficiency and Forage Consumption 14 III. DIETARY SELECTION OF ALPACAS GRAZING ALTIPLANO AND BOFEDAL RANGELAND SITES 18 Description of the Study Areas 18 La Raya, High Mountain Site 18 Chuquibambilla, Altiplano Site 23 Methods 24 Study Design 24 Forage Availability 26 Diet Sampling and Analysis 27 Results 30 Forage Availability 30 Diet Botanical Composition 32 Discussion 38 IV. DIETARY NUTRIENT CONTENT OF ALPACAS GRAZING ALTIPLANO AND BOFEDAL RANGELAND SITES 4 3 Methods 4 3 Results 45 Discussion 48

1 V V. INTAKE OF ALPACAS HOUSED IN METABOLISM CAGES AND FREELY GRAZING A NATIVE ANDEAN PASTURE 52 Methods 52 Intake in Metabolism Cages 52 Intake of Free-ranging Alpacas 53 Results 55 Intake in Metabolism Cages 55 Intake of Free-ranging Alpacas 56 Discussion 58

VI. SUMMARY AND CONCLUSIONS 62

LITERATURE CITED 66

APPENDIX

A. PLANT SPECIES LIST 74

B. FORAGE AVAILABILITY AND DIET COMPOSITION DATA 7 7

C. CP AND IVOMD DATA 86

D. INTAKE STUDY DATA 88

E. STATISTICAL TABLES 91 ABSTRACT

Fiber and meat production from alpacas are important for the livelihood of both subsistance and market-oriented producers using high elevation Andean grasslands in Peru. The study objectives were: 1) to seasonally measure alpaca diet quality and botanical composition on two rangeland sites; and 2) to estimate forage intake of alpacas grazing Andean rangeland. Diets were collected from free-ranging, esophageal fistulated alpacas during eight months between June 1983 and March 1984. The "bofedal" site (4,500 m) was a perennially green sedge community dominated by Eleocharis albracteata and Carex ecuadorica. Vegetation cover was 22% grass, 42% sedges and reeds, and 33% forbs. The "Altiplano" site (3,190 m) was predominately the grass species Festuca dolichophylla and Muhlenbergia fastigiata. Vegetation cover was 66% grass, 13% sedges and reeds, and 6% forbs. Diets at both sites were highest in grass during the wet and early dry season. As the dry season progressed bofedal alpaca diets became largely sedges and reeds (81%) while Altiplano alpaca diets increased in grass (68%). Forb consumption varied between 9 and 26% of the diet year-round.

vi Crude protein (CP) in bofedal alpaca diets (12.3%) averaged higher than on the Altiplano (10.2%). Values were lowest during June (7.1%) on the Altiplano and July (8.0%) on the bofedal. Dietary CP at the bofedal site was higher during the nutritional critical late dry season. Ln vitro organic matter digestibility of alpaca diets was lowest during August (49%) at the Altiplano site and in October (50%) on the bofedal. Alpacas grazing the Altiplano may be deficient in CP during the entire dry season (Jun.-Nov.) and energy deficient during the late dry season (Aug.-Oct.). Dietary CP may be deficient on the bofedal site only during the early dry season and energy deficient during the late dry season. Supplementation of alpaca diets is recommended during periods of protein and energy deficiency.

Alpacas freely grazing a Festuca dolichophylla / Muhlenbergia fastigiata range consumed daily 50.5 and 44.3 g organic matter per kg metabolic body weight during the dry and wet seasons. These values are lower than reported for sheep. Recommended stocking rates for alpaca may require re-assessment.

VI 1 LIST OF TABLES

Page 1. Sample dates, conditions and locations associated with collection of alpaca diets at the La Raya (bofedal) and Chuquibambilla (Altiplano) sites during 1983 and 1984 26 2. Percent forage foliar cover during eight months of 1983 and 1984 on the study sites in southern Peru.. 30 3. Bofedal site dietary botanical composition listed in order of annual contribution and then re-ranked by season 35 4. Altiplano site dietary botanical composition listed in order of annual contribution and then re-ranked by season 36 5. Frequency of grass seeds and flower parts in the diets of alpacas grazing the study sites 38 6. Number of fistulated alpacas needed to estimate CP and IVOMD with 95% confidence within 10% of the mean during five day collection periods 48 7. Organic matter intake (OMI) and digestibility (OMD) of two forages consumed by eight alpacas in metabolism cages at La Raya, Peru 55 8. Vegetation biomass (kg DM/ha) of a-La Raya valley bottom pasture during the dry (October) and wet (February) seasons 56 9. Dietary crude protein, IVOMD and organic matter intake (OMI) of free-ranging alpacas sampled during the wet and dry seasons at La Raya, Peru 57 10. Plant species found on the bofedal and Altiplano sites during 1983 and 1984 75 11. Bofedal vegetation cover and alpaca diet botanical composition on June 10 and July 10, 1983 78 12. Bofedal vegetation cover and alpaca diet botanical composition on August 22 and October 4, 1983 79

VI 11 13. Bofedal vegetation cover and alpaca diet botanical composition on November 7 and January 1, 1984 80

14. Bofedal vegetation cover and alpaca diet botanical composition on February 4 and March 3, 1984 81 15. Altiplano vegetation cover and alpaca diet botanical composition on June 10 and July 10, 1983 82 16. Altiplano vegetation cover and alpaca diet botanical compostion on August 22 and October 4, 1983 83 17. Altiplano vegetation cover and alpaca diet botanical composition on November 7 and January 1, 1984 84 18. Altiplano vegetation cover and alpaca diet botanical composition on February 4 and March 3, 1984 85 19. Mean CP and IVOMD with coefficients of variation (CV) in alpaca diets ' 87 20. Organic matter intake and fecal output of caged and free-ranging alpacas 89 21. Vegetation biomass (kg/ha) in a La Raya valley bottom pasture grazed during a intake study of free-ranging alpacas 90 22. Analysis of variance for dietary grass composition 92 23. Analysis of variance for dietary sedge and reed composition 92 24. Analysis of variance for dietary forb composition 93 25. Analysis of variance for dietary crude protein 93 26. Analysis of variance for dietary IVOMD 94

IX LIST OF FIGURES

Page 1. Mean daily precipitation (mm) recorded at La Raya and Chuquibambilla from April, 1983 to April, 1984. No data were recorded during April and May at Chuquibambilla 21 2. Average daily maximum and minimum temperatures (C) recorded at La Raya and Chuquibambilla between April, 1983 and April, 1984 22 3. Percent dietary botanical composition for alpacas grazing the bofedal and Altiplano sites 33 4. Percent crude protein with 95% confidence limits in the diets of alpacas grazing the bofedal and Altiplano sites 46 5. Percent in vitro organic matter digestibility (IVOMD) with 95% confidence limits in the diets of alpacas grazing the bofedal and Altiplano sites 47 CHAPTER I INTRODUCTION AND OBJECTIVES

Alpacas (Lama pacos) and (Lama glama) are the only indigenous domestic ungulates of South America. Both animals are members of the family Camelidae, which consists of the genus Lama of South America and Camelus of Asia and North Africa. Alpacas are raised primarily for wool fiber, while llamas are most often beasts of burden. Alpacas are grazed primarily on high elevation grassland in the Mountains of southern Peru and northern Bolivia. The Peruvian government estimates that southern Peru's 6.5 million hectares of high elevation rangeland supports approximately 3 million camelids, 11 million sheep and 4 million cattle (Holgado et al. 1979). Highland Indian populations of Latin America are among the economically poorest inhabitants of the continent. Most communities are located above 3,100 m in harsh alpine environments characterized by seasonally low temperatures and intense solar radiation. At these elevations less than 5 % of the land is suitable for cultivation (Gilles 1980), the remaining portion is used only by range livestock and wildlife. In most highland Peruvian communities, the people's livelihood depends on wool fiber and meat production from alpacas and sheep. Sheep are grazed up to 4,200 m elevation. Above this elevation they suffer respiratory infections. Alpacas can graze to 6,000 m without problems. High elevation rangeland in southern Peru can be characterized as two expansive grassland zones: the high plains (Altiplano) of the Lake Titicaca basin (3100 m elevation), and the high mountains surrounding the basin. Rangeland in the high mountains can exceed 6,000 m. The Altiplano is primarily a zone of sheep production while alpacas are grazed in the high mountain zone. Precipitation-in southern Peru is bimodal. November through April constitutes the wet growing season. During the dry season (May - October) there is no precipitation and little forage production. August through October are months of forage scarcity. During this season alpacas rely on perennially green, tundra-like, pastures called bofedales. Bofedales are common in the high mountain zone and less abundant in the Altiplano basin. Range management in the form of grazing systems, diet supplementation, or trucking animals to seasonal pastures is rare. Over most of their range, alpacas suffer from disease, impaired fiber production and low fertility which commonly are attributed to inadequate nutrition. It is the consensus of most experts that adequate nutrition during the dry season is the key to improving alpaca production in Peru (Fernandez Baca 1975, West 1981). Unfortunately, there is little information available regarding the nutritional quality, diet botanical composition or intake values for the varied range sites alpacas graze. In recent years, the market for alpaca fiber has increased. In 1984 alpaca fiber sold for over six times the price of sheep wool. In response to this market, livestock producers in the Altiplano are increasing alpaca numbers. As alpaca numbers increase, scarce Altiplano bofedales will be further overgrazed. Nutritional supplementation or trucking animals to high mountain bofedales may become necessary. This dissertation addresses three topics in addition to a literature review of alpaca management and biology. First it describes seasonal dietary selection of alpacas grazing typical bofedal and Altiplano rangeland sites (Chapter III). Secondly it contrasts the nutritional quality of diets collected at each site (Chapter IV). Last, dietary intake for alpacas is estimated in metabolism cages and in a free-ranging situation (Chapter V). The specific objectives were: 1. To describe and contrast the seasonal botanical composition of alpaca diets in an Altiplano pasture and high mountain zone bofedal. 2. To describe and contrast seasonal crude protein and organic matter digestibility of alpaca diets in an Altiplano pasture and high mountain zone bofedal. 3. To compare the dietary organic matter intake of alpacas housed in metabolism cages with free-ranging alpacas in dry and wet season pastures in the high mountain zone. CHAPTER II LITERATURE REVIEW

The world population of alpacas is now approximately 3.2 million, with 70% living in southern Peru (Fernandez Baca 1971). Peasant villagers own over 80% of Peru's alpacas. They are grazed in small household herds and on community land (Orlove 1977). The remaining 20% of Peru's alpacas are raised on large social cooperatives. Peru produced over 3,400 metric tons of alpaca fiber in 1980. Almost 80% was exported to Europe (West 1981). There are two distinct breeds of alpaca': the huacaya and the suri (Reiner and Bryant 1983). The huacaya is characterized by highly crimped wool which appears much like that of Lincoln sheep. Suri wool is straight with very little crimp. Condorena (1980) found five year old huacaya and suri alpaca to be approximately equal in size (61.7 vs 62.5 kg). Wool fiber production (kg/ha) in huacayas also was slightly greater. There is little selective breeding favoring one breed over the other. Huacayas are more common however, especially in colder climates (Tillman 1981). Reproduction The unique reproductive characteristics of alpacas mandate management practices unlike those employed with cattle and sheep. Alpacas are polygamous, but unlike cattle and sheep they are copulation-induced ovulaters (as are rabbits and cats). Thus, females have no defined breeding estrous cycles. Ovulation occurs 24 to 36 hours after copulation (Fernandez Baca 1971). Although breeding seasons exist, a female can ovulate and give birth at any time of the year (Schmidt 1973). In southern Peru alpacas are bred during January and February (wet season). Because gestation is roughly a year (340 to 350 days), most births occur during a time when forage is green and plentiful. Breeding ratios vary but normally are between 5 to 10 males per 100 females (Escobar Calle 1982). Estrus disappears within five days following copulation. The female will then reject the approach of a male. If fertilization does not occur, follicles again become active and estrus can be observed within 13 days. The absence of estrus is a diagnostic sign of fertilization (San Martin et al. 1968). Alpacas have a bi-cornuate uterus, but 98% of all pregnancies are carried in the left uterine horn. There are no records of twins. Birth almost always occurs in the early morning, possibly an adaption to low nighttime temperatures of high altitudes (Tillman 1981).

Fertility in alpacas appears to be low when compared to other livestock. Observations by Novoa (1970) show that alpaca fertility is approximately 50% on most alpaca ranches. Embryonic mortality may be a major cause of low fertility (Sumar 1983). West (1981) suggests that poor herd management is a underlying cause of low fertility. On the well managed ranch "Estancia Accoyo," which he investigated, fertility was above 80%.

