1987 San Martin Tesis Comp Bot de La Dieta PDF

COMPARATIVE FORAGE SELECTIYITY AND NUTRITION OF
SOUTH AMERICAN CAMELIDS AND SHEEP

by
FELIPE A. SAN MARTIN HOWARD, B.S. in V.M., 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
December, 1987
DI
r3
ACKNOWLEDGMENTS
I express my appreciation to my major professor Dr.
Fred C. Bryant for his invaluable advice, teaching and
friendship.
I am especially grateful to committee members
Dr. Jim Pfister, Dr. Bill E. Dahl, Dr. B. Frank Craddock,

and Dr. C. Reed Richardson for their guidance and careful
reviews of this manuscript.
I also wish to acknowledge the funding for this study
which was provided by the Small Ruminant-Collaborative
Research Support Program (SR-CRSP) of the U.S. Agency for
International Development (AID).
The Universidad Nacional
Mayor de San Marcos also provided support for my graduate

program.
Special appreciation is extended to Dr. Teresa
Arbaiza, Ing. Ramiro Farfan, and Ing. Timoteo Huisa,
without them the completion of this study could not have
been possible.
I am particularly indebted to Dr. Luis Carlos Fierro
for his encouragement and support throughout my graduate
program and to Dr. Benjamin Quijandria for his logistical
support during my field work in Peru.
The personnel of
the La Raya Research Station also deserve special

recognition.
My sincere gratitude goes to Dr. Enrique

ii
Franco, Dr. Julio Sumar, Ing. Juan Alpaca, Dr. Victor

Leyva, Dr. Nicanor Condorena, and Dr. Arturo Rosales for
their support of my research effort in Peru.
I also want
to thank Ms. Gretchen Scott for her valuable assistance in

the laboratory phase of my analyses.
My sincerest appreciation goes to international
graduate students of Texas Tech University, especially
Ing. Sergio Soltero, Ing. Francis Villena, Ing. Custodio
Bohorquez, Ms. Norma Bohorquez, Mr. Changgui Wan, and Ing.
Muslim Maiga, who in different ways helped me to
accomplish my goals for my graduate program.
Special thanks also goes to Dr. Juan Espinoza Blanco
for his support and honest friendship during the last 20
years.
Lastly, I fully express my gratitude for the
encouragement and support always provided by my family,
particularly, Edita and Ricardo, Julia and Augusto,
Guillermo and Marlin, Pedro and Iris, Jorge and Gabi, Aunt
Tomasita, Paul and Zenina, Pilar and Jose Antonio,
Eduardo, and my father-in-law Victor.
111
TABLE OF CONTENTS
ACKNOWLEDGMENTS
ii
ABSTRACT
vi
LIST OF TABLES
viii
LIST OF FIGURES
xii
CHAPTER
I.
II.
INTRODUCTION AND OBJECTIVES
DIETARY SELECTION BY LLAMAS, ALPACAS, AND

SHEEP GRAZING NATIVE RANGE AND IMPROVED
PASTURES
Introduction
Description of Study Area
Methods
Results
Forage Availability
Diet Botanical Composition
Plant Part Selectivity
Similarity of Diets with
Forage Availability
Diet Similarity Among Animal Species . . .
III.
7
12
15
21
21
22
26
29
29
Discussion
31
FEED INTAKE AND DIET QUALITY IN LLAMAS,

ALPACAS, AND SHEEP GRAZING NATIVE
RANGE AND IMPROVED PASTURES
43
Introduction
Methods
Results
43
45
48
Improved P a s t u r e
F e d o Range Site
Feri Range Site
48
48
52
Discussion
55
IV
IV.
FEED DIGESTIBILITY AND PASSAGE RATES

IN LLAMAS AND SHEEP
66
Introduction
Methods
66
70
Fluid and Particulate Passage

Rate Trial
Digestibility Trials
Results
77
Liquid and Particulate Passage

Rate Trial
Discussion
77
81
87
Liquid and Particulate Passage Rates . . .

Digestibility
Intake
V.
70
74
87
89
94
Conclusion
95
SUMMARY AND CONCLUSIONS
96
LITERATURE CITED
100
APPENDIX
115
ABSTRACT
Forage selectivity, diet nutritional quality, voluntary intake, digestibility, and passage rates were studied
in South American Camelids (SAC) and sheep.
Investiga-
tions focused on (1) contrasting the seasonal dietary

botanical composition and nutrient content, and intake of
llamas, alpacas, and sheep grazing two native ranges and
an improved pasture located in the highlands of Southern
Peru; and (2) contrasting digestibility and passage rates
of different rations fed to llamas and sheep under penned
conditions.
On the improved pasture, sheep consumed 2.6 times
more legumes than SAC.
On the range sites, llamas ate
more (P < 0.05) tall grasses than alpacas or sheep.

Alpacas and sheep showed greater (P < 0.05) selection for
short grasses and forbs than llamas.
Sheep always select-
ed more (P < 0.05) leaf material than llamas and alpacas,

while alpacas selected more (P < 0.05) leaves than llamas.
Llama diets were most similar to the forage available; sheep diets were least similar.
When ranked from
most selective to least selective, the order was sheep,

alpacas, and llamas.
Alpacas were the most opportunistic.
Diet similarity between species on improved pasture was

vi
highest between llamas and alpacas.
In the dry season on
a Festuca dolicophylla range site, alpacas and sheep diets

were most similar, but on Festuca rigida range site, llama
and alpaca diets were most similar.
Llamas showed the lowest dietary quality, sheep the
greatest, whereas alpacas were intermediate. Dry matter
75
intake (g per kg BW' ) on the improved pasture and on the
range sites was 36 and 26% lower (P < 0.05) in SAC than
sheep, respectively.
Comparing llamas to sheep under penned conditions,
particulate passage rate in the forestoraach was slower in
llamas (3.5 v. 4.6%/h); total mean retention time was
longer in llamas (63 vs_. 41h) .
Liquid passage rate in the
first two compartments of the forestomach was faster in

llamas (10.3%/h) than sheep (7.7%/h).
Llama digestibility of organic matter, neutral and
acid detergent fiber, and protein was greater (P < 0.05)
than sheep.
Only when animals ate high quality rations
were the digestibility values compared (P > 0.05) between

both species.
Vll
LIST OF TABLES
1.1
Estimated population and distribution of

South American Camelids
2.1
Average botanical composition (%) of llama,

alpaca, and sheep diets during the dry and
rainy seasons on an improved pasture
23
Botanical composition (%) by plant groups

of llama, alpaca, and sheep diets during
the dry and rainy seasons on a Festuca
dolicophylla range site
24
Botanical composition by plant groups of

llama, alpaca, and sheep diets during the
dry and rainy seasons on a Festuca
rigida range site
27
Plant parts (%) in the diet of llama,

alpaca, and sheep during the dry
and rainy seasons
28
Similarity indices (%) between llama,

alpaca, and sheep diets and the forage
available on three different pastures
in the dry and rainy seasons
30
Similarity indices (%) between llama,

alpaca, and sheep diets on three
different pastures in the dry and
rainy seasons
30
Plant species important in llama, alpaca,

and sheep diets on Festuca dolicophylla
and Festuca rigida range sites during the
dry and rainy seasons
40
Nutritive composition of esophageal

samples and intake for llamas, alpacas,
and sheep during the dry and rainy seasons
on an improved pasture
49
2.2
2.3
2.4
2.5
2.6
2.7
3.1
Vlll
3.2
3.3
4.1
4.2
4.3
4.4
4.5
4.6
A.l
A.2
A.3
A.4
A.5
Nutritive composition of esophageal

samples and intake for llamas, alpacas,
and sheep during the dry and rainy seasons
on a Festuca dolicophylla range site
51
Nutritive composition of esophageal samples

and intake for llamas, alpacas, and sheep
during the dry and rainy seasons on a
Festuca rigida range site
53
Ingredients and chemical composition

of experimental rations
71
Particulate passage rate and mean

retention time from experimental llamas
and sheep
79
Rumen fluid volume and dilution rate from

experimental llamas and sheep
\
80
Digestibility coefficients (%) and intake

(g/kg W* ) of sheep and llamas as a
function of level of dry matter consumption . .
82

function of dietary quality
84

function of dietary fiber level
86
Forage availability during the dry and

rainy seasons on an improved pasture
116
Forage availability during the study seasons

on a Festuca dolicophylla range site
117

on a Festuca rigida range site
118
Botanical composition (%) of llama, alpaca,

and sheep diets during the dry and rainy
seasons on a Festuca dolicophylla
range site
119
Botanical composition (%) of llama, alpaca,

and sheep diets during the dry and rainy
seasons on a Festuca rigida range site . . . .
120
IX
A.6
A.7
A.8
A.9
A.IO
A.ll
A.12
A.13
A.14
A.15
A.16
A.17
Scientific names, family and identification

codes of plants consumed by llamas, alpacas
and sheep
121
Dry matter fecal output (g/100 kg BW) in

llamas, alpacas, and sheep during the dry
and rainy seasons on an improved pasture,
Festuca dolicophylla and Festuca rigida
range sites
122
Comparative daily dry matter intake as a