Animal Management and the Andean Environment Animal management is usually in response to the distinct seasonality of precipitation and the reproductive status of female alpacas (Escobar Calle 1982). Intensity of herd management varies greatly between producers. In small villages alpaca owners employ only traditional management. Animals are herded in response to availability of grass and community customs (Tapia Nunez and Flores Ochoa 1984). Little attention is given to protecting vegetation from overgrazing or separating herds by specific classes. In contrast, one finds better managed herds on larger cooperative ranches. 8 A typical large-scale alpaca operation may graze from 1,000 to 50,000 alpacas (Fernandez Baca 1971). A herd usually contains between 100 to 500 animals. Shepherds are given charge of herds for a particular season and instructed where to graze the animals.

In the wet season, herds are kept close to the ranch headquarters because forage is abundant and this is the season of most animal management. During this period females give birth and are rebred. Yearling females (open females) are bred in January, and adults are bred from February to March. Early breeding of replacement females allows first time breeders to give birth in December the following year. The young (crias) are therefore older and stronger going into the dry season. After the termination of the cria season alpacas are dipped for control of ecto-parasites. The long dry season constitutes a major bottleneck for grazing animal production. Transhumance is sometimes practiced so herders can take advantage of remote bofedales which occur in high glacial valleys. During this season the herder lives with the animals and distributes them among bofedales. Grazing of bofedales during the dry season begins several hours after sunrise because the pastures often freeze solid during the night. No attempt is made to supplementally feed alpacas. It is commonly believed by many producers that alpacas are unable to consume supplements (personal observation). Others feel that alpacas will reject pelleted supplements and cultivated forages given choice over natural range forages. Alpacas weights peak near the end of April and decline throughout the dry season (Sumar, unpublished). Lactation continues until weaning in late September. In October, animals are returned to pastures near the ranch headquarters for shearing. At this time females are in the last trimester of gestation and arrival of rain is critical for proper pre-partum nutrition. Dipping for ectoparasites is repeated in November. Some operations now shower alpacas instead of dipping in order to reduce stress and chance of ear infections.

Alpaca Diets No literature is available on dietary quality of alpacas grazing native plant communities. Two studies examined the botanical composition of diets. Antezana (1972) listed species preferred by alpacas using ocular estimates of forages consumed. In his southern Peru study near Pitumarca and San Pablo, he identified 26 species of primary importance to alpacas. 10 Bryant and Farfan (1984) used microhistological examination of fecal material to determine diets of adult female and subadult alpaca grazing a Festuca / Calamagrostis association (Salaverry 1981) at La Raya Peru. They found that dry-season diets were largely grass, Carex sp. and Juncus sp. Among the most consumed grasses and sedges were Eliocharis albibracteata, Festuca dolichophylla and Stipa brachiphylla. Preference indicies showed Stipa brachiphylla and Calamagrostis vicunarum to be highly sought after grasses. Bryant and Farfan (1984) showed highest forb consumption in June (20% of the diet) and lowest in September (12%). Alchemilla pinnata, Trifolium amabile and Rununculus limoselloides received high preference scores. Large quantities of seed parts were found in the feces of alpacas during the dry season. A high of 21% seed was recorded during September. The authors point out that seed consumption could help explain how alpacas cope energetically during the dry season when forage is in short supply. When the studies of Bryant and Farfan (1984) and Antezana (1972) are compared, six species in common are listed as highly preferred alpaca forages. These are: Alchemilla pinnata, Calamagrostis vicunarum, Muhlenbergia 11 fastigiata, Muhlenbergia peruviana, Poa sp. and Trifolium amabile (Farfan 1982).

Alpaca Digestive Morphology and Physiology The unique digestive morphology and physiology of a grazing animal sets limits on the forage types it is able to consume and utilize. Although alpacas are ruminants in the strict sense of the word (that is they chew a cud) they evolved separately from "true ruminants" and their stomachs are not clearly divided into four compartments (Walker 1964). The camelid stomach consists of three major compartments. Vallenas et al. (1971) described the and stomach, which structurally are similar to each other and to the stomach of the alpaca. The forestomach of camelidae differs from "true ruminants" being divided into forward and rear sacs by a muscular pillar. Unlike the rumen, the major part of the mucosa in the central forestomach is lined with glandular epithelium. The rest is lined with stratified squamous epithelium without papillae (Vallenas and Stevens 1971b). Both sacs of the forestomach contain recessed glandular sacules which are extruded during the gastric contraction cycle. This complex muscular and mucosal arrangement has no counterpart in the advanced ruminant. 12 Function of the glandular lining of the forestomach has been suggested by several authors. Schmidt-Nielsen et al. (1957) and Moir (1968) speculated that the saccular mucosa contains cardiac glands. Schmidt-Nielsen (1964) suggested that the glandular lining function as accessory salivary glands. Vallenas et al. (1971) indicates that this mucosa may be adapted for absorption of volatile fatty acids (VFA). He also points out similarities in the structure of epithelial cells of the cardiac region in the llama to that of the gall bladder and small intestinal- epithelia that are known to maintain high rates of absorption.

Vallenas and Stevens (1971b) studied the relationship between forestomach pH and concentration of VFA, and compared their findings with sheep. This study suggested that the llama and guanaco are more efficient in neutralizing fatty acids than sheep. Rubsamen and Engelhardt (1978) confirm this hypothesis, finding a higher buffering capacity in the forestomach fluid of the llama compared to sheep. They also found VFA absorption rates to be 2 to 3 times faster than has been noted from the forestomachs of "true ruminants." Increased buffering capacity in camelids forestomachs could be the result of more rapid VFA absorption or greater salivary buffering capacity. Both mechanisms appear to operate in buffering the forestomach of "true ruminants." 13 Eckerlin and Stevens (1973) studied bicarbonate secretion and solute absorption in the llama forestomach. Bicarbonate concentration in the contents of the glandular sacs in compartment one was found to be higher than the rumen of "true ruminants." Further investigations showed that the bicarbonate concentration in a temporarily isolated second compartment increased rapidly after feeding. Ortiz et al. (1971) found alpaca saliva to have a pH of 8.6. In general, the alpaca saliva concentrations of K"*" and Na"*" were similar to those of sheep and cattle, although alpaca had higher salivary HCO3 and HPO4 than cattle, and lower HPO4 concentration than sheep. Feeding increased saliva flow five fold. The authors speculate that increased buffering ability of the camelid forestomach when compared to sheep may be because of greater salivary flow in relation to the stomach size. Lucio (1969) examined the tissues of a llama stomach for digestive enzymes. He found a relatively high concentration of neutral proteases in the glandular tissue of the second compartment, but noted no cellulase or amylase in those tissues or the remaining glandular tissues of compartment one, two, and the proximal 4/5 of compartment three. 14 Besides unique digestive properties, South American camelids have other physiological differences when compared to sheep and cattle. Blood glucose and gluconeogenesis activity in the liver are higher in camelids than true ruminants (Huaman 1974). Camelids also express special respiratory adaptations to high altitudes. These adaptations include smaller elliptically shaped red blood cells (Reynafarje et al. 1968) and increased blood affinity for oxygen (Bartels et al. 1963).

Digestive Efficiency and Forage Consumption Although controversy exists, camelids appear to have greater digestive efficiency than cattle or sheep when consuming low quality forages. Fernadez Baca and Novoa (1966) conducted caged digestibility trials with three sheep and three alpacas. The animals, in separate experiments, were fed oat hay followed by a species of aquatic Scirpus. Dry matter digestion coefficients were found to be 50.0% and 75.1% for sheep and alpacas on oat hay, respectively, and 57.9% and 75.2% for Scirpus sp. Bardales Ramirez (1969) also reported higher digestibility coefficients for dry matter and crude fiber by alpacas. San Martin et al. (1982) and Oyanguren (1969) in similar studies reported higher 15 means for alpaca digestibility of dry matter and crude fiber but failed to show significant differences compared to sheep.

Hintz et al. (1972), conducted digestibility trials with four ponies, two sheep, one llama, and one guanaco fed alfalfa pellets and another complete pelleted diet. They found the camelids were significantly more efficient in digestion of dry matter, neutral-detergent fiber, acid-detergent fiber and cellulose. In a later study, Hintz et al. (1976) failed to find increased ability of camelids to digest fiber. The feed used in this study, however, contained lower fiber than feeds of earlier studies. In addition, they used the lignin ratio technique which requires caution when interpreting results due to variable digestibility of lignin. Vallenas and Stevens (1971a) found that compartments one and two of the camelid stomach undergo cyclic activity similar to that of the reticulum and rumen in cattle and sheep, but with greater frequency of contraction and rumination cycles. Hintz et al. (1972) suggested that the increased motility of the camelid digestive tract may be a major reason for any increase observed in digestive efficiency of camelids on high fiber diets. 16

Sillau et al. (1973) found greater In vitro activity (gas production) from the rumen microbes of sheep than alpacas, suggesting that increased efficiency of alpaca digestion may not be the result of superior fermentation by rumen microbes. Bardales Ramirez (1969) conducted tests with alpaca and sheep rumen fluid to compare the j_n vitro digestibility of several forages. No differance was reported. Hungate et al. (1959) ruled out the possibility of significant large intestine or cecum (equid-like) digestion in South American camelids when he found little fermentation in these organs in 25 Old World camels. In contrast, Esquerre and Samaniego (1980) measured the proximal colon of alpacas to be ten times as large as the cecum, suggesting its possible importance in a large intestine microbial digestion process. In a penned study of llama and sheep forage consumption, Riera and Cardozo (1970) found lower dry matter consumption for llamas than sheep when given free access to alfalfa. Llamas consumed 2.1% of their body weight per day and sheep consumed 4.3%, thus indicating greater efficiency in llamas. Fernandez Baca and Novoa (1966) also found lower consumption on a per weight basis for alpacas eating hay and Scirpus sp. 17 Decreased consumption per body weight likely is related to slower passage of ingesta through the gastro-intestinal tract of camelids. Flores and Valdivia (1973) compared the mean retention time of marked digesta within the alimentary tract of alpacas and sheep. Mean passage time for alpacas was 50.3 hours compared to 43.2 hours for sheep. Efficient digestion of high fiber grasses may be an adaption to high altitude life where fibrous grasses abound. Physiological adaptions may be: 1) increased motility of the camelid stomach, 2) increased ability to buffer and absorb VFA, and 3) decreased passage- rate in the gastro-intestinal tract. CHAPTER III DIETARY SELECTION OF ALPACAS GRAZING ALTIPLANO AND BOFEDAL RANGELAND SITES

Bofedal and Altiplano rangeland plant communities are major sites grazed by alpacas in the Andes mountains of southern Peru. Knowledge of alpaca diet selection may help explain seasonal dietary nutrient deficiencies encountered on these sites. This study describes forage availability and diet botanical composition for each site.

Description of the Study Areas La Raya, High Mountain Site The study site, chosen as representative of alpaca production in the high mountain zone of southern Peru, was the 6,323 hectare National Center for South American Camelids at La Raya. This experimental ranch is located in the rugged mountains along the roadway between the cities of Cuzco and Puno (14° 30'S 71° W). The ranch is jointly operated by the Institute for High Altitude and Tropical Veterinary Research (IVITA) and the Peruvian Ministry of Agriculture (CIPA). The La Raya facility provided access to a herd of 4,349 alpacas and served as a center of operation for the project.

18 19 Topography of La Raya consists of a small flat valley floor with steeply sloping sides (30-60%). The valley floor elevation is 4,200 m and the surrounding mountains rise to 5,472 m. Abundant bofedales are located in high glacial cirques at approximately 5,000 m. Melt water from ice fields which lie above 5,200 m irrigate the bofedales year-round. Valley floor soils are characterized by dark highly organic epipedons. They are low in available nitrogen and phosphorus due to slow organic matter decomposition imposed by the cold climate (Drosdoff et al. 1960, Molina and Little 1981). Bofedal soils described by Wilcox (1982) were classified as typic cryohemists. They were poorly drained and moderately permeable. Organic matter was between 60 and 70%. The pH was lower than on upland sites. Upland soils at La Raya appear to be similar to those described by Wilcox (1982) as cryoboralfs. The pH was between 4.6 and 6.0. Organic matter in upper epipedons ranged from 4 to 8%. In a normal year approximately 80% of the precipitation falls during the period of October through May. Xeric conditions exist from July though September. Southern Peru's climate in 1983 was drier than normal because of "El Nino," a warm ocean current which periodically approaches 20 the Peruvian coast. Mean annual temperature at La Raya (1972-1978) is 6.52 C, and annual precipitation averages 956 mm (Holgado et al. 1979). Figure 1 shows precipitation at La Raya during the study period. Monthly average maximum and minimum temperatures are shown in Figure 2.

The dominant vegetation at La Raya was tall, robust perennial bunchgrasses interspersed with very short mat forming layers of forbs and sedges. Annuals contributed very little to the flora. Bofedales were primarily low mat-like communities of muscoides, Carex ecuadorica and Plantago sp. Boundaries between bofedales and other plant communities were distinct. Islands of the grasses Calamagrostis rigescens and Calamagrostis antoniana were common. To date there is no comprehensive floral classification of La Raya. Salaverry (1981), in a non-quantitative survey of La Raya vegetation, classified the major grass species into 4 general associations. These were: 1) Festuca dolichophylla / Calamagrostis vicunarum located in the valley bottom between 4,100 and 4,300 m elevation, 2) Festuca riqida / Stipa obtusa located on the sloping hillsides up to 4,800 m elevation, 3) Calamagrostis antoniana / Scirpus rigidus located on slopes of 30° to 45° between the altitude of 4,120 m and 4,600 m and 4) 21 280s- 220-

200- La Raya ^ ISO­ Chuquibambilla 's J 160- I 140 H GO a 120- o a. lOOH

60-

A S 0 N Months

Figure 1. Mean daily precipitation (mm) recorded at La Raya and Chuquibambilla from April, 1983 to April, 1984. No data were recorded during April and May at Chuquibambilla. 22

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14-

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G '0-1 La Raya - S Chuquibambilla «

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Months

Figure 2.' Average daily maximum and minimum temperatures (°C) recorded at La Raya and Chuquibambilla between April, 1983 and April, 1984. 23 Calamagrostis antoniana / Scirpus rigidus located on rocky soils with a greater component of lichens in the vegetation. Bofedales were not included in his survey.

Chuquibambilla, Altiplano Site Chuquibambilla was chosen as a site representative of the Altiplano. The ranch functions as an experimental center for the University of the Altiplano at Puno (UNTA) as well as a commercial sheep operation. It is located 65 km southwest of La Raya on the road to Puno. Research at Chuquibambilla was conducted in the Buena Vista section of the ranch. Elevation is 3,190 m and topography is flat. Precipitation was similar to that of La Raya (Figure 1) although Chuquibambilla received slightly less rain during the wet season. Chuquibambilla had lower minimum temperatures and higher maximum temperatures during the dry season (Figure 2). Vegetation was predominately robust cool season bunch grasses with low growing sedges and forbs filling the interspaces. Velarde Vilca and Astorga (1984) described three range sites at Chuquibambilla: 1) a site dominated by Festuca dolichophylla located on flat topography with deep dark red soils of sandy-clay texture (Pucara series), 2) a site dominated by Calamagrostis antoniana also located on 24 flat areas but with relatively deep, dark loamy soils (Sorani series) and 3) a site dominated by Festuca dichlocada located on hills with red, sandy loam soils (Pusi series).

Range sites in the Buena Vista pasture were predominatly Pacara and Sorani. Above-ground plant biomass dry matter in 1982 was 1,237 kg DM/ha on the Pacara site, 1,957 kg/ha on the Sorani site and 1,240 kg/ha on the Pusi site (Velarde Vilca and Astorga 1984). Soils appeared to be poorly drained. Standing water was common in the wet season.

Methods Study Design Alpaca diets were sampled under typical herding conditions at the Altiplano and high mountain bofedal sites by selecting a representative band of animals as a study herd in each of these locations. For the bofedal site I chose to observe the La Raya alpaca herd which grazed during the dry season in the high bofedales of Jatun Cuchu (bofedale #1) and Juchi Cuchu (bofedale #2). These sites appeared similar to alpaca dry season grazing sites at the expansive alpaca producing ranch at Marangani. Marangani, which borders La Raya to the northeast, is the second 25 largest alpaca producer in Peru. Over 50,000 alpacas were grazed at Marangani in 1983 and 1984. Because alpaca were not numerous in the Chuquibambilla area, a band of approximately 100 sheep was selected to represent typical herding practices in this region. This band remained in the Buena Vista pasture during the entire study. During sampling periods, the experimental alpacas were allowed to graze freely within the study bands at both sites. Sampling periods were chosen to correspond with major management or physiological events in the annual life or production cycles of alpacas. Diets were sampled during eight periods in 1983/1984 at the La Raya and Chuquibambilla sites. All collections at Chuquibambilla were in the Buena Vista pasture. La Raya collections were made at the Jatun Cuchu bofedal until the October sampling when the shepherd moved the band to the Juchi Cuchu bofedal. Vegetation cover was sampled before diet collections for all sampling periods at Chuquibambilla. Cover was not measured preceding the November 1983 and March 1984 collection periods at La Raya. Sampling at each site was separated by one week because it was not possible to work at both study sites simultaneously. Table 1 lists the sample dates and major events associated with these collection periods. 26 Table 1. Sample dates, conditions and locations associated with collection of alpaca diets at the La Raya (bofedal-^) and Chuquibambilla (Altiplano) sites during 1983 and 1984.

DATE CONDITIONS LOCATION SAMPLED

Jun. 10 early dry season, Altiplano lactation bofedal #1 Jul. 10 dry season, lactation Altiplano bofedal #1 Aug. 2 2 dry season, Altiplano period of weight loss bofedal #1 Oct. 4 late dry season, weaning, Altiplano dipping, shearing bofedal #2 Nov. 7 early wet season, Altiplano last trimester of gestation bofedal #2 Jan. 1 wet season, mid-parturition Altiplano season, breeding bofedal #2 Feb. 2 mid-wet season, breeding Altiplano bofedal #2 Mar. 3 end wet season, late Altiplano parturition, breeding, early bofedal #2 lactation, dipping

^ Bofedal #1 is Jutun Cuchu and #2 is Juchi Cuchu.

Forage Availability Percent foliar, soil and litter cover was estimated before each diet collection period. Cover was determined on La Raya bofedales by the inclined point method (Levy and Madden 1933, Wilson 1960) because it is recommended for 27 short plant communities with single layer canopies (Brown 1954). An inclined rack with ten points angled at 45° and separated by 10 cm was placed at 1 m intervals along two, 50 m transects across the bofedal. Only the first hit on foliage, soil or litter was recorded for each point position. During each sampling period 1000 points were recorded. Percent foliar, soil and litter cover was estimated at the Altiplano site using a modification of the Daubenmire technique (Mueller-Dombois and Ellenberg 1974). Cover was recorded by species within 36, 0.5 m^ quadrats randomly located across the pasture. When estimating cover, the technician was given no requirement that total cover in each quadrat equal 100%. Composition of foliar, litter and soil cover was later adjusted to 100%.

Diet Sampling and Analysis Diets were sampled from four male alpacas at each site. All eight animals were of the huacaya breed. Each was castrated and then underwent four weeks of training. Training involved walking each animal daily for one mile on halter and lead. Alpacas age ranged from two to three years. The experimental animals were maintained on cultivated pastures when they were not involved in diet collections. 28

The alpacas were surgically fitted with esophageal cannulas (Torell 1954, Cook et al. 1958, McManus 1981) and allowed to recover for three weeks before sampling was initiated. Intervenus injected Combelin (0.5 cc) was used as the anesthesia in combination with locally administered Xylocane (2.0%). Cannulas were constructed of 1 inch (outside diameter) PVC tubing (Taylor and Bryant 1977). A length of 3 inches was needed to accommodate for swelling. The cannulas were later shortened by 1 inch after swelling had diminished. Daily injection of 2.5 cc of Tantum (Benzidamina clorhidrate, Instituto Sanitas, S.P.S.A., Peru) helped reduce swelling problems. Five days preceding a sampling period, experimental animals were moved from cultivated pasture and combined with the study herd. This allowed the animals to become accustomed to both the study herd animals and the native vegetation. Diets were collected using screen bottom bags each morning at 9:00 AM for five days while the experimental alpacas grazed freely within the study herd. A collection period lasted 30 to 60 minutes depending on forage availability.

After collections the experimental alpacas grazed with the free-ranging herd until dusk. At night the free-ranging 29 herd and the experimental alpacas were penned as is typical for the area. Fasting at night encouraged grazing during sampling.

Upon collection, each extrusa sample was thoroughly hand-mixed and air dried. Samples were then individually ground to pass through a Wiley mill fitted with a 0.5 mm screen. Botanical composition was determined by the microhistological technique (Sparks and Malecheck 1968). Two slides were made per sample and 10 fields read per slide. Plant species frequency was recorded for each slide following the procedure of Scott and Dahl (1980). One slide per sample was later re-read for occurance of seed and flower parts. Only grass seed and flower part frequencies were recorded. A completely randomized, split plot analysis of variance was used to test for the main effects of site and month on diet composition within forage classes (Appendix Tables 23-25). Variation between animals was considered experimental error and days as sample variation. Site (high mountain bofedal vs Altiplano) was designated the treatment. Sample periods (months) were split out as repeated measures. The GLM procedure of the SAS (1982) computer program was used for the analysis. Mean comparisons were by Least Significant Difference Tests (Snedecor and Cochran 1980). 30 Results Forage Availability Altiplano forage availability was distinctly different than on the bofedal. The Altiplano was predominately grass year-round; forbs were abundant only during the wet season. In contrast, the bofedal site was dominated by sedges and forbs year-round (Table 2).

Table 2. Percent forage foliar cover during eight months of and 1983 and 1984 on the study sites in southern Peru.

Months Sites J J A 0 N J F M Forage Class Dry Season Wet Season

Bofedal Grass 26 21 17 18 * 22 28 * Sedges and Reeds 46 46 48 34 * 36 42 * Forbs 23 30 30 44 * 38 27 * Moss 2 2 4 3 * 2 1 • Litter and Soil 3 1 1 1 * 2 2 * Altiplano Grass 68 52 72 75 67 72 60 60 Sedges and Reeds 15 25 7 11 11 10 13 9 Forbs 6 1 1 1 3 16 20 Moss ^ 1 1 T 2 2 3 7 Litter and Soil 14 16 19 13 19 13 8 4

T represents values less than 0.5%. No data available for this period. 31 Thirty-five plant species were identified at the bofedal site (Appendix Table 11). Averaged across collection periods, vegetation cover was 22% grass, 42% sedges and reeds, and 33% forbs. Important grasses included Calamagrostis antoniana, Calamagrostis rigescens and Festuca dolichophylla. Grass cover was highest during the wet season and lowest during the late dry season. The major sedge species included; Eleocharis albibracteata, Distichia muscoides and Luzula peruviana. Juncus sp. (reeds) were abundant in the wet season.

Bofedal forbs were most abundant during the late dry season. At this time, forbs were largely Plantago tubulosa, Hypochoeris taraxacoides and Werneria pygmaea. Alchemilla diplophilla was the major wet season forb. Forty-seven plant species were identified on the Altiplano pasture; 20 were also found on the bofedales. Vegetation cover averaged 66% grasses, 13% sedges and reeds, and 6% forbs. The predominate grasses at the Altiplano site were Festuca dolichophylla, Calamagrostis antoniana and Muhlenbergia fastigiata. Sedges were mostly Carex ecuadorica. Small percentages of Alchemilla pinnata, Lepidium sp., Hypochoeris stenocephala and Geranium sessiliflorum represented the dry season forbs. Larger percentages of Arenaria sp., Nototriche sp., Trifolium amabile and Oxalis sp. were present in the wet season. 32 Moss and lichen availability ranged from 2 to 4% on both sites except in March on the Altiplano when 7% moss was recorded. Exposed soil cover and litter were greater at the Altiplano site. Soil cover in the Altiplano reached a high of 4% in the late dry season. Soil on the bofedales was relatively unexposed year-round.

Diet Botanical Composition Alpaca diets on the bofedal were consistently higher in sedges and reeds, and lower in grasses than in diets on the Altiplano. Year-round contribution of species to bofedal alpaca diets were highest for sedges and reeds (59%), followed by grass (26%), and forbs (14%). Alpaca diets on the Altiplano site were highest in grass (51%), followed by sedges and reeds (33%), and forbs (15%). ANOVA by forage class was significant (P<0.05) for the main effects of site and month for grass and sedges and reeds. There were also significant (P<0.05) site/month interactions for these forage classes. The F test for site within the forb forage class was not significant (P>0.05) (Appendix Tables 23-25). Dietary composition during the early dry season (June and July) did not differ (P>0.05) between sites for each forage class (Figure 3). As the dry season progressed dietary composition increasingly differed between the two sites. August grass consumption on the Altiplano 33

70- BOFEDAL 50- Forbs ALTIPLANO • -#

30-

10- 1 I T" —I 1 r- A 0 N J F M DRY SEASON - WET SEASON •

Figure 3. Percent dietary botanical composition for alpacas grazing the bofedal and Altiplano sites. 34 increased to a high of 67% while grass consumption by alpacas on the bofedal greatly decreased from 47% in July to only 6% in August. In August the bofedal diets were largely sedges and reeds (81%). During the late dry season (August, October and November), sedge and reed consumption was higher (P<0.05) on the bofedal than on the Altiplano site. Highest consumption of sedges and reeds on the Altiplano site was during the mid-wet season (February).