percentage of body weight in llamas and
sheep under penned conditions reported
in the literature
123
Comparative daily dry matter intake as a

percentage of body weight in alpacas and
sheep under penned conditions reported
in the literature
124
Daily intake values for llamas, alpacas,

and sheep under grazing conditions in
Southern Peru reported in the literature
Nutritive composition (%) of diets
selected by llamas, alpacas, and sheep
. . .
....
125
126
Dry matter intake (DMI) and digestible

energy intake (DEI) by llamas, alpacas,
and sheep
129
Average body weight of fecal collector

llamas, alpacas, and sheep used during
dry and rainy seasons on three pastures . . . .
132
Particulate passage rates for sheep and

llamas as affected by diet quality
133
Liquid dilution rate and rumen volume in

sheep and llamas as affected by diet
quality
134
Coefficients of digestion (%) by sheep

and llamas as affected by level of intake . . .
135

and llamas as affected by diet quality
136
. . . .
A.18
A.19

and llaraas as affected by level of neutral
detergent fiber
Organic matter intake (g/kg W* 75 ) by sheep
and llamas in Experiments 1, 2, 3, and 4
A.20
A.21
A.22
137
...
138
Comparative coefficients of digestion in

llamas, sheep, and bison
141
Coraparative coefficients of digestion

in alpaca and sheep
142
Sumraary of comparative digestibility

estiraations in alpaca and sheep
XI
(%)
145
LIST OF FIGURES
2.1
2.2
Mean,raaximumand miniraum temperatures

during January 1985 to January 1986, at
the La Raya Experiment Station, Peru
14
Distribution and amount of rainfall during

January 1985 to January 1986 (actual), and
the 10-year average for each raonth at the
La Raya Experiraent Station, Peru
16
xii
CHAPTER I
INTRODUCTION AND OBJECTIVES
The llama (Laraa glaraa), the alpaca (Laraa pacos), the
guanaco (Laraa guanicoe) and the vicuna (Laraa vicugna) forra
the group known as South Araerican Camelids (SAC).
Of
these, the tv70 former are doraesticated.

The South Araerican Caraelids belong to the faraily
Caraelidae of the order Artiodactyla (ungulates).
They are
separated frora the other rurainants into the infraorder

Tylopoda, while cattle and sheep belong to the infraorder
Pecora.
The SAC and Old World Camelids (Camelus drora-
ediarius and Caraelus bactrianus) appear to stem frora a

coraraon ancestor in North America some 16 million years
ago.
The ancestors of SAC raigrated through the Panama
Isthmus to South America with the coraing of the Ice Age.

The distribution and population of the SAC are shown
in Table 1.1.
countries:
These aniraals are mainly found in four
Peru, Bolivia, Chile and Argentina.
The
largest population of llama and alpaca are found mainly in

the Altiplano regions of Bolivia and Peru.
Alpacas are raised primarily for their wool.
There
are two distinct breeds of alpaca, the "huacaya" and the

"suri."
Huacaya, the coraraon breed, are characterized by
Table 1,1.
Estiraated population and distribution of South

American Camelids
Country
Llaraa
Alpaca
Vicuna
Peru
900.0
3,020.0
50.0
5.0
2,500.0
300.0
2.0
0.2
Argentine
75.0
0.2
2.0
109.0
Chile
85.0
0.5
1.0
13.0
Bolivia
Colurabia
0.2
USA
3.0
Source:
Novoa and Wheeler (1984)
Guanaco
highly criraped fiber which appears much like that found on

Lincoln sheep.
crimp.
Suri fiber is straight with very little
The adult body weight in huacaya and suri variety
is 62 and 64 kg, respectively (Condorena 1980).
Fiber
production in the huacaya appears to be slightly superior

to that of the suri variety.
Alpaca meat is an important
source of protein in the diet of the local people.

Llamas are tall robust aniraals. They are the largest
of the four species (120 cm tall and 110 kg). Typically,
they are used as beasts of burden.
Almost all the raeat
frora llaraa is consuraed by the people of the Altiplano.

The leather is suitable for shoes, sandals and bags.
The
fiber is long and coarse, varying in color from white to

black, and is used to make ponchos and blankets.
Peru has about 27raillionhectares of rangeland with
approximately 51% located in the Southern region.
The
rangelands of Southern Peru support about 8.6 million

sheep, 3.3 million caraelids and alraost 2 million cattle
(Martinez 1984) . Within the region, the Peruvian "Altiplano" is situated over 3,800 meters above sea level.
The
Altiplano has seasonally low teraperatures and intense

solar radiation.
Precipitation in Southern Peru is
erratic and liraited.
Approxiraately 75% of the annual
precipitation coraes between Deceraber and March, with a

relative short growing season.
During the dry season
(May-October), no precipitation occurs and little forage
production.
At these elevations, less than 5% of the land
is suitable for cultivation (Gilles 1980) , the reraaining

portion is used only by range livestock and wildlife.
The
dominant vegetation are bunchgrasses, known locally as

"ichu."
They belong to several genera including Stipa,
Festuca, and Calamagrostis.

The priraary husbandry systera is the extensive production of sraall rurainants (SAC and sheep).
At present, in
the Altiplano of Southern Peru, all llaraas, 80% of all

alpacas and 70% of total sheep are under control of
traditional pastoralists.
The reraaining alpacas and sheep
are kept in large herds which belong to the organized

types of land tenures system produced by the land reforra
program of the 1970's where pastoralism manageraent also is
practiced.
Fernandez Baca (1975) reported that approxi-
mately 200,000 campesino farailies depend in one way or

another upon alpacas.
The sraall holders raaintain flocks of 30 to 1,000
aniraals including llaraas, alpacas and sheep on coraraunal
grazing lands.
Tapia and Flores (1984) reported that the
coraposition of typical faraily herds vary according to the

altitude, land area, araong other factors.
Thus, in the
high altitude ranges near Cusco, the average composition

of a faraily herd varied from about 80 to 300 alpacas, 80
to 200 llamas, 50 to 100 sheep, and 3 to 4 horses (Tapia
and Flores 1984) .
These numbers eraphasize the multiple aniraal use of

the rangeland and the importance that llaraas have in the
small holder livestock production system.
Also, it is
iraportant to point out that, despite the Peruvian llaraa

population (Table 1.1), this species is not considered in
the livestock census when stocking rates of this region
are estiraated.
On the other hand, in recent years the
price of alpaca wool has increased on the world raarket in

relation to the value of sheep wool; thus, the size of
alpaca herds has grown.
However, increases in the alpaca
population has been conducted without a clear understanding of basic differences among these animal species.
To obtain an ecological balance araong aniraal species
and plant coraraunities,raanageraentraustbe based on knowledge of how these aniraals use and partition resources
available to thera. Unfortunately, there is a lack of
knowledge of differences in dietary partitioning araong
ruminant species sharing the Altiplano rangeland, and the
ways in which those species might react to changing
pasture conditions.
Also, inforraation is needed about how
these aniraal species, because of differences in the

gastrointestinal dynaraics, are less or raore able to raake
efficient utilization of the food resources available in
those rangelands of diverse vegetation.
The objectives of this study were:

1.
To describe and contrast the botanical coraposi-
tion of the diets of llaraas, alpacas, and sheep grazing

different pasture types.
2.
To describe and contrast the nutritive value of
diets and forage intake of llaraas, alpacas, and sheep

grazing different pasture types.
3.
To determine and contrast the digestion efficien-
cy and kinetics of SAC and sheep.
CHAPTER II
DIETARY SELECTION BY LLAMAS, ALPACAS,
AND SHEEP GRAZING NATIVE RANGE
AND IMPROVED PASTURES
Introduction
Knowledge of range herbivores' diet coraposition is
basic in understanding and planning the manageraent of
rangelands.
this type of inforraation is iraportant to
deterraine key species and optiraal forage allocation, among

other important factors related to grazing management
(Fierro 1985) .
Further, Scott and Dahl (1980) pointed out
that aniraal production frora rangelands is influenced by

plant species selected, and accurate evaluation of a
grazing aniraals diet facilities application of range
raanageraent principles.
The diets selected by grazing aniraals primarily are
dictated by animal preferences.
Moreover, aniraal prefer-
ences are the result of interactions between a pasturebased liraitation of the selection opportunity (how plant
species are distributed in both tirae and space which
influence accessibility and ease of prehension) and an
aniraal-based liraitation, or the extent to which dietary
8
preferences are raodified by the size of mouth parts and
mode of biting (Grant et al. 1985) .
Detailed data on diets of sheep indicate they prefer
leaf to stera, and green and young raaterial in preference
to dry and old (Arnold 1960, Arnold et al. 1966, Hodgson
1981) . Also, diets selected corapared with the raaterial
offered are usually higher in nitrogen and lower in fiber.
In the Altiplano region of Peru, few data exist which
focus on sheep diet selection.
Bejar (1969) , Rojas (1977)
and Fierro (1985) studied diets selected by sheep on this

treeless grassland dominated by the bunchgrasses Festuca
dolicophylla, Calamagrostic antoniana and F. dichoclada.
The vegetation complex also contained short, postrate
plants from the Cyperaceae, Juncaceae, Rosaceae and
Legurainosae farailies (Fierro 1985) . Sheep diets reported
by Bejar (1969) and Fierro (1985) were sirailar during the
dry season in terms of percent grass species in the diet.
Both studies found sheep selected Festuca dolicophylla and
Muhlenbergia fastigiata.
But the results were different
in terms of selection of grasslike species.
Among grass-
like species, Bejar (1969) reported dietary proportions of