Although forbs did not contribute greatly to the annual diet, they consistently were present throughout the year. Forb consumption at both sites was highest during the early dry season (June and July) (21-26%) and remained between 9% and 16% of the diet during the late dry and wet seasons. Tables 3 and 4 rank plant species within forage class in order of their annual contribution to alpaca diets on the bofedal and Altiplano sites. Plant species, averaged across seasons, which made up more than 5% of the diet or more than 10% in one collection period were included. The were re-ranked by their seasonal dietary contribution. Grasses in alpaca diets on the bofedal were largely Calamagrostis antoniana and Festuca dolichophylla (Table 3) while on the Altiplano, Festuca dolichophylla and Muhlenbergia fastigiata were predominant (Table 4). During the wet season Calamagrostis rigescens was important to 35

Table 3. Bofedal site dietary botanical composition listed in order of annual contribution and then re-ranked by season.

Rank by season Annual Early Late Early Late Forage class rank dry dry wet wet

Grass Calamagrostis antoniana 1 1 2 3 3 Festuca dolichophylla 2 2 1 2 1 Calamagrostis rigescens 3 3 3 1 2 Sedges and Reeds Juncus sp. 1 1 1 2 1 Eleocharis albibracteata 2 2 2 1 2 Carex ecuadorica 3 4 5 3 5 Distichia muscoides 4 5 3 4 3 Luzula peruviana 5 3 4 5 4 Forbs Hypochoeris taraxacoides 1 1 4 2 1 Werneria heteroloba 2 3 1 4 Stylite¥ andicol¥ 3 5 2 1 Taraxicum officinale 4 2 5 5 AlchemiTTa diplophlla 5 4 3 3

•'• Seasons; Early Dry (Jun., Jul.), Late Dry (Aug., Oct.), Early Wet (Nov., Jan.), Late Wet (Feb., Mar). * Species not found in the diet during this season. 36

Table 4. Altiplano site dietary botanical composition listed in order of annual contribution and then re-ranked by season.

1 Rank by species'^ Alinua l Early Late Early Late Forage class iran k dry dry wet wet

Grass Festuca dolichophylla 1 2 1 1 2 Muhlenbergia fastigiata 2 1 2 2 1 Calamagrostis antoniana 3 3 3 4 4 Calamagrostis vicunarum 4 5 5 5 • Poa gymnantha 5 4 4 3 3 Sedges and Reeds Eleocharis albibracteata 1 2 2 1 1 Carex ecuadorica 2 1 1 2 2 Juncus sp. 3 3 * 3 3

Forbs Alchemilla pinnata 1 1 1 1 1 Trifolium amabile 2 2 • 3 * Gomphrena mexicana 3 * * 2

^ Seasons; Early Dry (Jun., Jul.), Late Dry (Aug., Oct.), Early Wet (Nov., Jan.), Late Wet (Feb. Mar.). * Species not found in the diet during this season. 37 alpacas on the bofedal, while Muhlenbergia fastigiata and Festuca dolichophila were ranked highest on the Altiplano. An increase of Poa sp. was observed in Altiplano diets during the wet season.

Juncus sp. and Eleocharis albibracteata were the dominant sedge species in bofedal diets year-round. The large increase in dietary sedge composition recorded in August on the bofedal site was primarily due to increased consumption (60%) of Juncus. Eleocharis albibracteata and Carex ecuadorica were the dominant dietary sedges at the Altiplano site. Forb species composition varied more between seasons than other forage classes. Alpacas grazing the Altiplano site consumed Alchemilla pinnata in greater amounts than other forbs during all seasons and consumption peaked in July at 20%. Top ranked bofedal forb species changed each season in relation to forage availability. Hypochoeris tarxacoides comprised 10% of the diet during July, whereas Werneria pygmaea comprised 11% of bofedal diets during the nutritionally critical month of August.

Frequency of seeds and flower parts in the diets of alpacas grazing the bofedal site was greatest during the dry season. Diets of alpacas grazing the Altiplano contained the most seeds during the late wet season (Table 5). 38

Table 5. Frequency of grass seeds and flower parts in the diets of alpacas grazing the study sites.

Month JJAONJFM Site Dry season Wet season

Bofedal 14 22 20 7 13 7 5 8 Altiplano 11 3 5 7 9 15 20 17

Discussion High year-round availability of grass on the Altiplano was consistent with greater amounts of grass in the diets of Altiplano alpacas. Similarly, on the bofedal where sedges and reeds were abundant, they were the major dietary component. The rapid decline in dietary grass on the bofedal during July (Figure 3) may be due to depletion of grasses on this site. In October the shepherd moved the animals to Juchi Cuchu (bofedal #2) and grass in the diet again increased. 39 Diet composition at the Altiplano site was similar to diets reported by Bryant and Farfan (1984) in their study on a Festuca / Calamagrostis pasture at La Raya. Grass was the major component of dry season diets in both studies with Festuca dolichophylla being the major species. Similarly, Eleocharis albibracteata and Carex ecuadorica were the major sedges and Alchemilla pinnata was the major dry season forb. Bofedal forage availability and alpaca diets from my study showed little similarity to those of Bryant and Farfan (1984).

Bofedales described in my study were similar to communities described by Weberbauer (1936) as Distichia moor, by Tovar (1973) as bofedales and by Wilcox (1982) as a Plantago tubulosa / Hypochoeris taraxacoides community type. The bofedales in my study differed from those described by Wilcox (1982) by having a greater component of Distichia muscoides and Calamagrostis antoniana. Abundance of bofedal forbs during the late dry season was mostly due to increases in unpalatable forbs such as Plantago tubulosa. Bryant and Farfan (1984) also found Plantago tubulosa to be relatively unpalatable. Similarly, Altiplano forb cover increased during the late wet season due to a flush in growth of Arenaria sp. growing in the pasture. In disagreement with a report by Tapia Nunez and 40 Flores Ochoa (1984) who described this species as palatable, alpacas in my study consumed very little. Abundance of forbs during different seasons at the two sites may be due to soil moisture differences. Because bofedal pastures are saturated with water year-round, the major constraint on plant growth is likely temperature. Increase of forbs on the bofedal during the late dry season corresponds well with increasing minimum daily temperatures in October (Figure 2). Altiplano soils have very low water potential during the dry season (Velarde Vilca and Astorga 1984). In • addition, minimum daily temperatures are lower during this season than at the bofedal site. Plant growth is likely limited by both factors. Forbs in this environment appear adapted for rapid growth as soil water recharges during the warm wet season. Mat-like plant communities were found at both sites but were more common on the bofedales. Dwarfism and mat development are alpine adaptions which protect plants from the cold and keep them out of reach from grazers (Mooney and Billings 1960, Whittaker 1975). Low growing forbs composed up to 44% of the bofedal vegetative cover but were not a major dietary component. Low forb consumption on the bofedal may be partially the result of forb unavailablity due to their short stature. 41 The number of species found in alpaca diets was similar to numbers reported for cattle, sheep, and goats (Squires 1982). A relatively small number of species (8-12) composed the bulk of alpaca diets at both sites. The optimal foraging model of Owen-Smith and Novellie (1982), predicts that animals widen the range of acceptable plant species in their diets as available forage declines. This was not apparent in my study. On both study sites alpaca diets contained more species diversity during seasons of highest forage availability. Bryant and Farfan (1984) reported that alpacas diets contained large amounts of seeds during September. They hypothesized that seeds may be energetically important for alpacas during the dry season. Alpacas grazing the bofedal site in my study also consumed large amount of seeds during the dry season; however, seed consumption peaked in July and August. Grass seed consumption by alpacas on the Altiplano site followed a dissimilar pattern. At this site, grass seed consumption was greatest in the late wet season when seeds were most abundant. Decline of seeds in Altiplano alpaca diets during the dry season may be due to seeds becoming unavailable after heavy grazing.

Alpacas appear to be highly adaptable grazers and vary their diets according to forage availability. When grass is 42 available, it comprises the bulk of alpaca diets. On sites with low grass availability and abundant sedges, sedges dominate the diet. Forbs were not major dietary components at either site, but their importance can not be ignored. I did not determine the nutritional value of each forage class independently; therefore, the small amount of forbs consumed in the dry season could have been of major nutritional importance. CHAPTER IV DIETARY NUTRIENT CONTENT OF ALPACAS GRAZING ALTIPLANO AND BOFEDAL RANGELAND SITES

No published literature on the nutrient content of free-ranging alpaca diets exists. This study was designed to determine the seasonal diet nutritional quality of alpacas while grazing two important rangeland sites in the Andes Mountains of southern Peru.

Methods The study design and diet collection techniques were discussed in Chapter III. Extrusa samples were air dried, ground to 0.5 mm in a Wiley mill and then chemically analyzed for percent nitrogen and percent ^ vitro organic matter digestibility (IVOMD). Percent nitrogen was determined by the micro-Kjeldahl procedure (A.O.A.C. 1970). Crude protein (CP) was estimated from nitrogen by using a multiplication factor of 6.25. Percent IVOMD was determined according to the procedures of Van Soest (1970). Samples were inoculated for 48 hours with steer inocula and later washed with neutral detergent solution. Each batch of samples was standardized

by concurrently analyzing six sorghum samples of known vn

43 44 vivo digestibility for cattle. Duplicates of all diet samples were analyzed. A sample was reanalyzed if its two results were not within 10% of each other (Harris 1970). A completely randomized, split plot analysis of variance was used to test alpaca diets for differences in CP and IVOMD. Individual alpacas were considered the unit of replication. Site (bofedal vs Altiplano) was designated the treatment. Sample periods were split out as repeated measures.

The number of fistulated alpacas needed for precision sampling of CP and IVOMD (95% confidence, within 10% of the mean) was estimated by analyzing intra-animal variability in dietary CP and IVOMD by the following formula:

t2 s2 n=

where t is the value of the t statistic at n-1 degrees of freedom, s is the standard deviation, and d is the half-width of the desired confidence interval (Steel and Torrie 1980). 45 Results Crude protein (CP) in alpaca diets increased during the wet season and declined during the dry season at both sites. ANOVA showed that the main effects of site and month were significant (P<0.01). There was also a significant site/month interaction (P<0.01). CP in alpaca diets on the bofedal (12.3%) was higher than Altiplano diets (10.2%) when averaged across collection periods. Dietary CP was lowest during June (7.1%) and July (8.0%) on the Altiplano pasture and bofedal, respectively (Figure 4). CP in alpaca diets was highest during February in the Altiplano and in March on the bofedal. Diets on the bofedal had higher (P<0.05) -CP content than in the Altiplano during June, August, October, November, January and March. Bofedal alpaca diets were lower (P<0.05) in CP than on the Altiplano site during July. Dietary IVOMD was greatest during the wet season and gradually decreased during the dry season on both sites (Figure 5). ANOVA of these data showed the main effects of site and month were significant (P<0.01). A site/month interaction was also significant (P<0.01). IVOMD was lower at the bofedal site (63%) than in the Altiplano (64%) when averaged across collection periods. IVOMD was lowest in Altiplano and bofedal alpaca diets during August (49%) and October (50%), respectively. 46

•J) 4J 0) CC -r-4 < Ti Q) SI +J C m •H LU cn +J •H

-reH '-' -< a en u a c ^ a ^ U} > •H U-( 0 c c o CO 0 (TJ u r-l ^- a c e»P •H LD 4J o

c i-H •H fTJ 0) -O < +J o 0 V4-I u, 0 aja 0) 0) TJ x: 3 4J i-i U Cn c 4-1 •rH c N OJ (TJ u V-, Wl D^ 0) ^ w <0 I—I 1—I—I \—I—I 1 1—I—r u • 03 ^ a r^cDio^pocvj — oooor^cD 1—1 0) (t! u 3 U-J (%) u!9jojd epnjQ D^ 0 47

dp in a: ON < X 2 an d ) wi t c .-^ m S 03 LU n T! Li. (IV ( bofe (

Z 4-> Q) < •H x: -D in g t > U) N CJ 03 o CO •H cr ^Z T3 ^.- W J-i 03 c O U -u 03 O o o 2 i c m a o f a l c ^o (T3 cn < die t o or q U\ 0 -J 4JI^ 3 -D c c •H -H cn 2 4-» ^ • 3 ~D u -- 4J u <-^• H (D cn CL 0 I I I I I I I I I I I I I I I I I I I I I I I I I I I I I I » u o c C Q CO CM CD ^ o ^ ^ ® • !i; ^ 00 1^ 1^ U) in T: .—' CO (O ^ ^ ^ •H Q. Q U-. •r^ J-1 c *J (%) ai/^OAi 3 0 .— (T> U < 48 Animal numbers needed to precisely sample CP were generally higher than required to sample IVOMD. Less alpacas were required to sample CP during the wet season than during the dry season (Table 6).

Table 6. Number of fistulated alpacas needed to estimate CP and IVOMD with 95% confidence within 10% of the mean during five day collection periods.

Month Site J J A 0 N J F M

Altiplano

CP 8 4 7 0 C N 7 4 2 3 IVOMD 2 3 4 5 3 3 1 Bofedal CP 8 6 11 7 5 3 4 2 IVOMD 2 1 7 3 2 2 2 1

Discussion No nutritional requirements for alpacas are available for comparison with my data. The major importance of this study was to provide a seasonal view of nutritional trends encountered by alpacas on the Altiplano and bofedal sites. Throughout the study, dietary CP and IVOMD from both sites closely paralleled seasonal precipitation. 49 Alpaca management, because of low female fertility, should be directed towards proper nutrition of the reproducing female (Fernandez Baca 1971). A critical period in the female alpacas annual life cycle is the last trimester of gestation (September-January). In southern Peru, females enter this period in poor condition after surviving the long dry season while carrying a developing fetus and lactating. The return of wet season forage is critical for improving the female's condition before parturition and breeding in January. CP maintenance requirements of a dry pregnant ewe and lactating ewe (50 kg with twins) are 9.3 and 11.5%, respectively (NRC 1975). Similarly, goats require 10.9 and 13% CP (NRC 1981). In my study Altiplano alpaca diets were at or below 9.3% CP from June to early November. Alpacas grazing the Altiplano site were deficient in dietary CP for optimal lactation or fetal growth if requirements are similar to sheep or goats. Bofedal diets were below 9.3% CP only in June and July (second trimester of gestation). An important seasonal difference in dietary CP between sites was in August when Altiplano alpaca diets plunged to a yearly low of 6.1% while bofedal alpacas had values up to 11.0%. Bofedal alpaca diets remained higher than on the Altiplano for the next 50 three months. These data indicate, that for optimal CP nutrition, alpacas grazing the Altiplano site could have been moved to the bofedal in mid July or protein should be supplemented otherwise.

Rittenhouse et al. (1971) reported a strong relationship between organic matter digestibility and in vivo digestible energy. Depressed digestibility at both sites during the nutritionally critical mid and late dry season might indicate energy deficiency during these months. Depressed digestibility of late dry season forage may be the result of low dietary CP (Jones 1972). Protein acts as a nitrogen and sulfur source for rumen microflora, altering its cellulose digesting ability. Cook and Harris (1968) demonstrated that adding protein to sheep diets increased gross energy, dry matter and protein digestibility. In contrast to CP results, dietary IVOMD means were similar between sites. One explanation may be related to high sedge consumption at the bofedal site. Digestibility of dry season sedges may be similar to the grasses which made up a large part of the Altiplano diet. Although the bofedal site offered higher CP nutrition than the Altiplano, diets may be energy limited due to sub-optimal organic matter intake. The very low mat-like 51 nature of bofedal plants likely limits the amount of leaf material alpacas can harvest. Grazing management of a bofedal might include daily or bi-daily rotation of the band to nearby upland bunchgrass sites. These plant communities are more easily grazed and appear energy rich but protein poor. Similarly, cultivated pasture might by used in rotation with the Altiplano site to provide a better balance of protein and energy. The minor increase in CP observed during July on the Altiplano site can be explained by a sudden increase in algae consumption (6% of the diet). During July alpacas consumed algae which was growing in several flooded irrigation ditches which crossed the pasture. Algae and other aquatic plants should not be overlooked as a possible source of supplemental dry season protein for Altiplano livestock. Aquatic plants and algae comprise a large percentage of diets of cattle grazing along the shores of Lake Titicaca in southern Peru (personal observation). Plant material might be harvested from the lake and transported to Altiplano livestock during the dry season. CHAPTER V INTAKE OF ALPACAS HOUSED IN METABOLISM CAGES AND FREELY GRAZING A NATIVE ANDEAN PASTURE

Little is known about forage intake of alpacas. Recently, there has been controversy among alpaca producers over what intake values to assign alpacas when estimating proper stocking rates for native ranges. This study directly measured intake of low quality oat hay and higher quality ryegrass in metabolism cages and compared those values to estimated intake of free-ranging alpacas on wet and dry season native rangeland.

Methods Intake in Metabolism Cages Organic matter intake (OMI) was estimated for alpacas in metabolism cages during June of 1983 at La Raya, Peru. Procedures followed the methods outlined by Harris (1970) for digestion balance trials. Eight alpacas were confined in metabolism cages for 14 days preceding the trial in order to accustom them to the feed and unusual surroundings. Four alpacas were fed oat hay (Avena fatua) ad libitum and four were fed ryegrass hay (Lolium perenne). The oat hay had been rained on and was

52 53 similar to low quality dry season forage. The ryegrass was harvested in an immature stage to simulate native wet season forage. Forage was presented each morning at 7:15 AM. The trial lasted for seven days following the accustoming period. Feces were collected using waterproof canvas bags attached to each animal with straps. Placement of the bags was to avoid contamination by urine. Forage, orts, and fecal material were weighed every 24 hours of the trial and 100 g aliquots were saved for oven drying. The stem/leaf ratio of oat hay orts was compared to the feed ratio to test whether alpacas were selecting for particular forage parts in the feed. Ryegrass offered no opportunity for plant part selection because it was entirely leaf. The orts nutritive content was therefore assumed to be the same as feed.

Intake of Free-ranging Alpacas Dry season OMI of freely grazing alpacas was estimated in October (dry season) and again in February (wet season) in a Festuca dolichophylla / Muhlenbergia fastigiata pasture at La Raya. The pasture had been moderately grazed by vicunas before the experiment. Vegetation biomass was measured before each trial by randomly clipping ten, 0.5 m^ plots to a height of 2 cm. The clipped material was air dried, sorted by species, and weighed. 54 OMI was determined by the total fecal collection procedure (Schneider et al. 1955). Six castrated male alpacas were trained to accept sheep fecal collection bags modified to fit alpacas. Preceding each collection period the animals were allowed to graze the pasture for five days. After this accustoming period total feces were collected for five days (Cordova et al. 1978). OMI was calculated by rearrangement of the digestibility equation as follows:

Organic matter fecal output OMI= 1- IVOMD

IVOMD of the diets was determined by concurrently collecting diets with four esophageally fistulated alpacas (see Chapter IV for details). Oat hay and ryegrass from the metabolism cage study were used as standards in the j_n vitro digestibility procedure to adjust wet and dry season forage samples, respectively. Organic matter intake of the alpacas consuming oat hay and ryegrass in metabolism cages were compared to free-ranging OMI during the dry and wet season using t tests. 55 Results Intake in Metabolism Cages No difference was found (P>0.05) between the stem/leaf ratio of the oat hay feed and orts. It was thus concluded that the alpacas were not selecting for plant parts and that nutritional differences between feed and orts were minimal. Therefore, chemical analysis was not performed on the orts.

Intake of eight alpacas in metabolism cages did not stabilize until 12 days into the accustoming period. During the seven day trial alpacas consuming oat hay lost 1.3 kg (60.8 kg starting) and those consuming ryegrass lost 0.7 kg (50.5 kg starting). Percent organic matter digestibility of ryegrass was greater (P<0.10) than for oat hay (Table 7).

Table 7. Organic matter intake (OMI) and digestibility (OMD) of two forages consumed by eight alpacas in metabolism cages at La Raya, Peru.

CP OMD OMI OMI OMI Forage (%) (%) (g/day) (% BW) (g/kg BW^-^S)

Oat Hay 9 64.2 a-^ 688.0 a 1.13 a 31.6 a Ryegrass 14 68.9 b 545.0 b 1.08 a 28.8 a

^ Means in the same column followed by a common letter are not significantly different (see text for alpha). 56 OMI was greater (P<0.05) for oat hay than for ryegrass. When OMI was adjusted as a percent of body weight, the treatment groups were not different (P>0.05).

Intake of Free-ranging Alpacas Vegetation biomass of the pasture grazed by the experimental free-ranging alpacas was greater during wet season sampling (Table 8). Grass production was largely Festuca dolichophylla and Muhlenbergia fastigiata. Sedges and reeds were primarily Scirpus rigidus and Carex sp. Wet season forbs were mostly Oxalis sp. and Trifolium amabile (Appendix Table 21).

Table 8. Vegetation biomass (kg DM/ha) of a La Raya valley bottom pasture during the dry (October) and wet (February) seasons.

Class Dry Season Wet Season

Grass 1721 3016 Sedges and reeds 122 142 Forbs T1 18 Total 1843 3177

^ T represents less than 1 kg/ha 57 Diets collected by esophageal fistulated alpacas were 8.1% CP in the dry season and 12.6% CP in the wet season. IVOMD of diets were 53 and 64% for the dry and wet seasons, respectively. Average alpaca weight was 61.7 kg at the time of dry season sampling and 62.0 kg during the wet season. OMI of the free-ranging alpacas was greater (P<0.10) during the dry season than during the wet season (Table 9). OMI also was greater (P<0.10) during the dry season when expressed as a percent of body weight. When compared to intake values from the metabolism cage study, OMI of free-ranging alpacas was greater (P<0.01) during the dry season than for caged animals consuming oat hay. OMI during the wet season was also greater (P<0.01) than caged alpacas consuming ryegrass.

Table 9. Dietary crude protein , IVOMD and organic matter intake (OMI) of free-ranging alpacas sampled during the wet and dry season at La Raya, Peru.

CP IVOMD OMI OMI OMI Season (%) (%) (g/day) (% BW) (g/kg BW^-^S)

Dry 8.1 53 1112 a^ 1.8 a 50.5 a Wet 12.6 64 979 b 1.6 b 44.3 b

^ Means in the same column followed by a common letter are not significantly different (P>0.10). 58 Discussion There is clearly a need for sound intake estimates for alpacas. Alpaca intake reported in the literature ranges from 1.2 to 2.4% of live body weight (BW) (Fernandez Baca and Novoa 1966, Florez and Valdivia 1973, San Martin et al. 1982). Past studies have used metabolism cages and it is doubtful if these results can be used as a base for practical management recommendations. Results of my metabolism cage study are low compared to other studies but it should be recognized that the alpacas lost weight during this trial. The alpacas never became adjusted to confinement and this may have lowered intake. Excessive weight loss by the alpacas eating oat hay can be explained by examining daily protein intake. Alpacas eating oat hay consumed more organic matter but less protein than those eating ryegrass. Alpacas eating oat hay consumed approximately 2.8 grams protein per kg metabolic body weight (MBW) daily while those eating ryegrass consumed 4.0 g protein/kg MBW. Huasasquiche (1974) found in a nitrogen balance study that alpacas maintained weight while consuming 2.13 g digestible protein/kg MBW per day. In my trial, even if high digestibility of protein is assumed, CP intake of alpacas consuming oat hay was well below the value reported by Huasaquiche (1974). 59 During our study with free-ranging alpacas the animals appeared to behave normally. Intake estimates from this study were towards the upper end of those reported for caged alpacas but were lower than most intake estimates for free-ranging sheep (Cordova et al. 1978) and goats (Pfister 1983). Jeffery and Holder (1971) reviewed intake data on 137 tropical forages by sheep and concluded that the upper boundary for dietary intake was about 83 g DM/kg MBW. This figure is considerably higher than my upper intake estimates expressed on a dry matter basis (60.5 g DM/kg MBW for the dry season and 53.7 g DM/kg MBW for the wet season). Organic matter intake/kg MBW for our study is also lower than reported by McCammon-Feldman (1980) and Pfister (1983) for free grazing goats. Lower consumption may be related to reduced retention time of digesta in the gastro intestinal tracts of alpacas (Flores and Valdivia 1973). Intake of CP in the free-grazing trial was 4.09 and 5.58 g CP/kg MBW. If approximately 50% digestibility of protein is assumed for my study, these values more closely resemble digestible protein values reported by Huasasquiche (1974). Energy intake can be estimated using the familiar factor of 4.4 Meal digestible energy per kg TDN if we assume OMD closely resembles TDN values. This is likely a valid 60 assumption because OM values included all fractions of TDN except the multiplication of ether extract by 2.25, and ether extract is normally a very small percentage of range forages. Using TDN calculations, and IVOMD of the diets, energy intake was 117.8 Meal digestible energy/kg MBW in the wet season and 124.0 Meal digestible energy/kg MBW per day in the dry season. These values were lower than reported for sheep grazing an Altiplano pasture (Fierro 1985). Dry season OMI estimates were greater than during the wet season. This may be the result of two factors. First, forage cell wall contents during the wet season are saturated with water; thus, wet season forage is high in bulk. Intake may be limited by bulk restrictions related to the size of the gastro-intestinal passage. Secondly, alpacas grazing wet season pastures may satisfy their energy needs with less forage because caloric content of wet season forage is greater. Intake regulation in this case is chemostatic (Arnold and Dudzinski 1978). Behavior studies by Rios et al. (1984) support both theories by showing that alpacas spend less time grazing during the wet season.