35 and 31% for Elocharis albibracteata and Carex praegraciallis, respectively.
Whereas Fierro (1985) found sheep
diets averaged only 7% for Elocharis spp. and 16% for

Carex ecuadorica.
The initial alpaca diet studies in Peru were done by

Tapia and Lascano (1970) through direct observation.
They
observed that alpacas consuraed mainly tall grasses in the

wet season and short grasses in dry season.
The species
listed as preferred were Festuca dolicophylla, Distichia

rauscoides, Trifoliura araabile and Broraus unioloides.
Bryant and Farfan (1984) worked with adult female
alpaca and tuis (7-8 months of age) grazing on a
Festuchetura-Calaraagrosetura association.
Diet botanical
coraposition was estiraated frora June-Deceraber using fecal

material and the microhistological technique.
They found
that grass consuraption was highest during the driest

raonths (June through August); consumption of grass-like
species was inversely related to grass in the diet.
Forbs
in the diet increased in November and December (early

rainy season).
Adult alpaca diets were consistently high
in leaf material and the araount of leaf increased as the

wet season progressed.
Tuis had higher dietary levels of
grass-like plants than did adults.

more forbs than did adults.
Also, tui alpacas ate
The pattern of intake of
leaf, stera and seed by tuis alpacas was sirailar to adults.

However, tuis ate raore seeds than did adults during the
driest months of the year.
The first diet study using esophageally fistulated
alpacas was done by Barcena (1977) on FestucaCalaraagrostis range.
The raost iraportant wet season
10
dietary coraponents were Hipochoeris stinophala (18%) and
Eleocharis albibracteata (15%) .
In the dry season,
Festuca dolicophylla (56%) and Calaraagrostis vicunarura

(28%) were dorainants.
Huisa (1985) , working with alpaca on a Festuca-Stipa
range, classified the species by strata and found selection of tall grasses increased and use of short grasses
declined as the dry season progressed.
The dorainant
species in the diets were Festuca dolicophylla (21%),

Muhlenbergia peruviana (20%) , F. rigida (17%) , and Stipa
brachiphylla (15%).
Reiner and Bryant (1986) collected diets frora freeranging esophageally fistulated alpacas frora June to March
on two sites.
The bofedal site was perennially green,
tundra-like pasture, and the altiplano site (3,190m

altitude) was dorainated by Festuca dolicophylla and
As the dry season progressed
(frora June to August), bofedal alpaca diets becarae largely

sedges and reeds (78%), while altiplano alpaca diets were
raostly grass (60%).
Reiner and Bryant (1986) pointed out
that alpaca appear to be highly adaptable grazers; where

grass is available, it coraprised the bulk of the alpaca
diet.
But on a site with low grass availability and
abundant sedges, sedges dominated the diet.

Comparisons of dietary botanical composition between
alpaca and sheep has been reported only by Bryant et al.
11
(1987).
This study was done in the altiplano region of
Southern Peru on a Festuca dolicophylla range site.
The
authors reported alpaca selected more tall grasses and

less short grasses than did sheep.
Grass-like plants were
important to alpaca from October to March (early rainy

season to rainy season) (41%) , whereas sheep ate grasslike plants throughout the year (24%) . Forb consumption
reraained near 10% of the diets throughout the year for
both livestock species.
The highest index of diet sirai-
larity (70%) was during the interraediate months between

the dry and rainy season.
During the dry season, this
index was 46%; during the rainy season, 49%.

No work has been done on dietary botanical composition of the llaraa. Franklin (1982), based on visual
observation, reported that llaraas prefer to graze tall and
coarse bunchgrasses raore than other herbivores.
Further,
alpaca showed a rauch greater use of theraoistbottomlands 1

than did sheep and llamas.
In Peru, close to 70% of the total livestock are held
by small holders and sraall comraunities.
Llamas and
alpacas are mixed with the sheep and managed together.

Very little is known about diet similarities of those
species under the same grazing conditions.
This informa-
tion is important to determine forage niche separation and

feeding ecology in the Andean Region.
Furthermore,
inforraation on the dietary interrelationships between
12
sheep and South Araerican Caraelids when using different
types of pastures could illustrate differences in their
biological efficiency.
Last, range types are classified in Peru based on the
percentage of plant species preferred mostly by sheep and
alpacas.
Those preferred species were estimated from
observation and experience (Tapia and Flores 1984).

Therefore, this classification systera, although valuable,
needs to include reliable inforraation on plant preference
of llaraas.
The objective of this study was to describe and
contrast the seasonal botanical coraposition of llama,
alpaca and sheep diets under three different pastures
during two seasons.
Description of Study Area
The study area was located at the National Center for
South American Camelids Research Station at La Raya, in
the highlands of Southern Peru, departraent of Cusco, near
coordinate 1430' southern latitude and 71 western longitude.
The altitude ranges frora 4,000 to 5,000m above sea
level.
According to Holdridge (1967), La Raya is classified
as very wet subalpine life zone.
Beck (1982), taking into
account the longitudinal belts and subregions of huraidity,

classified La Raya as very cold therraic region and subhuraic subregion of huraidity.
13
The Peruvian Andes are part of the Central Andes,
which are found in Peru, Northern Chile, a great portion
of the Bolivian Sierra and Northwestern Argentina between
the Equator and the Tropic of Capricorn.
This chain of
raountains separates in Central Peru into two chains, one

which extends into Eastern Bolivia, and the other into
Northern Chile.
This division gives way to broad, high-
altitude valleys and plateaus known collectively as the

Altiplano.
Climatically, the Altiplano is characterized by
differences in diurnal temperatures.
Intense solar
radiation through a thin atmosphere produces warra afternoon teraperatures; however, loss of heat by radiation at
night reduces night teraperatures to freezing or below.
This variation in teraperatures is reflected in the
occurrence of frost which can occur on any day of the
year.
Meteorological data frora the La Raya Experiraent
Station for 1974 to 1984 demonstrated a mean maxiraura

teraperature of 14.1C and minimura temperature of -1.5C,
with an overall raean of 6.3C.
Figure 2.1 shows the raean,
raaxiraura and rainimum temperatures at La Raya Research

Station during the study period (January, 1985 to January,
1986) .
In a normal year, approximately 80% of the precipitation falls during the period of October through May,
reaching a peak intensity in January and February.
14
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15
Thus, there are two raarked seasons in rainfall distribution, the dry season (May through August) and the rainy
season (Deceraber through April).
Precipitation data frora
the La Raya Experiraent Station for 1974 to 1984 showed a

raean annual precipitation of 914.5 rara. Figure 2.2 shows
the distribution of rainfall during the study period.
Valley floor soils at La Raya have dark highly
organic epipedons.
They are low in available nitrogen and
phosphorus due to slow organic raatter decoraposition

iraposed by the cold cliraate (Molina and Little 1981).
pH is between 4.6 and 6.0.
The
Organic raatter in upper
epipedons ranges frora 4 to 8%.

Orlove (1977) pointed out that dominant vegetation
forms are bunchgrasses, known locally as "ichu."
They are
classified among several genera including Stipa, Festuca,

and Calaraaqrostis.
Each contains a nuraber of species but
the pattern of growth is sirailar, i.e., dense, deeply

rooted cluraps 5 to 20 cra in diaraeter and 15 to 60 cra in
height, and coraposed of raany single blades.
Shorter
grasses and forbs (1-5 cra) grow in favorable microsites

around the base of each clump.
Methods
Dietary botanical coraposition of llaraas, alpacas and
sheep were obtained on an iraproved, irrigated pasture and
two range sites.
The improved pasture was 5-ha of Festuca
16
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17
rubra, Loliura perenne, and Trifoliura repens.
The 5-ha
pasture was divided into six paddocks and raanaged as short

duration grazing.
Rest periods were approxiraately 60 and
40 days for dry and rainy seasons, respectively.

paddock was grazed 3 to 4 days.
Each
The pasture was irrigated
once every 10 days during the dry raonths. This pasture

was located in a flat valley floor at an elevation of
4,000ra.
The soil is a Pachic cryoboroll (heavy sandy
loara).
The two range sites were 1-ha each and were fenced as
exclosures.
These range sites were exclosed 12 months
before this study was initiated.
The Festuca dolicophylla
range site (Fedo range site) was located in a relatively

level area (< 5% slope) at 4,000m and had a constant
supply of soil water (Aguirre 1985) .
The dominant species
present at this site were Festuca dolicophylla, Muhlenbergia fastigiata, and Alchemilla pinnata.
The soil
supporting this vegetation also was classified as a Pachic

cryoboroll (light sandy loam).
Before sampling was
initiated in the rainy season, sheep accidently were

allowed to graze this range site for 3 to 4 days.
The Festuca rigida range site (Feri range site) was
located on a hillside at 4,200m with a 15 to 20% slope;
this site was rauch drier than the Fedo range site.
The
dorainant species were Festuca rigida, Stipa obstusa, and