Low intake findings for alpacas support claims by Fernandez Baca and Novoa (1966) and others that alpacas require less forage than other livestock on a percent BW basis. The Peruvian Ministry of Agriculture recoimnends a 61 conversion ratio of 0.67 alpaca per sheep on range where proper stocking is known (Holgado 1979). Fierro (1985) reported wet and dry season OMI for sheep (47 kg) grazing at Chuquibambilla as averaging 1269 g/day. My wet and dry season estimates for 60 kg alpacas averaged 1046 g OM/day. These data indicate that government stocking recommendations for alpaca may be low. Furthermore, using government guidelines to restock properly stocked alpaca range with sheep may result in overstocking sheep.

It should be noted that estimates of intake are highly dependent on accurate fecal output and forage digestibility measurements. Handal and Rittenhouse (1975) reported that use of a neutral detergent step (Van Soest and Wine 1967) which I incorporated into the basic Tilley and Terry in vitro procedure may cause inflated IVOMD determinations. They reported that higher IVOMD would in turn underesti.nate intake by approximately 10%. Even if adjusted upward by 10% my estimates are lower than reported for other livestock. CHAPTER VI SUMMARY AND CONCLUSIONS

Alpacas are important fiber and meat producers for both subsistance and market-oriented ranchers using high elevation grasslands in southern Peru. Alpacas are able to utilize rangeland above 4,000 m which is unavailable to other livestock. Alpaca management using established sheep production techniques is inappropriate because of the alpacas unique physiology. Southern Peru's high elevation rangeland can be characterized as two grassland zones; the high plains (Altiplano) of the Lake Titicaca basin and the zone of high mountains surrounding the basin. The long Andean dry season (June - October) constitutes a period of nutritional stress for grazing animals. During the dry season alpacas grazing the high mountain zone graze perennially green pastures called bofedales. Bofedales are uncommon in the Altiplano and livestock graze dormant pastures throughout the dry season. Little is known concerning the nutritional quality of native range for alpaca. Furthermore, there are no estimates available for dietary intake of alpacas grazing rangeland on which to base stocking rate recommendations.

62 63 Alpacas in both zones suffer from impaired fiber production, low fertility, and disease attributed to inadequate nutrition.

The objectives of this study were to seasonally describe botanical selection and nutritional content of alpaca diets on typical high mountain bofedal and Altiplano grassland sites. A second objective was to estimate organic matter intake for alpacas grazing native rangeland. The Altiplano site, located at Chuquibambilla, was dominated by the grasses Festuca dolichophylla and Muhlenbergia fastigiata. The bofedal site, located at La Raya, was dominated by the sedges Eleocharis albibracteata and Carex ecuadorica with pockets of the grass Calamagrostis antoniana and the forb Plantago tubulosa. Forbs were most abundant on the Altiplano site during the late wet season (March) and during the late dry season (October) on the bofedal. High availability of grass on the Altiplano was consistent with finding mostly grass in alpaca diets. Similarly, on the bofedales where sedges and reeds were abundant, they were the major dietary component. Grass seeds were abundant in alpaca diets on the bofedal site during the dry season. Seed consumption on the Altiplano site was highest in the late wet season but dropped as the dry season progressed. 64 Crude protein (CP) in alpaca diets increased during the wet season and fell during the dry season at both sites. Year-round dietary CP averaged higher (12.3%) at the bofedal than at the Altiplano site (10.2%). Dietary CP was higher (P<0.05) on the bofedal site during the nutritionally critical late dry season. Altiplano CP values dropped to a yearly low of 6.2% during August when female alpacas entered their last trimester of gestation. Improved protein nutrition during the dry season may improve problems of low fertility common to alpacas. In vitro organic matter digestibility (IVOMD) of diets followed the same seasonal trend as CP. Contrary to CP findings, average IVOMD of alpaca diets on the Altiplano site (64%) were similar to those on the bofedal (63%). Dietary IVOMD reached its lowest values during August (49%) at the Altiplano site and in October (50%) at the bofedal. Energy intake may be deficient at both sites during these periods. Several authors suggest that alpacas consume less forage than sheep or cattle when compared on a body weight basis. My study supports this theory. Alpacas freely grazing a Festuca / Muhlenbergia range consumed daily 50.5 and 44.3 g organic matter per kg metabolic body weight during the dry and wet seasons, respectively. Average wet 65 and dry season intake for 60 kg alpacas was only 82% of the average intake reported for 47 kg sheep grazing similar range. My study suggests that the present stocking exchange ratio (0.67 alpaca per sheep) recommended by the Peruvian Ministry of Agriculture for properly stocked sheep range is low. Furthermore, using government guidelines to restock properly stocked alpaca range with sheep may result in overstocking the range. LITERATURE CITED

Antezana, C.E. 1972. Estada y tendencia de las pasturas alpaqueras en el sur oriente Peruano. Tesis. Programa Academ. de Agronomia y Zootecnia, Universidad Nacional del Cuzco, Peru.

Arnold, G.W., and M.L. Dudzinski. 1978. Ethology of free- ranging domestic animals. Elseiver Scientific Pub. Co., New York. 198p. Association of Official Agricultural Chemists. 1970. Official methods of analysis. (11th ed). Ass. Off. Agr. Chem., Washington, D.C. Bardales Ramirez, J.R. 1969. Estudio comparative de la digestibilidad de la materia seca e fibra cruda entre alpacas y ovinos jji vitro y jji vivo. Tesis. Univ. Nac. Mayor de San Marcos, Fae. de Med. Vet., Lima, Peru. 20p. Bartels, H., P. Hilpert, Ba. Klaus, Be. Klaus, R. Klaus, E: Lang, and J. Metcalfe. 1963. Respiratory functions of the blood of the yak, llama, Dybowski deer, and African elephant. Am. J. Physiol. 205:331-336. Brown, D. 1954. Methods of surveying and measuring vegetation. Bull. 42. Commonwealth Agr. Bur. Bucks, England. Bryant, F.C., and R.D. Farfan. 1984. Dry season forage selection by alpaca (Lama pacos) in southern Peru. J. Range Manage. 37(4):330-333. Condorena, N. 1980. Algunos indices de produccion de la alpaca, bajo el sistema de esquila anual establecido en La Raya. Rev. Inv. Pec. (IVITA), Univ. Nac. Mayor San Marcos. Lima, Peru. 5:50-55. Cook, C.W., and L.E. Harris. 1968. Effect of supplementation on intake and digestibility of range forages. Utah State Agric. Stat. Bull. #475. 38p. Cook, C.W., L.E. Harris, and L.A. Stodart. 1958. Measuring the nutrative content of a foraging sheep's diet under range conditions. J. Animal Sci. 7:170-180.

66 Corciova, F.J., J.D. Wallace and R. Pieper. 1978. Forage intake by grazing livestock: a review. J. Range Manage. 31(6):430-438.

Drosdoff, M., F. Queuedo, and C. Zamora. 1960. Soils of Peru. Trans. 7th Int. Congr. Soil Sci. 4:97-104. Eckerlin, R.H., and C.E. Stevens. 1973. Bicarbonate secretion by the glandular saccules of the llama stomach. Cornell Vet. 63:436-445. Escobar Calle, R. 1982. Produccion y mejoramiento de la alpaca. Banco Agrario del Peru, Lima. 334p. Esquerre C., and G. Leonor Samaniego. 1980. Peso relative del tejido del tracto alimenticio de la alpaca adulta. Rev. Inv. Pec. (IVITA). Univ. Nac. Mayor de S. Marcos, Lima, Peru. 5(l):3-9. Farfan, R.D. 1982. Dry season forage preferences of alpaca (Lama pacos) during the dry season in southern Peru. M.S. Thesis, Texas Tech Univ., Lubbock. Fernandez Baca, S.A. 1971. La alpaca, reproduction y •crianza. Univ. Nac. Mayor San Marcos. Lima, Peru. Bol. de divulgacion #7. Fernandez Baca, S.A. 1975. Alpaca raising in the high Andes. World Animal Rev. FAO, United Nations. 14:1-8. Fernandez Baca, S.A., and C. Novoa. 1966. Estudio comparative de alpacas. In: Rev. la Digestibilidad de las Forrajes en Ovinos y XTpacas. Fae. Med. Vet., Univ. Nac. Mayor de San Marcos. Lima, Peru. 18:88-96. Fierro, L.C. 1985. Forage intake, diet composition and bioenergics of grazing sheep in southern Peru. Dissertation, Texas Tech. Univ., Lubbock. Flores A., and R. Valdivia. 1973. Veloeidad de pasaje de la ingesta en alpacas y ovinos. In: IV Congreso Nac., Med. Vet. Huancayo, Peru. Gilles, J. 1980. Andean peasant economics and pastorialism. Small Ruminants CRSP, Dept. of Rural Sociology, Univ. of Miss.-Columbia. Columbia, Missouri. Pub. #1, 135p. 68 Handal, W.P., and L.R. Rittenhouse. 1975. A comparison of three methods of estimating digestibility for determining intake of grazing cattle. J. Range Manage. 28(5): 414-416.

Harris, L.E. 1970. Nutrition research techniques for domestic and wild animals. An. Science Dept. Utah State Univ., Logan. Vol. 1. Hintz, H.F., H.F. Schryzer, and M. Halbert. 1972. A note on the comparison of digestion by New World camel, sheep and ponies. Anim. Prod. 16:303-305. Hintz, H.F., C.J. Sedgewick, and H.F. Schryver. 1976. Some observations on digestion of a pelleted diet by ruminants and non-ruminants. International Zoo. Yearbook. 16:54-57. Holgado, D., R.D. Farfan, and M.E. Tapia. 1979. Evaluacion agrostologica de los pastizales de La Raya, Puno, Peru. Rev. Inv. Pec. (IVITA). Uni. Nac. Mayor San Marcos. Lima, Peru. 4:32-37. Huaman, J. 1974. Regulacion hormonal de la glicolisis y gluconeogenesis en higado de alpacas. Tesis Mag. Bioquimica. Univ. Nac. Mayor San Marcos, Lima, Peru. 33p. Huasasquiche Schwarz, A.E.G. 1974. Balance del nitrogeno y digestibilidad en alpacas y ovinos. Tesis de Bachiller. Univ. Nac. Mayor de San Marcos, Lima, Peru. 48p. Hungate, R.E., G.D. Phillips, A. McGregor, D.P. Hungate, and H.K. Buechner. 1959. Microbial fermentation in certain mammals. Science. 130:1192. Jeffery, H., and J.M. Holder. 1971. The nutritive values of some pasture species examined in a subtropcal environment. Trop. Grassl. 5:117-122. Jones, G.M. 1972. Chemical factors and their relation to feed intake regulation in ruminants: a review. Can. J. Anim. Sci. 52:207-239. Levy, E.B., and E.A. Madden. 1933. The point method of pasture analysis. New Zeal. Jour. Agr. 46:267-279. Lucio, E.Z. 1969. An investigation of the digestive enzymes of the llama. Thesis. Cornell Univ. New York. 69 McCammon-Feldman, B. 1980. A critical analysis of tropical savana forage consumption and utilization by goats. Ph.D. Diss. Univ. Illinois, Urbana. 297p. McManus, W.R. 1981. Esophageal fistulation techniques as an aid to diet evaluation of the grazing ruminant, IT\: Forage evaluation: concepts and techniques, J.L. Wheeler and R. Mochrie (eds.), CSIRO, Australia. Moir, R. 1968. Ruminant digestion and evolution. In: Handbook of Physiology. Williams and Wilkins Co., Baltimore. Chap. 126, p2673-2694. Molina, E.G., and A.V. Little. 1981. Geoecology of the Andes: the natural science basis for research planning. Mountain Res. and Devel. 1:115-144. Mooney, H.A., and W.D. Billings. 1960. The annual carbohydrate cycle of alpine plants as related to growth. Amer. J. Bot. 47:594-598. Mueller-Dombois, D., and H. Ellenberg. 1974. Aims and methods of vegetation ecology. John Wiley & Sons, New York. Novoa, C. 1970. Review, reproduction in Camelidae. J. Reprod. Fert. 22:3-20. NRC. 1975. Nutrient requirements of sheep. #5. National Academy of Sciences, Washington D.C. 72p. NRC. 1981. Nutrient requirements of goats: Angora, dairy and meat goats in temperate and tropical countries. #15. National Academy Press, Washington D.C. 91p. Orlove, B.S. 1977. Alpacas, sheep and man. Academic Press. 270p. Ortiz, S.G., J. Cavero, LL. Sillau, and S. Cueva. 1971. The parotid saliva of the alpaca. Rev. Vet. Sci. 16:54-56. Owen-Smith, N. and P. Novellie. 1982. What should a clever ungulate eat? Amer. Nat. 19:151-178. Oyanguren, F. 1969. Ensayo comparative de la digestibilidad del ensilaje de Avena sativa y Scirpus totora en ovinos y alpacas. Tesis. Univ. Nac. Tec. Altiplano, Puno, Peru. 33p. 70 Pfister, J.A. 1893. Nutrition and feeding behavior of goats and sheep grazing deciduous shrub-woodland in northeastern Brazil. Diss., Utah State Universtity, Logan. 130p.