Lepechinia raeyeni. The soil for this range site also was
18
a Pachic cryoboroll.
Both range sites have been clas-
sified according to livestock use; the Fedo range site was

classified as fair for alpacas, good for sheep, and fair
for cattle, while the Feri range site was classified as
poor for alpacas, poor for sheep and very poor for cattle
(Aquirre and Oscanoa 1985).
The vegetation of the range sites was divided into
five forage classes to enhance interpretation of results.
These were total grasses, tall grasses (greater than 40 cra
in height), short grasses (less than 40 cra in height
and/or postrate), grass-like plants, and forbs.
The iraproved pasture, Fedo range site, and Feri range
site all were sarapled for livestock diets and standing
crop during the raonths of June-July 1985 (dry season), and
Deceraber, 1985-January, 1986 (rainy season).
The biraonth-
ly raean precipitation for those periods was 9.1 and 219.7

rara, respectively (Fig. 2,2).
Diet saraples during the dry season were obtained frora
four castrated, esophageal fistulated llaraas, five
alpacas, and five sheep.
increased to five.
In the rainy season, llaraas were
The llaraas were adults (5-6 years old)
weighing an average of 89 4.9 kg.
The alpacas were
adults (4-5 years old) of the Huacaya type with an average

weight of 68 1.5 kg.
The sheep were approximately 2-3
year old Corriedale, with an average weight of 51 2.0

kg.
19
Llamas and sheep were surgically fitted with esophageal cannulas and allowed to recover for 20 and 30 days,
respectively, before the first sampling in the dry season.
Alpacas were fistulated at least two months before the
sarapling period.
animals were used.
During the rainy period, the same

The additional llama used during the
rainy season sarapling period was fistulated 2 raonths prior

to collection of diets.
The experimental aniraals in both
sarapling periods were maintained on improved pastures when

they were not involved in diet collections.
During both
collection periods, the sequence of sampling was to

collect diets from all livestock species on the improved
pasture, followed by collection from the Fedo range site,
and then the Feri range site.
Prior to the 5-day sampling period, a 10-day adaptation in each pasture was allowed to accustom the aniraals
to the experimental pastures.
Diets were collected for
0.5h using screen bags each morning at 0900h.

Upon collection, each extrusa sample was allowed to
drain, air dry, and then was hand mixed.
Each daily dried
extrusa saraple was coraposited across individual animals to

obtain one 5-day saraple per aniraal.
Saraples were ground
through a Wiley raill fitted with a 1-mra screen.
Botanical
composition was determined by the microhistological

technique (Sparks and Malechek 1968) .
Five slides were
made per sample and twenty fields read per siide, based on
20
a reference collection of plants frora the study areas.
Plant species frequency was recorded for each slide
following the procedures of Scott and Dahl (1980) . Frora
those twenty fields, five were randomly chosen to read for
occurrence of leaf, stem, seed, and flower parts.
Analy-
sis of botanical composition was conducted using a lOOX

microscope at La Raya Experiraental Station.
Forage availability (DM kg/ha) of the pastures was
assessed in each season by hand-clipping all herbaceous
vegetation within 10, Ira2 quadrats randoraly located across
each pasture before each sarapling period.
Vegetation
saraples were separated by species and oven dried at 60 C

for dry weight deterraination.
Kulczynski's sirailarity coefficient, calculated for
each pasture and season, was used to corapare diets between
llama, alpaca, and sheep, and to compare llama, alpaca,
and sheep diets with plant species availability (Grant et
al. 1985).
Kulczynski's sirailarity coefficient (S) as
modified for two sets of proportions, X and Y (e.g., X =

diet % of llama and Y = diet % of sheep), is defined by:
S=
(2W/a+b)
X 100
where W = the lesser value only coraparing X and Y for each

coraponent, suramed over all components; a = sum over all
coraponents of the X proportions = 100; and b = sura over
all coraponents of the Y proportions = 100.
For
21
interpretation, when S = 100, then there is coraplete
similarity; S = 0 implies coraplete dissirailarity.
A corapletely randomized, split plot analysis of
variance per pasture was used to test for differences
between aniraal species in diet coraposition, including
plant species, plant groups, and plant parts.
Species
(llama, alpaca, and sheep) was designated the treatment,

with individual aniraals nested within treatraents. Variation between individual animals of all species was considered the experimental error.
Sampling seasons were
considered as repeated raeasures. The GLM procedure of the

SAS coraputer prograra was used for the analysis. Mean
coraparisons were done by protected Least Significant
difference test after significant F Tests were deterrained
(Steel and Torrie 1980) .
Results
Forage Availability
Forage available on the improved pasture was slightly
lower (10% less) during the rainy season than the dry
season, but the proportion of grasses and forbs was
approximately the sarae during both seasons.
Festuca rubra
was the dorainant grass, and Trifoliura repens the dominant

forb (Table A.l).
On the Fedo range site, Festuca dolicophylla comprised 74 and 82% of the total biomass during both seasons
(Table A.2).
Muhlenbergia fastigiata in both seasons
22
was the dominant shortgrass species, followed by Bromus
unioloides and Calaraagrostis heterophylla in the dry
period.
Araong grass-like plants, Carex sp. was dorainant
in the dry season and Juncus sp. was dorainant in the rainy
season,
Araong the forbs, Alcherailla pinnata was dominant
in both seasons, followed by Lepechinia meyeni and

Trifoliura amabile in the rainy season.
On the Feri range site, Festuca rigida was the most
abundant species with 48 and 39% of the biomass in the dry
and rainy season, respectively (Table A.3).
Muhlenbergia
peruviana and Calaraagrostis heterophylla were the raost

iraportant shortgrasses species during the dry season; Poa
candaraoana was dorainant in the rainy season.
Lepechinia
raeyeni was the raost abundant forb in both seasons.

Diet Botanical Coraposition
Iraproved Pasture.
Llaraa and alpaca diets during both
periods were consistently greater (P < 0.05) than sheep in

grass coraposition (Festuca rubra and Loliura perenne)
(Table 2.1). Sheep consuraed 2.6 tiraesraoreTrifoliura
repens than did South American Camelids.
Fedo Ranqe Site.
Llaraas consuraed more (P < 0.05)
tall grasses across both seasons than did alpaca and

sheep, while tall grass consumption by alpaca was greater
(P < 0.05) than by sheep (Table 2.2). During both seasons, llamas consistently selected grasses, grass-like
23
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25
plants, and forbs at levels of 88, 6, and 6%, respectively.
Alpaca consumption of shortgrasses and forbs was
similar to that of sheep in the dry season.
However, in
the rainy season, alpaca selected less (P < 0.05) short

grasses and more (P < 0.05) forbs than did sheep (Table
2.2).
Sheep in the rainy period decreased their selection
for forbs and increased selection for shortgrasses (P <

0.05) (Table 2.2).
Grass species consumed by llamas, alpacas, and sheep
were largely Festuca dolicophylla, Calamagrostis heterophylla, Muhlenberqia fastigiata, Poa candamoana, and
Broraus unioloides (Table A.4).
Proportions of the species
in the diets were different between aniraal species.

Festuca dolicophylla was greater in llaraa diets than in
sheep diets (P < 0.05), while Muhlenbergia fastigiata was
raore iraportant to sheep (P < 0.05).
In the rainy season,
sheep showed a greater preference (P < 0.05) for Broraus

unioloides than either alpaca or llaraa (P < 0.05).
Also,
alpacas ate raore (P < 0.05) B. unioloides than llaraas. In

the case of forbs, Plantago sp. was raore preferred by
alpacas in the rainy season than llamas or sheep (P <
0.05), whereas Geranium sessiliflorura was highly preferred
by sheep in the dry season (P < 0.05) (Table A.4).
Feri Range Site.
Llaraas selected large araounts of
total grasses during both seasons, whereas alpacas

selected larger (P < 0.05) araounts of grass during the dry
26
season than in the rainy season (Table 2.3). Sheep
increased (P < 0.05) selection for grasses frora the dry
season to the rainy season by consuraing raore shortgrasses.
With respect to forbs, llaraas ate less (P < 0.05) forbs
than sheep in both seasons, and less than alpacas in the
rainy season (Table 2.3). Alpacas substantially increased
(P < 0.05) selection for forbs in the rainy season, by
decreasing consuraption of tall and short grasses.
Sheep
showed a high consuraption of forbs in both periods (Table

2.3) .
Grasses in llaraa, alpaca, and sheep diets in both
seasons were largely Festuca rigida, Stipa brachiphylla,
Calaraagrostis heterophylla, Poa candamoana, and Bromus
unioloides (Table A.5).
In the dry season, Muhlenbergia
peruviana was iraportant to all aniraal species.
In gen-
eral, all these plant species, with the exception of B.

unioloides and M. peruviana, were raore iraportant to llaraas
than alpacas and sheep.
In the case of forbs, Trifoliura
araabile was highly selected in the rainy season by all

three animal species (Table A.5).
Plant Part Selectivity
Sheep selected more leaf material (> 80%) than did
either llaraas (68%) or alpacas (73%) (P < 0.05) on all
three pastures (Table 2.4). On the Fedo range site, the
selection for leaves by all aniraals decreased from the dry
27
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29
to the rainy season; in the case of the Feri range site,
selection for leaves increased.
A possible reason for
this observed behavior could be that the Fedo range site

was grazed prior to the rainy season while the Feri range
site was not.
The selection of seeds and flower parts in
both seasons and in the three pastures was low.

Sirailarity of Diets with
Forage Availability
Of all the livestock species, llama diets were most
\
similar to the forage available while sheep diets were the

least sirailar (Table 2.5). Frora the dry to the rainy
season, the indices of sirailarities for all livestocks on
the iraproved pasture and Fedo range site increased, while
in the Feri range site these indices decreased.
Diet Sirailarity Araong Aniraal Species
The index of sirailarity between aniraal species on the
iraproved pasture was very high between llaraas and alpacas
in both seasons (Table 2.6). In the dry season, alpacas
and sheep diets were most sirailar on the Fedo range site,
but on the Feri range site, the highest index was between
llama and alpaca.
In the rainy season, from both range
sites, the index of similarities across species were lower

than the dry season.
Also, these indexes did not show
notorious differences between species.
30
Table 2.5.
Sirailarity indices (%) between llama, alpaca,

and sheep diets and the forage available on
three different pastures in the dry and rainy
seasons.
Improved
Aniraal
Festuca
dolicophylla
Range Site
Dry
Ramy
Festuca rigida
Range site
Rainy
Dry
Dry
Ramy
Llama
78
90
39
53
44
35
Alpaca
77
88
31
35
40
22
Sheep
63
63
26
27
36
20
Table 2.6
Similarity indices (%) between llama, alpaca,

and sheep diets on three different pastures in
the dry and rainy seasons.
Iraproved
Animal
Dry
Rainy
Festuca
dolicophylla Festuca ricgida
Range Site
Range site
Dry
Rainy
Dry
Rainy
Llaraa vs. Alpaca 99
94
67
59
84
51
Llaraa vs. Sheep
75
73
60
55
61
60
Alpaca vs. Sheep 76
74
83
61
70
59
31
Discussion
Forage biomass on the improved pasture was similar
between seasons.
This pasture was irrigated at 10-day
intervals during the dry raonths to allowraaintenanceof a

constant availability throughout the year.
On both range
sites, the forage production showed a sirailar trend to

data reported by Aguirre (1985) . The greater bioraass on
the Feri range site compared with Fedo range site was
presumably a reflection of the physiological adaptation of
the species to water availability.
Even though the Fedo
site is wetter, the water available to the plant is

greater on the Feri site due to less clay and organic
matter content in the soil (Aguirre 1985).
On both range
sites, the availability of grasses was above 80%, while

forbs and grass-like plants availability coraprised the
reraaining 20%.
There is no inforraation on the coraparative forage
selectivity on improved pastures of llaraa, alpaca, and
sheep.
The fact that South Araerican Caraelids selected
less legumes than sheep in this study might explain in

part the fact that cases of bloat have not been reported
for South American Caraelids (Suraar, personal communication).
Llama forage selectivity data on Altiplano region is
very liraited.
Llaraas selection of forage classes on both
range sites were in general agreeraent with observations
32
raade by Franklin (1982) who reported that llaraas prefer to
graze tall, coarse bunchgrasses.
Selection for coarse
raaterial is also reflected in selection by llaraas of more

steras and less leaves corapared wit alpacas and sheep.
Alpacas showed a high preference for shortgrasses and
forbs in both range sites.
Tapia and Lascano (1970)
reported alpacas consuraed raainly tall grasses in the rainy

season and shortgrasses during the dry season.
This trend
was observed on the Fedo range site, but the consuraption

of forbs was greater on both range sites corapared to the
results of Tapia and Lascano (1970).
On the Fedo range
site, the overall raean for forbs in alpaca diets across

both seasons was 38%. On the Feri range site, consuraption
of forbs increased frora 15 to 5 6% frora the dry to the
rainy season.
This high proportion of forb selection in
alpacas diets also was reported by Reiner and Bryant

(1986) and Bryant and Farfan (1984) . Bryant and Farfan
(1984) , working on a Festuchetura-Calaraagrosetum range
site, reported that forb consumption increased in the
early wet season and reached a peak in Deceraber (early
rainy season).
These data suggest that alpacas eat raore
forbs when cliraatic conditions favor forb growth.

Alpaca consuraption of grass-like plants on the Fedo
range site was 2% across both seasons.
Bryant et al.
(1987) reported high consuraption by alpacas of grass-like

plants frora early rainy to rainy season on a Festuca
33
dolicophylla range site.
The low consuraption of grass-
like plants by alpacas in this study on the Fedo range

site may be due to the relative availability of forbs
which were preferred by alpacas instead of grass-like
plants.
This may also be true for sheep, as sheep also
had low levels of consuraption of grass-like plants.

The intermediate position of alpacas between llamas
and sheep in selection of leaf and steraraaterialeraphasizes that alpacas are raore selective than are llaraas.
These results agree with those found by Reiner and Bryant
(1986) , Bryant and Farfan (1984) and Huisa (1985) , who
have shown that the alpaca is a highly adaptable grazer.
Alpacas vary diet selection on native range according to
forage availability.
Thus, with the exception of alpaca
diets on the Feri range site in the dry season, when forb
availability was high (Tables A.2, A.3), forbs were
iraportant in alpacas diets (Tables 2.2, 2.3).
Corapared to the findings of Bryant and Farfan (1984)
and Reiner and Bryant (198 6) , the araount of seeds consumed
by alpacas in this study was low.
Bryant and Farfan
(1984) found high seed consuraption, mainly in tui alpacas,

and indicated that seeds could compensate for the lack of
forage quality in critical months.
The low seed consump-
tion in this study might be due to the abundance of forage

found in the exclosures (Tables A.2, A.3).
Further,
Bryant and Farfan (1984) analyzed fecal samples to
34
estimate dietary botanical coraposition.
Fecal analyses
tend to underestimate fractions which are digested to such

an extent that fecal cellular residues are too small to be
detected.
On the contrary, indigestible fractions (like
seed coat) tend to be overestimated (Theurer et al. 1976,

Vavra et al. 1978) .
Sheep, like alpacas, showed greater forb intake than
llaraas. On the Feri range site in the rainy season, forb
intake by sheep wasraainlybecause of its high intake of
Trifoliura araabile, a sraall native legurae that grows only
in the rainy season.
It is not possible to conclude
whether the sheep preference for leguraes, observed on the

improved pasture and on the Feri range site, is because
sheep had more preference for this plant species or their
increased selection ability contributed legume ingestion.
The low selection for tall grasses and high selection
for leaves by sheep has been reported by others (Grant et
al. 1985, Arnold et al. 1966, Arnold and Dudzinski 1978).
Furtherraore, the high forb consuraption by sheep has been
deraonstrated in previous studies (Kothraann 1968, Bryant et
al. 1979).
This dietary selection could be due to the
height at which this species grazes. Arnold (1964)

pointed out that it is raore difficult for sheep to graze
tall, dense stands than short dense stands.
Charabers et
al. (1981) in a study coraparing sheep and cattle diets

found that the ratio of jaw movements to bites was greater
35
in sheep than cattle, which suggests that sheep manipulate
the forage with their lips and jaws before biting, to a
greater extent than do cattle.
This would allow sheep to
select the preferred coraponent better than cattle, and

perhaps caraelids, when preferred and non-preferred species
are growing together in fine-scale raixtures.
Even though differences between the raouth parts of
South Araerican Caraelids and sheep are not large, both
aniraal groups have an upper lip divided by a middle grove
(leporine lip) and the lower lip is relatively large.
However, mouth parts differ in size.
Sheep have thinner,
raobile lips while alpacas and llaraas would have thicker

lips.
Selection differences between sheep and cameiids
could be a function of the size and shape of body and

mouth parts and thus affect grazing strategy.
Several factors influence species and plant part
selection such as palatability, plant species associations, kind of aniraal, animal physiology, climate, soil
and topography, physical plant characteristic, presence of
secondary corapounds, prior experience, and their interactions (Heady 1964, Senft 1985).
Thus, the act of
selectivity, as influenced by all these factors, is very

difficult to interpret.
Even so, if selective grazing is
defined as the difference between the coraposition of the

raaterial seiected by the grazing animal and the
36
composition of the material on offer, this study showed
that sheep are more selective than are llamas.
Sheep showed a greater ability to graze all comraunities selectively under all circurastance as reflected in
the low sirailarities indices between diet and botanical
coraposition of the forage available.
The greatest indices
of sirailarity between species was found in the dry season

and could be because of the low availability of green
forage during this season.
Langlands and Sansora (1976) ,
coraparing sheep and cattle selectivity, reported the

differences between aniraal species declined as the availability of green herbage increases.
The high dietary
overlap in the dry season could be associated with

increased forage bulk in this period, offering raore
opportunities for selective grazing, which is associated
with a decline in the nutritive value of the forage on
offer (Chapter III). Also, Pfister and Malachek (1986)
reported that the highest sirailarities indexes between
goats and sheep on a deciduous woodland of Northeastern
Brazil was in the dry season, providing indirect evidence
of severe inter-species corapetition during this season.
Thus, corapetition between livestock species apparently
increased during this season.
The grazing selection of llaraas in this study could |
be interpreted as an adaptation to an arid environment.
Over 70% of world's llama population is found on the
31
Bolivian Altiplano region where the precipitation fluctuates between 250 to 450 rara. In the Peruvian Altiplano
region, with only 25% of world's llaraa population, the
precipitation fluctuates between 500 to 900rara(Tapia
1971).
Furtherraore, the majority of the Peruvian llaraa
population is found in the raore arid habitat (dry puna)

than the other South Araerican Caraelids (Novoa and Wheeler
1984, Tapia and Flores 1984).
Today, llaraasraainlyrange
frora the Central Ecuadorian highlands to Northwestern