Reiner, R.J., and F.C. Bryant. 1983. A different sort of sheep. Rangelands. 5(3):106-108. Reynafarje, C, J. Faura, A. Paredes, and D. Villavicencio. 1968. Erythrokinetics in high-altitude adapted animals (llama, alpacas, vicunas). J. Appl. Physiol. 24(l):93-97.

Riera, S.G., and A. Cardozo. 1970. Consume de ensilaje de alfalfa y agua en llamas y ovinos. Alpa Mem. 5:49-54. Rios M., A. Schlundt, and F.C. Bryant. 1984. Comportamiento de alpacas bajo cuatro intensidades de pastoreo en la sierra sur de el Peru. IT\: L.C. Fierro and R. Farfan (eds), Investigacion sobre pastes y forrajes de Texas Tech University en el Peru. Technical Article T-9-338, Col. of Agr. Sci. Texas Tech Univ., Lubbock. 116p. Rittenhouse, L.R., C.L. Streeter, and D.C. Clanton. 1971. Estimating digestible energy from digestible dry and organic matter in diets of grazing cattle. J. Range Manage. 24:73-75. Rubsamen, K., and W.V. Engelhardt. 1978. Bicarbonate secretion and solute absorption in the forestomach of the llama. Am. J. Physiol. 235:E1-E6. Salaverry, E. 1981. Clasificacion y cuantificacion de las pasturas del predio "La Raya." Ministerie de Agricultura, Peru. San Martin, M., M. Copaira, J. Zuniga, R. Rodriguez, G. Bustinza, and L. Acostas. 1968. Aspects of reproduction in the alpaca. J. Reprod. Fert. 16:395. San Martin, F., A. Huasasquiche, R. Farfan, 0. Del Valle, D. Holgado, T. Arbaiza, M. Navas and C. Villarroel. 1982. Consume y digestibilidad de pastes cultivados entre alpacas y ovinos. Resumenes de proyectos de investigacion. Univ. Nac Mayor San Marcos. Lima, Peru. p225. SAS, Institute Statistical Analysis System. 1982. SAS users guide. SAS Institute. Gary, North Carolina. 71 Schmidt, CR. 1973. Breeding seasons and notes on some aspects of reproduction in captive camelids. Int. Zoo. Yb. 13:386-390. Schmidt-Nielsen, K. 1964. Desert animals. Oxford Clarendon Press, New York. Schmidt-Nielsen, K., Schmidt-Nielsen, B., S.A. Jarnum, and T.R. Houpt. 1957. Body temperature of the camel and its relation to water economy. Am. J. Physiol. 188:102-112. Schneider, B.H., B.K. Soni, and W.E. Hamm. 1955. Methods for determining consumption and digestibility of pasture forages. Washington Agr. Exp. Sta. Tech. Bull. #16. 42p. Scott, G., and B.E. Dahl. 1980. Key to selected plant species of Texas using plant fragments. Occasional papers. The Museum. Texas Tech Univ, Lubbock. #64 37p. Sillau, H., D. Chauca, and A. Valenzuela. 1973. Evaluacion de la actividad microbiana del fluido ruminal del ovino y la alpaca. Rev. Inv. Pec. (IVITA). Univ. Nac. Mayor San Marcos, Lima, Peru. 2(1):15-21. Snedecor, G.W. 1980^ Statistical methods. Iowa State Univ. Press Ames, Iowa. 626p. Sparks, D.R., and J.C. Malechek. 1968. Estimating percentage dry-weight in diets using a microscope technique. J. Range Manage. 21:264-265. Squires, V. 1982. Dietary overlap between sheep, cattle and goats when grazing in common. J. Range Manage. 35:116-119. Steel, G.D., and J.H. Torrie. 1980. Principles and procedures of statistics. McGraw-Hill Book Co. 633p. Sumar J. 1983. Studies on reproductive pathology in alpacas. Thesis. Fae. of Vet. Med., Swedish Univ. of Ag. Sci., Uppsala, Sweden. 90p. Tapia Nunez, M.E., and J.A. Flores Ochoa. 1984. Pastoreo y pastizales de los Andes del sur de Peru. Small Ruminants Collaborative Research Program. U.S.A.I.D. Lima, Peru. 321p. 72 Taylor, C. A., and F.C. Bryant. 1977. A durable esophageal cannula for sheep and goats. J. Range Manage. 30(5):397-398.

Tillman, A. 1981. Speachless brothers, the history and care of llamas. Early Winters Press, Seattle, Wa. 120p. Torell, D.T. 1954. An esophageal fistula for animal nutrition studies. J. Anim. Sci. 13:879-884. Tovar, 0. 1973. Communidades vegetales de la reserva nacional de vicunas de Pampas Galeras, Ayacucho, Peru. Mus. Hist. Natur. Javier Prado. Lima,Peru. Sec. Bot. #27.

Vallenas, A.P., and C.E. Stevens. 1971a. Motility of the llama and guanaco stomach. Am. J. Physiol. 220(l):275-282. Vallenas, A.P., and C.E. Stevens. 1971b. Volatile fatty acid concentrations and pH of the llama and guanaco forestomach digesta. Cornell Vet. 61:(2):239-252. Vallenas, A.P., J.F. Cummings, and A. Munnell. .1971. A gross study of the compartmentalized stomach of two New-World camelids, the llama and guanaco. J. of Morph. 134(40):399-407.

Van Soest, P.J. 1970. Chemical basis for the nutritional evaluation of forages. Proc. Nat. Conf. on Forage Qual. Evsal. and'Util. Lincoln, Neb., 1969. Van Soest, P.J., and R.H. Wine. 1967. Use of detergents in the analysis of fibrous foods. IV. Determination of plant cell-wall constituents. J. Ass. Off. Agr. Chem. 50:50-55. Velarde Vilca, R., and J. Astorga. 1984. Relaciones suelo-planta in praderas alto-Andinas en el sur de el Peru. lT\: L.C. Fierro and R. Farfan (eds), Investigacion sobre pastes y forrajes de Texas Tech University en el Peru. Technical Article T-9-338, Col. of Agr. Sci. Texas Tech University, Lubbock. 116p. Walker, E.P. 1964. Mammals of the world. John Hopkins Press, Baltimore. Vol 2. 1355p. Weberbauer, A. 1936. Phytogeography of the Peruvian Andes. In: J.F. McBride (ed.) Flora of Peru. Field Mus. of Nat. Bot. Ser. Pub. #351, 13. 73 West, T.L. 1981. Alpaca production in Puno, Peru. Small Ruminant Collaborative Research Support Program. Dept. of Rural Sociology, Univ. Missouri-Columbia. Pub. #3. Whittaker, R.H. 1975. Communities and ecosystems. Macmillian Pub. Co., New York. 385p. Wilcox, B.P. 1982. Plant communities and soils of the central Andes of Peru. Thesis. Texas Tech Univ., Lubbock. 106p. Wilson, J.W. 1960. Inclined Point quadrats. New Phytologist. 59:1-8. APPENDIX A PLANT SPECIES LIST

74 75 Table 10. Plant species found on the bofedal and Altiplano sites during 1983 and 1984.

Species Site

GRASS

Bremus uniloides Altiplano Calamagrostis amoena Bofedal Calamagrostis antoniana Altiplano, Bofedal Calamagrostis rigescens Altiplano, Bofedal Calamagrostis curvula Altiplano Calamagrostis vicunarum Bofedal Distichlis humilis Altiplano Festuca dichoclada Altiplano Festuca dolichophylla Altiplano, Bofedal Festuca rigida Bofedal Hordium muticum Altiplano Muhlenbergia fastigiata Altiplano, Bofedal Muhlenbergia peruviana Bofedal Paspalum pigmaeum Altiplano, Bofedal Poa gilgiana Altiplano Poa gymnantha Altiplano, Bofedal Poa lilloi Bofedal Stipa ichu Altiplano Stipa inconspicua Altiplano, Bofedal Stipa mucronata Altiplano Stipa obtusa Bofedal

SEDGES AND REEDS Carex ecuadorica Altiplano, Bofedal Carex sp. Altiplano, Bofedal Distichia muscoides Bofedal Eleocharis albibracteata Altiplano, Bofedal Eleocharis retroflexa Bofedal Luzula peruviana Bofedal Juncus sp. Bofedal Scripus rigidus Bofedal

FORBS Alchemilla pinnata Altiplano, Bofedal Alchemilla diplophlla Bofedal Arenaria sp. Altiplano Capsell"a bursa-pastoris Altiplano Chenopo^ium pallidicaule Altiplano 76 Cotula mexicana Altiplano, Bofedal Daucus monticcDla Altiplano Hipsella reniformes Altiplano Hypochoeris stenocephala Altiplano Hypochoeris taraxacoides Altiplano, Bofedal Plantago tubulosa Altiplano, Bofedal Gentiana postrata Altiplano Gentian"ella bruneo-tincta Altiplano, Geranium sessiliflorum Altiplano Bofedal Gnaphalum sp. Altiplano Gomphrena mexicana Altiplano, Guilleninea densa Altiplano Bofedal Lepichinia sp. Altiplano Liabum ovatum Altiplano Liphidium meyeniane Altiplano Nothoscordium andicola Altiplano Nototriche sp. Altiplano Ophioglusum sp. Altiplano Oxalis sp. Altiplano, Bofedal Ranunculus sp. Altiplano Sisyrinchium sp. Altiplano Stylites andicola Bofedal Tarasa sp. Altiplano Taraxicum officinale Altiplano, Bofedal Trifolillm amabile" Altiplano, Bofedal Uritica sp. Altiplano Werneria heteroloba Altiplano, Bofedal Werneri"¥ pygmaea Bofedal APPENDIX B FORAGE AVAILABILITY AND DIET COMPOSITION DATA

77 78 Table 11. Bofedal vegetation cover and alpaca diet botanical composition on June 10 and July 10, 1983

% COVER % DIET SPECIES Jun. Jul. Jun. Jul

GRASS Calamagrostis antoniana 14 9 27 29 Calamagrostis rigescens 7 8 2 8 Calamagrostis vicunarum 3 1 T 4 Festuca dolichophylla 2 1 13 6 Muhlenbergia fastigiata 1, 1 1 1 Paspalum pygmaeum T1 0 T 0 Unknown grass T 0 T 0 SEDGES AND REEDS Distichia musccpides 12 13 6 1 Eleocharis albibracteata 11 10 7 13 Carex sp. 11 13 10 T Carex equadorica 5 4 T 2 Luzula peruviana 4 4 2 1 Scirpus rigidus 2 T T T Eleocharis retroflexa 1 0 T 0 Juncus sp. T 2 9 11 FORBS Plantago tubulosa 12 10 T 0 Hypochoeris taraxacoides 4 6 4 10 Cotula mexicana 3 2 0 1 Werneria pygmaea 1 5 2 3 Alchemilla diplophylla 1 3 2 0 Alchemilla pinnata 1 2 T 1 Taraxicum officinale 1 1 3 4 Stylites~andicola 1 1 0 T Trifolium amabile T 0 T 0 Werneria heteroloba T 0 0 0 Castilleja fissifolia T 0 T 0 Unknown forbs T T 9 T MOSS 2 2 T 5 LITTER T T SOIL T T

^ T repesents values less than 0.5%. 79 Table 12. Bofedal vegetation cover and alpaca diet botanical composition for August 22 and October 4, 1983.

% COVER % DIET SPECIES Aug. Oct. Aug. Oct

GRASS Calamagrostis rigescens 8 T^ 1 T Calamagrostis antoniana 5 6 1 6 Festuca dolichophylla 2 10 1 8 Calamagrostis vicunarum 2 T 0 1 Muhlenbergia fastigiata 1 0 0 0 Poa gymnantha 0 T T T SEDGES AND REEDS Carex sp. 16 13 0 0 Distichia musccpides 12 12 5 7 Eleocharis albibracteata 8 1 12 31 Carex equadorica 4 2 1 1 Juncus sp. 5 6 60 33 Luzula peruviana T 1 3 0 Scirpus rigidus T 0 T 0 FORBS Plantago tubulosa 13 26 0 0 Werneria pygmaea 6 5 11 2 Hypochoeris taraxacoides 4 11 T 2 Alchemilla diplophlla 3 T 1 1 Cotula mexicana 2 T 0 1 Alchemilla pinnata 1 0 0 0 Stylites andicola 1 3 T 4 Taraxicum officinale 1 0 0 0 Unknown forbs T T 1 0

MOSS 4 3 1 2 LITTER T T SOIL 1 T

^ T represents values less than 0.5% 80 Table 13 . Bofedal vegetation cover and diet botanical composition on November 7 and January 1, 1984.