Argentina.
Llaraas are largely concentrated within 350 kra
of Lake Titicaca, Peru, whereas the distribution center

for alpacas is within 50 to 100 kra radius of Lake Titicaca
(Franklin 1982) . Moreover, it has been observed that
llaraas are susceptible to foot rot when grazing raoist and
raarshy soil-plant types, a problera which never has been
reported in alpacas (Suraar, personal coraraunication).
Cardozo (1954) reported that llaraas prefer graze dry
ranges, whereas alpacas prefer wetter ranges.
Shiraada and Shiraada (1985) based on ethnografic,
archaezoological, physiological, and etnohistoric evidence
suggest that llamas were successfully bred and maintained
on the desert North Coast of Peru from Early Middle
Horizon (ca. A.D. 600) and perhaps since the Early Horizon.
They indicated that llamas are physiologically well
adapted for the coastal environment and the rarity of

llaraas on the Peruvian coast today is due to competition
38
with animals introduced by the Spanish.
The llaraa dietary
selection observed in this study is similar to that

reported for the desert camel.
Desert camels often
disregard the dense, succulent vegetation and raove to

seeraingly dried-out pasture, avoided by other grazing
aniraals.
In the dry raonths, relatively dry plants are
often chosen over green ones in contrast to sheep and

cattle which seek out succulent vegetation (Yagil 1985).
The high diet sirailarity between llaraa and alpaca
diets on the Feri range site, and between alpaca and sheep
diets on the Fedo range site, are attributed to the alpaca
ability to shift plant species preferences according to
In this study, llaraa and sheep
continued to graze preferred species even when the availability was low.
These results suggest that a corapleraentary grazing
systera with llaraas and sheep raay offer the most efficient
way to harvest forage and reduce inter-species competition.
This is particularly true during the dry season.
Llamas would be raore appropriately grazed on rangeiands

with taller, coarse forage.
Sheep, because of their
grazing strategy, would be able to better use plant

species in the low stratura of these range sites.
For
single-species use of these rangelands, alpacas, because

of their flexibility in diet selection and potential diet
overlap at tiraes with both llaraas and sheep, seem to be
39
the raost appropriate species to singularly graze all the
ranges in the Altiplano region.
Because of the shortcoraings of the preference index
(Loehle and Rittenhouse 1982) and because coraponents
representing a substantial proportion of the forage
available cannot be as highly ingested as rainor components
(Grant et al. 1985), no attempt was made to construct a
forage preference index.
Thus, a siraple species ranking
by their percentages in the diet on both range sites

(Table 2.7) is presented to underscore and discuss the
plants raost selected for by llaraa, alpaca, and sheep
during the dry and the rainy seasons.
Calaraagrostis heterophylla.
A perenniai grass
considered in Peru as highly desirable species (Tapia and

Flores 1984) .
Bryant and Farfan (1984) reported this
species to represent 5.9% of the dry season diets in tuis

alpacas.
This species was highly preferred by llamas in
both seasons on both range sites in this study.
Alpacas
and sheep decreased their selection for this during the

rainy season on both range sites.
Festuca dolicophylla.
A tall perennial grass spe-
cies, it is reportedly preferred by cattle, horses, sheep,

and alpacas in that order (Tapia and Flores 1984).
Its
importance in herbivore diets on the Altiplano range has

been shown in several studies (Bejar 1969, Tapia and
40
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41
Lascano 1970, Barcena 1977, Rojas 1977, Bryant and Farfan
1984, Huisa 1985, Reiner and Bryant 1986, Fierro 1985).
Its nutritive quality decreases sharply as it matures
(Kalinowsky et al. 1970).
Since this species was dorainant
on the Fedo range site, its high consuraption could be

attributed to its high availability.
Fierro (1985)
reported sheep consuraed 31% of this plant species during

the rainy season.
Broraus unioloides.
A biannual or short-lived peren-
nial grass, this species is considered a desirable species.
In this study, it was highly consumed by sheep and
alpacas in the rainy season.
Huisa (1985) reported this
species to be present in alpaca diets in the late dry and

late rainy seasons.
Tapia and Lascano (1970) indicated
this species is highly preferred by alpacas.

Stipa brachiphylla.
A perennial tallgrass considered
by Tapia and Flores (1984) to be a less desirable species.

It was highly consuraed by llaraas and was found to be an
important part of alpaca and sheep diets during the dry
season in this study.
Huisa (1985) reported a high
consumption of this species by alpacas (9 to 20%) during

the rainy season on a Festuca-stipa range site.
grass.
A perennial rhizomatous
It is considered excellent forage and grows
abundantly in the Altiplano region.
It was reported as an
important plant species in alpaca and sheep diets (Bryant
42
and Farfan 1984, Reiner and Bryant 1986, Bejar 1969, Rojas
1977, Fierro 1985).
Poa candamoana.
A perennial grass widely distributed
in the Altiplano region.
It is considered a desirable
species (Tapia and Flores 1984).
It was highly consumed
by llaraas and to a lesser degree by alpacas and sheep in

this study.
Huisa (1985) reported it in alpaca diets.
Festuca rigida.
A tall perennial grass considered to
be less desirable species (Tapia and Flores 1984).
It
becomes very coarse and fibrous as the dry season advances, thereby rendering it unpalatable.
iraportant to llamas in this study.
overall proportions of 14%.
It was very
Alpaca consumed it in
Huisa (1985) reported a
consuraption of up to 56% in the late dry season for

alpaca.
Trifoliura araabile.
A perennial legurae considered an
excellent forage, but its growth is limited to the rainy

season and on acidic soils (Tapia and Flores 1984).
In
this study, T. amabile was found in high proportion in

sheep diets on the Feri range site (28%).
CHAPTER III
FEED INTAKE AND DIET QUALITY IN LLAMAS,
ALPACAS, AND SHEEP GRAZING NATIVE
RANGE AND IMPROVED PASTURES
Introduction
Information on forage intake and diet quality is an
important tool for better formulation of range and animal
manageraent strategies.
However, there is a lack of
detailed knowledge of differences in intake and diet

quality between different animal species grazing particular rangeland comraunities, and the manner in which these
different animal species respond to changing pasture types
and conditions.
In the Altiplano region, there are few studies on
diet quality under range conditions.
Reiner and Bryant
(1986) and Huisa (1985), working with alpacas, indicated

that during the dry and late dry seasons, the diet quality
reach the lowest level of crude protein and digestibility.
Whereas in the rainy season those indicators of quality
reach the highest levels.
Sheep diet quality studies in
the Altiplano region are also scarce.
Rojas (1977) and
Fierro (1985) reported dietary quality for sheep also

resembled the seasonal variation observed in alpacas.
43
44
In Peru, there are no data for forage intake for the
three most important livestock species which were taken
simultaneously under the same pasture conditions.
Compar-
ative intake studies between llamas and alpacas (Ravillet

et al. 1985) and between alpacas and sheep (Espinoza 1975)
have been reported.
Ravillet et al. (1985) reported daily
intakes in the late dry season on a Festuca-Calamagrostis

range site of 1.9 and 1.4 kg DM/head for llaraas and
alpacas, respectively.
Also on a Festuca-Calaraagrostis
range site, Espinoza (1975) reported dry season intakes of

1.4 and 1.5 kg DM/head for alpacas and sheep, respectively.
For alpacas grazed alone, Reiner et ai. (1987)
75
reported alpacas ate 50.5 and 44.3 g OM/kg W
in the dry
and the rainy season, respectively, grazing Festuca
dolicophylla-Muhlenbergia fastigiata range site.
Farfan
et al. (1986) reported that llaraas grazing a Festuca

rigida range site had intake values of 42 and 40 g OM/kg
W'^^ for the dry and the rainy season, respectively.
For
sheep on a Festuca-Calaraagrostis range site, Fierro (1985)

75
obtained the lowest intake values (67 g OM/kg W
) in
July (driest raonth) and the highest intake (127 g OM/kg
W
) in February (wettest raonth).

Coraparative information on feed consumption between
llama and sheep (Riera and Cardozo 1968, Riera and Cardozo
1970, Caraargo and Cardozo 1971), and between alpaca and
45
sheep (Fernandez Baca and Novoa 1966, Oyanguren 1969,
Florez 1973, Huasasquiche 1974, San Martin et al. 1982a,
San Martin et al. 1982b) have been obtained under penned
conditions.
In general, the consumption per day of South
American Caraelids has been found to be lower than sheep.

The objective of this study was to estiraate and
contrast the intake and the diet quality selected by
llaraas, alpacas, and sheep grazing two different range
sites and an iraproved pasture during two seasons.
Methods
Details about the study area and design were presented in Chapter II.
Five raature castrated raale llaraas,
alpacas, and sheep were trained to wear fecal collection

canvas bags.
Preceding each collection period of 5 days,
a 20-d adaptation period was allowed to accustora the

aniraals to the experiraental pasture.
Forage DM intake (DMI) was calculated using the fecal
excretion:indigestibility ratio (Cordova et al. 1978).
The forraula used was as follows:
UtAI = fecal output (DM basis) /1- DM digestibility cf ficient.
Fecal dry matter output was determined gravimetrically by total fecal collection.
Fecal bags were
eraptied twice a day, and feces were weighed fresh and then
raixed thoroughly to take an aliquot saraple (5% of the
total fresh weight).
The aliquot sample of feces were
dried to constant weight.
46
The digestibility estimate used was the value generated frora the extrusa raaterial collected frora
esophageally-fistulated aniraals described in Chapter II.
These animals were grazed simultaneously with the fecal
collection aniraals.
Daily DMI was calculated as percentage of body weight
and relative toraetabolicweight (kg W ^ ^ ) .
These values
were used in the statistical analysis.

Diet collection technique and saraple preparation were
described in Chapter II. Dry coraposite extrusa saraple by
aniraal were analyzed for nitrogen (expressed as crude
protein) (AOAC 1975) , neutral detergent fiber, and acid
detergent fiber (NDF and ADF) (Goering and Van Soest
1970).
For the fiber analysis (NDF and ADF), decalin and
sodiura sulfite were oraitted as recoraraended by Van Soest

(1982) . Digestibility of dry raatter (DMD) was determined
using two stage iii vitro technique (Tilley and Torrie
1963).
Rumen inoculura was obtained frora rurainally-
fistulated llaraa, alpaca, and sheep grazing the same

pastures in both periods grazed by esophageally-fistulated
and fecal collector aniraals. All samples were analyzed in
duplicate and averaged.
Because they were unavailable for
those species, forage standards were not used in calculating DMD values.
Gross energy was estimated using three estrusa and
three fecal samples per species, season, and pasture using
47
a Parr adiabatic bomb calorimeter,
The fecal gross energy
for each animal was used to estimate the fecal gross

energy excreted.
The gross energy from the three estrusa
samples were averaged and used to estimate gross energy

consumed.
The digestible energy consuraed (DEI) was
obtained using the following forraula:

DEI (Mcal/day) = { (A)(B)-(C)(D) }
v/here:
A = DM intake (kg/day)
B = Average gross energy (Mcal/kg) of consuraed DM
C = DM fecal output (kg/d)
D = Gross energy (Mcal/kg) of excreted DM.
Frora the daily DEI and body weights, the DEI was calcu75
lated on araetabolicweight basis (kg W
) and this value
was used for the statistical analysis.
A corapletely randomized, split plot analyses of
variance was used to test the raain effect of aniraal
species on intake and diet quality within pasture conditions.
Species (llaraa, alpaca, and sheep) was designated
the treatraent. Variation between aniraals was considered

experiraental error.
Sarapling seasons were considered as
repeated raeasures. The GLM procedures of the SAS coraputer

prograra was used for the analyses.
Mean comparisons were
conducted using the protected Least Significant Difference

Test after significant tests were determined (Steel and
Torrie 1980) .
48
Results
Iraproved Pasture
Dietary dry matter digestibilities (DMD) were lower
(P < 0.01) in the rainy season than in the dry season for
all species (Table 3.1). DMD were similar across species
within seasons (P > 0.05).
Dietary crude protein (CP)
content was similar (P > 0.05) across species between

seasons.
Sheep diet CP levels were higher (P < 0.05) than
were values for alpacas.
Dietary neutral detergent fiber
(NDF) and acid detergent fiber (ADF) across species

increased (P < 0.01) from the dry season to the rainy
season, and were sirailar (P > 0.05) between species.
Dry raatter intake as percentage of body weight (BWI),
andraetabolicweight (MWI), and digestible energy intake
perraetabolicweight (DEI) across species, decreased (P <
0.05) frora the dry to the rainy season (Table 3.1). The
reduction of MWI was 24, 14, and 6% for llaraas, alpacas,
and sheep, respectively.
In the case of DEI, the reduc-
tion was 49, 38, and 10% for llaraas, alpacas, and sheep,
respectively.
Sheep BWI, MWI, and DEI values were greater
(P < 0.05) than those for llaraas or alpacas averaged

across seasons.
Fedo Range Site
For all livestock species, dietary dry matter digestibility increased (P < 0.01) from the dry season to
49
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50
the rainy season (Table 3.2). Alpaca and sheep diets
showed greater (P < 0.05) DMD than llaraa diets.
Content
of CP in diets were lower (P < 0.01) in the dry season

than in the rainy season.
Sheep diet CP content was
greater (P < 0.05) than CP in alpacas and llamas diets.

Alpaca dietary CP was intermediate between the CP content
of llaraa and sheep diets. NDF and ADF dietary values
across species were greater (P < 0.01) in the dry season
than in the rainy season.
The average ADF content across
season generally was lower in sheep diets than llaraa and

alpaca diets, although significant differences in ADF (P <
0.05) were found only between sheep and alpaca diets.
Dry raatter intake as percentage of body weight in
alpacas increased (P < 0.05) by 37% frora the dry season to
the rainy season, whereas BWI in llaraas and sheep was
constant in both seasons.
BWI across seasons was greater
(P < 0.05) in sheep than llaraas and alpacas. Further,

alpacas BWI was greater (P < 0.05) than llaraas in the
rainy season.
The season x species interaction for MWI was significant (P < 0.05).
In the dry season, MWI was similar (P >
0.05) between llaraas and alpacas and both species were

lower (P < 0.05) than sheep, but in the rainy season, MWI
was different between the three species.
Sheep MWI in the
rainy season was also greater (P < 0.05) than llamas and
alpacas, whereas alpacas MWI was greater (P < 0.05) than
heep
51
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52
llaraas.
Again, only alpacas showed a significant (P <
0.05) increase in MWI frora the dry season to the rainy

season (34%).
For DEI, an interaction effect (P < 0.05) was found
between seasons x species.
DEI for all the three species
increased (P < 0.05) from the dry season to the rainy

season.
The DEI increase from the dry to the rainy season
was 25, 51, and 19% for llamas, alpacas, and sheep,
respectively.
In the dry season, alpaca DEI was similar
to that in llaraas but both were lower (P < 0.05) than

sheep.
Vhereas, in the rainy season, alpacas DEI was
sirailar to that in sheep but both were greater than DEI

values for llaraas.
Feri Range Site
The season x species interaction effect for DMD was
significant (P < 0.01).
DMD was higher (P < 0.05) during
the rainy season than during the dry season (Table 3.3).
In the dry season, sheep DMD was greater (P < 0.05) than
for llaraas and alpacas, and alpacas DMD was greater (P <
0.05) than llaraas.
In the rainy season, DMD of alpacas
and sheep diets was similar and both species had greater
(P < 0.05) values than did llaraas.
Dietary CP content for all species was greater (P <
0.01) in the rainy season than for the dry season. Again,
sheep dietary CP content was greater (P < 0.05) than for
O4
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53
-p
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54
llaraas and alpacas.
Crude protein in alpaca diets was
intermediate between llama and sheep diets.

Dietary NDF values were lower (P < 0.01) in the rainy
season than in the dry season.
NDF content of sheep and
alpacas diets were sirailar (P > 0.05) and both were lower
(P < 0.05) than values for llaraas diets.
The interaction
effect between season x species for ADF dietary content

was significant (P < 0.01); only ADF in llaraas diets
decreased (P < 0.05) frora the dry to the rainy season.
In
the dry season, ADF in llaraa diets was greater than for
alpacas and sheep diets, whereas in the rainy season no
differences (P > 0.05) among species were found.
Dry matter intake as a percentage of body weight was
similar (P > 0.05) between llaraas and alpacas, and both
were lower (P < 0.05) than BMI values for sheep (Table
3.3).
Only sheep BWI values decreased (P < 0.01) from the
dry to the rainy season, and this was by a factor of 22%.