% COVER % DIET SPECIES Nov. Jan. Nov. Jan

GRASS Festuca dolichophylla NA-^ 2 13 4 Calamgrostis antoniana NA 11 6 0 Poa lilloi NA T^ 3 0 Calamagrostis vicunarum NA 0 IT Calamagrostis rigescens NA 7 T 23 Poa gymnantha NA 2 0 T SEDGES AND REEDS

Juncus sp. NA 9 29 16 Eleocharis albibracteata NA 9 24 31 Distichia muscoides NA 9 5 0 Luzula peruviana NA 9 3 T Carex ecuadorica NA 6 2 13 Eleocharis retroflexa NA 1 0 T FORBS Stylites andicola NA T 9 0 Alchemilla diplophylla NA 8 3 2 Werneria pygmaea NA 0 10 Hypochoeris taraxacoides NA 10 0 6 Plantago tubulosa NA 10 TO MOSS NA 2 LITTER NA 2 SOIL NA T

^ Not available. ^ T represents values less than 0.5%. 81 Table 14. Bofedal vegetation cover and alpaca diet botanical composition on February 4 and March 3, 1984.

% COVER % DIET SPECIES Feb. Mar. Feb. Mar

GRASS Calamagrostis rigescens 10 NA^ 3 2 Calamagrostis antoniana 6 NA 1 1 Festuca dolichophylla 10 NA 14 18 Poa gymnantha 2. NA 1 1 Muhlenbergia fastigiata T2 NA 6 1 Poa qiliana" T NA 0 2 Unknown grass T NA T T SEDGES AND REEDS Eleocharis albibracteata 5 NA 24 27 Juncus sp. 9 NA 39 30 Distichia muscoides 9 NA 1 4 Carex sp. 5 NA 0 3 Luzula peruviana 2 NA 0 1 Eleocharis retroflexa 1 NA 0 0 Carex ecuadorica 1 NA T T FORBES Plantago tubulosa 13 NA 0 0 Hipochoeris taraxacoides 8 NA 2 1 Alchemilla diplophlla 2 NA 1 0 Cotula mexicana 2 NA 1 3 Werneria heteroloba 2 NA 0 0 Gentianella bruneo-tincta 1 NA 1 0 Trifolium amabile 1 NA 3 2 Oxalis sp. T NA T 2 MOSS 1 NA 2 1 LITTER T NA SOIL 0 NA

^ Not available. 2 T represents values less than 0.5% 82 Table 15. Altiplano vegetation cover and alpaca diet botanical composition on June 10 and July 10, 1983.

% COVER % DIET SPECIES Jun. Jul. Jun. Jul.

GRASS Festuca dolichophylla 21 29 16 4 Muhlenbergia fastigiata 7 20 21 14 Distichlis humilis 5 0 0 0 Poa gymnantha T"*- 0 2 6 Calamagrostis vicunarium 0 3 0 2 Calamagrostis antoniana 0 0 0 14 Hordium muticum 0 0 18 Poa Gilgiana 0 0 8 2 Stipa inconspicua 0 0 11 Calamagrostis curvula 35 0 12 T SEDGES AND REEDS Eleocharis albibracteata 0 13 3 10 Carex ecuadorica 15 7 20 2 Juncus sp. 0 4 T 2

FORBS Alchemilla pinnata 0 2 5 20 Lepidium sp. 0 3 0 0 Hypocheoris taraxacoides 0 T 0 1 Gentiana postrata 0 T 0 1 Trifolium amabile 0 T T T Werneria'heteroloba T T 2 1 Hypochoeris stenocephala 2 0 Unknown forbs T T 6 5

MOSS T 1 3 1 ALGAE 0 0 0 6 LITTER 7 9 SOIL 5 6

^ T represents values less than 0.5% 83 Table 16. Altiplano vegetation cover and alpaca diet botanical composition on August 22 and October 4, 1983.

% COVER % DIET SPECIES Aug. Oct. Aug. Oct.

GRASS Festuca dolichophylla 32 31 31 16 Muhlenbergia fastigiata 17 18 9 15 Calamagrostis antonania 16 15 10 11 Calamagrostis vicunarum 5 5 1 Poa gilgiana 0 6 0 Poa gymnanta T 3 0 0 Hordium muticum T 1 2 5 Distichlis humilis T 1 5 0 Paspulum pigmaeum T 0 T 0

SEDGES AND REEDS Carex ec^uadorica 5 8 16 22 Eleocharis albibracteata 1 3 7 17 Unknown Sedge T T 0 0

FORBS Geranium sessiliflorum 1 0 0 0 Alchemilla pinnata T T 8 11 Unknown forb T T 0 0 MOSS 1 T 1 2 LITTER 15 12 SOIL 4 1

^ T represents values less than 0.5%. 84 Table 17. Altiplano Vegetation cover and alpaca diet botanical composition on November 7 and January 1, 1984.

% COVER % :DIE T SPECIES Nov. Jan. Nov. Jan.

GRASS Festuca dolichophylla 25 22 31 19 Muhlenbergia fastigiata 18 23 8 5 Calamagrostis antoniana 22 15 11 9 Calamagrostis vicunarum 0 8 0 0 Gilgiana sp. °i 1 0 2 Hordeum muticum 1 0 1 Poa gymnanta 1 T T 4 Paspulum pigmaeum 0 T 0 0 Stipa inconspicua 0 T 0 T Calamagrostis curvula 1 0 T 0 Unknown grass T T 0 T SEDGES AND REEDS Carex ecuadorica 8 9 18 18 Carex sp. 1 1 0 0 Eleocharis albibracteata 3 T 18 25 FORBS Oxalis sp. 0 1 0 T Nototriche sp. 0 1 0 0 Trifolium amabile 0 1 0 1 Hypochoeris stenocephala 0 T 0 0 Liabum ovatum 0 T 0 0 Geranium sessiliflorum 0 T 0 0 Nothoscordium andicola 0 T 0 0 Alchemilla pinnata 1 T 10 16 Cotula mexicana 0 T 0 T Gomphrena mexicana 0 T 0 T Unknown forb T T T 2 2 MOSS 2 2 1 LITTER 11 10 SOIL 3 4

^ T represents values less than 0.5%. 85

Table 18. Altiplano vegetation cover and alpaca diet botanical composition on February 4 and March 3, 1984

% COVER % 1DIE T SPECIES Feb. Mar. Feb. Mar.

GRASS Festuca dolichophylla 40 29 10 15 Muhlenbergia fastigiata 10 18 16 Calamagrostis antoniana 4 2 4 Poa gilgiana 2 1 0 12 Hordium muticum 2 1 1 1 Poa gymnantha 1 0 1 0 Paspalom pigmaeum 1 0 0 T Stipa mucronata T 1 0 0 Distichlis humilis T 0 1 0 Calamagrostis vicunarum T 0 4 0 Festuca dichoclada 0 13 T 0 Unknown grass 0 0 0 7 SEDGES AND REEDS Carex ecuadorica 8 8 16 16 Juncus sp. 0 0 11 T Eleocharis albibracteata 3 1 16 21 FORBS Nototriche sp. 6 0 4 T Trifolium amabiie 4 2 0 T Geranium sessili^lorum 2 1 0 0 Oxalis sp. 3 1 2 T Hypochoeris stenocephala 1 2 0 0 Alchemilla pinnata 1 2 5 1 Gomphrena mexicana T 0 4 0 0 0 Arenaria sp. 1 11 unknown torbs T T 3 6 T 3 MOSS 3 7 LITTER 6 4 SOIL T T

^ T Represents values less than 0.5%. APPENDIX C CP AND IVOMD DATA

86 87 Table 19. Mean CP and IVOMD with coefficients of variation (CV) in alpaca diets.

BOFEDAL ALTIPLANO CP CV IVOMD CV CP CV IVOMD CV

Collection Peri od 1 8.7 13.9 0.58 6.4 7.1 13.7 0.62 6.9 2 8.0 11.5 0.62 5.7 9.4 10.1 0.62 7.9 3 11.0 15.6 0.56 12.2 6.2 13.1 0.49 12.5 4 12.8 12.8 0.50 8.2 9.4 13.3 0.57 6.1 5 10.4 10.7 0.65 6.0 9.2 12.9 0.64 10.4 6 15.9 8.1 0.67 6.4 13.8 9.2 0.74 8.9 7 15.3 9.7 0.75 6.7 15.3 6.6 0.75 7.6 8 16.5 6.6 0.77 3.2 11.9 8.8 0.76 2.3 APPENDIX D INTAKE STUDY DATA

88 89

Table 20. Organic matter intake and fecal output of caged and free-ranging alpacas.

Pre­ Post- X g X g Animal trial trial O.M. O.M. fecal # kg kg consumption output CV Oat Hay 1 62 59 397., 4 296.6 * 2 59 57 841.. 0 277.3 * 3 62 62 861.. 1 216.2 * 4 60 60 652.. 3 195.4 19.7 Ryegrass 5 60 57 508.. 5 241.2 * 6 46 46 516,. 8 198.3 * 7 45 45 429,. 4 97.1 * 8 51 51 725,. 0 140.5 37.3

Dry Season (free--ranging ) 1 58 * * 475.1 •k 2 69 * * 569.3 •k 3 64 * * 441.6 •k 4 62 * • 603.6 •k 5 56 * • 538.7 k 6 61 * * 506.1 11.5

Wet Season (free--ranging ) 1 58 * * 401.2 k 2 69 • • 307.1 k 3 65 * • 413.7 k 4 63 • • 408.3 k 56 * * 296.2 k 5 * 6 61 * 291.9 17.1 90 Table 21. Vegetation biomass (kg DM/ha) in a La Raya valley pasture grazed during a intake study of free-ranging alpaca.

Species Dry Season Wet Season

GRASS Festuca dolichophylla 664.0 2400.0 Festuca rigidus 314.4 0.0 Muhlenbergia fastigiata 360.0 411.0 Calamagrostis rigidus 55.2 125.6 Stipa sp. 298.4 60.8 Other Grass 28.8 18.4

SEDGES Carex sp. 19.2 115.2 Scirpus rigidus 103.2 27.2 FORBS Oxalis sp 0.0 4.8 Trifoliun amabile 0.0 2.4 Other forbs 0.1 11.2 APPENDIX E STATISTICAL TABLES

91 92 Table 22. Analysis of variance for dietary grass composition.

Source df SS F-ratio

Sites (t--1) 1 625103 85.8 Error a t(r-•1) 6 43763 Samp. a t*r([s -1) 32 247621

Month (m-1) 7 4396005 271.4 M*S (m-l)(S-l) 7 925194 57.1 Error b t(m-l)(r-l) 42 97183 Samp b t*r(m-l)s-l) 224 735421

Total t*r*m*s-l 319

Table 23. Analysis of variance for dietary sedge and reed composition.

Source df SS F-ratio

Sites (t-1) 1 734196 82.9 Error a t(r-l) 6 53217 Samp, a t*r(s-l) 32 346632

Month (m-1) 7 5170143 540.6 M*S (m-l)(S-l) 7 98(5276 103.1 Error b t(m-l)(r-l) 42 57385 Samp b t*r(m-l)s-l) 224 946221

Total t*r*m*s-l 319 93 Table 24. Analysis of variance for dietary forb composition.

Source df SS F-ratio

Sites (t-1) 1 27861 1.8 Error a t(r-l) 6 89046 Samp, a t*r(s-l) 32 492245 Month (m-1) 7 857587 4.0 M*S (m-l)(S-l) 7 596762 2.8 Error b t(m-l)(r-l) 42 1267561 Samp b t*r(m-l)s-l) 224 Total t*r*m*s-l 319

Table 25. Analysis of variance for dietary crude protein

Source df SS F-ratio

Sites (t-1) 1 395.9 102.4 Error a t(r-l) 6 21.0 Samp, a t*r(s-l) 32 101.2 Month (m-1) 7 2595.9 248.1 M*S (m-l)(S-l) 7 326.3 31.2 Error b t(m-l)(r-l) 42 62.7 Samp b t*r(m-l)s-l) 224 519.0

Total t*r*m*s-l 319 94 Table 26. Analysis of variance for dietaty IVOMD.

Source df SS F-ratio

Sites (t-1) 1 0.0122 10.1 Error a t(r-l) 6 0.0073 Samp, a t*r(s-l) 32 0.2300 Month (m-1) 7 2.2423 420.4 M*S (m-l)(S-l) 7 0.1737 32.5 Error b t(m-l)(r-l) 42 0.0320 Samp b t*r(m-l)s-l) 224 0.2451 Total t*r*m*s-l 319