For MWI, llamas and alpacas raaintained their levels
throughout both seasons, but sheep MWI values decreased (P
< 0.05) by 21% frora the dry to the rainy season.
All
livestock species raaintained (P > 0.05) DEI through both

seasons; however, sheep showed reduction of 16% frora the
dry season to the rainy season in its energy intake.
Sheep DEI across seasons was greater (P < 0.05) than DEI
values in llamas and alpacas.
55
Discussion
With the exception of diets from the improved pasture, dietary quality for all livestock species improved
from the dry season to the rainy season.
For the improved
pasture from the dry to the rainy season, dietary DMD

decreased (P < 0.01), dietary NDF and ADF increased (P <
0.01), and dietary CP content increased slightly in all
species.
The improved pasture, managed as short duration
grazing, was irrigated once every 10 days during the dry

season toraaintainthe forage supply and a constant
stocking rate throughout the year.
However, higher
arabient teraperatures (Fig. 2.1 Chapter II) and lower

levels of solar radiation during the rainy season compared
to the dry season, could have reduced forage quality
during the rainy season.
Wilson (1981) demonstrated that
high temperatures proraote raore rapid growth with an

increase in cell wall content, less accuraulation of
soluble carbohydrates and a reduction in digestibility.
In addition, the vegetation of the altiplano region,
because of the thin atraosphere, receives intense solar
radiation.
Tapia and Flores (1984) reported solar radia-
tion in this high altitude region reaches values which are

15 to 20% higher than regions with the sarae latitude but
at sea level.
This high radiation, however, is reduced in
the rainy season because of clouds.
Thus, during the
rainy season, the shading effect could influence the
56
dietary nutritional quality of the iraproved pasture.
Deniura and Dirven (1972) reported that shading reduced
plant soluble carbohydrate levels and usually there is an
accorapanying increase in cell wall content and reduction
of digestibility, although this detriraental effect on
herbage quality under shade often does not affect the CP
level.
The dietary quality across species on the range sites
was lower (P < 0.01) in the dry season than the rainy
season.
The reduced dietary quality observed in the dry
season is because of vegetation maturity, which is accompanied by reduced digestibility, CP levels and an increase
in cell wall constituents (Cogswell and Kamstra 1976, Rao
et al. 1973, Kilcher 1981, Mitchell 1973).
When dietary quality was corapared between species in
all pastures, llaraas showed the lowest dietary quality,
sheep the highest, and alpacas were interraediate between
both species.
The higher dietary quality observed for
sheep is apparently due to its greater selection capability than alpacas and llaraas (Chapter II). Sheep showed a
high degree of selectivity for leaves, forbs and shortgrasses, which are known to be higher in nutritional
quality (Norton 1981, Ulyatt 1981).
On the contrary,
llaraas showed lower selection for these plant species and

plant parts than did sheep and alpacas.
Alpacas dietary
quality is a conformation of their intermediate
57
selectivity for leaf and steraraaterialand plant species
observed in Chapter II. The higher selectivity by sheep
which was translated to higher diet quality is, in part,
explained by the sraall size and high metabolic rate of
sheep.
The metabolic rate of small animals is higher than
for larger animals per unit of body tissue, and they

require more protein and energy per unit weight (Bell
1971, Schwartz and Ellis 1981).
Thus, small sized rura-
inants were predicted to have feeding strategies that are

raore selective corapared to large herbivores.
Factors affecting forage intake raay act through three
control raechanisms: (1) metabolic, (2) physical, and (3)
behavioral (Hodgson 1985).
Conrad et al. (1964) and
Baumgardt (1970) suggested that at digestibility levels

above 67%, the voluntary intake in ruminants eating
roughages declines at a rate that maintains a constant
intake of digestible energy.
The change-over point at
which raetabolic signals becorae raore important than physical ones would, however, raove up and down the digestibility scale with a rise or fall in the potential energy
demand of the animal.
However, results frora a wide
variety of herbage diets shows a linear increase in

voluntary intake as digestibility increases up to at least
82% (Freer 1981), which contradicts the theoretical
assuraptions of Conrad et al. (1964) and Bauragardt (1970).
58
With roughage diets norraally available to grazing
animals, the signals for intake control are likely to be
dominated by the effect of bulk in the alimentary tract.
Thus, evidence suggest that the raain liraitation to intake
appears to be related to ruraen size and the araount of
digesta that can pass daily through the alimentary tract
(Freer 1981, Van Soest 1982, Ellis 1978).
In addition,
the intake by grazing aniraals involves an additional level

of coraplexity such as the botanical and raorphological
composition of vegetation that influences ingestive
behavior, and hence, the intake of grazing animals
(Hodgson 1981) .
It is within this framework that the
results in intake obtained in this study will be discussed.

With respect to the dry matter intake across species
on the three pastures in the dry and the rainy season, the
results are quite different between the improved pasture
and the two range sites.
On the improved pasture, intake
decreased from the dry to the rainy season (P < 0.05)

because of an increase in levels of cell wall contents and
a reduced DMD during the rainy season.
Minson (1981)
reported that forage intake is affected mainly by the

level of fiber when the protein, vitarains andraineralsare
available in sufficient quantity.
Further, Van Soest
(1982) indicated that the relationship of various forage

constituents to aniraal intake ultiraately depends on their
59
association with plant structure.
Thus, the indicator of
plant structure, NDF, is the most consistent fraction

related to intake.
Therefore, the higher dietary NDF
content across species in the rainy season appears to be a

raajor contributing factor in the reduced intake observed
in the iraproved pasture during the rainy season.
Voluntary forage intake wasraaintainedat a relatively constant level on the range sites, except for
alpacas grazing the Fedo range site.
In addition, the dry
raatter fecal output (FO) (Table A.7) was greater in the

dry season than in the rainy season, also with the exception of alpacas FO on the Fedo range site. The observed
increase in FO in the dry season, coupled with the sirailar
intake between seasons (even when the dietary quality in
the dry season was lower than in the rainy season), could
be explained because aniraals during the dry season
increased their gut capacity in response of the lower
quality forages they were consuraing (Kahn and Spedding
1984, Chesson and Orskov 1984).
Although this increase in
gut capacity has been reported as an effect of a prolonged

period of low quality forage consuraption, McCollura and
Galyean (1985) reported that steers increased gut fill as
the grazing season progressed frora early to late growing
conditions.
Another factor which raay have been important to the
relatively constant intake across seasons is the high
60
plant water content during the rainy season.
Although the
addition of water per se to the rumen has little effect

upon intake because it is largely absorbed and removed
(Holmes and Lang 1963) . Van Soest (1982) pointed out that
water retention, due to the sponge effect of coarse
structural components of ingested forage can have an
inhibitory effect on intake.
On the other hand, the dietary CP levels reached
during the dry season (above 7%) probably did not affect
voluntary intake.
This level of CP supplied the minimum
nitrogen required for the microbial activity in the

forestomach of most ruminant species (Van Soest 1982,
Carapling 1966, Milford and Minson 1965).
The observed increase in alpaca intake and fecai
output in the rainy season on the Fedo range site (Tables
3.2 and A.7) was apparently due to alpacas had a greater
selection for forbs during the rainy season (42%, Chapter
II) corapared to llaraas (7%) and sheep (13%) . Forbs
usually are raore digestible, remain for a shorter time in
the forestomach, and may allow increased density of the
digestive tract, thereby allowing the aniraal to accommodate raore fill (Minson 1981, McCollura and Galyean 1985).
The significant reduction in sheep intake (P < 0.05)
in the rainy season on the Feri range site (Table 3.3)
could be a result of the influence that pasture structure
and distribution of its coraponents has on intake (Hodgson
61
1981).
The Feri range site during the rainy season
increased in the proportions of tall grasses (Table A.3)

which raight have inhibited sheep selectivity and, thus,
resulted in the observed low intake during this season.
Arnold (1964) has commented that it is more difficult for
sheep to graze a tall dense stand than a short dense
stand.
Dry raatter intake values for llaraas and alpacas, as a
percentage of body weight (BWI) on the iraproved pasture
are very sirailar to those reported for these species under
penned conditions (Tables A.8 and A.9, respectively).
San
Martin et al. (1985a), reviewing the literature on alpaca

intake obtained under penned conditions, divided the
research results into those in which dietary CP was less
than 7.5% and greater than 10.5%.
In diets with less than
7.5% CP, the average BWI was 1.4%, very sirailar to the
values obtained for alpacas grazing the range sites.
When
dietary CP levels were above 10.5%, the average BWI was

2.2%, which is close to that obtained on the iraproved
pasture.
In the case of MWI intake for sheep, NRC (1987)

75
reported values of 90.5 g/kg W*
for fine diets, a value
within the range of those observed in this study for

improved pasture.
For coarsely chopped forage, NRC (1987)

75
reported a value of 57 g/kg W'
similar to those obtained
on the range sites.
62
Alpaca and llaraa intake values are within the range
of those reported by Reiner et al. (1987) and Farfan et
al. (1986) (TableA.lO), respectively.
Unfortunately,
some intake studies (Espinoza 1975, Ravillet et al. 1985)

only reported intake per day without giving the actual
weight of the experiraental aniraals used, thus making
interpretation difficult (Cordova et al. 1978).
On the
other hand, Fierro (1985) working with sheep on a FestucaCalaraagrostis range site reported organic matter intakes
higher than the dry raatter intake values obtained in this
study.
Sheep BWI and MWI were greater (P < 0.05) than noted
for llaraas and alpacas across pastures and seasons.
The
greater intake by sheep than by SAC agrees with coraparative intakes studies under penned conditions between
llaraas and sheep (Table A.8) and between alpacas and sheep
(Table A . 9 ) .
Also, Espinoza (1975) under range conditions
reported greater voluntary intake for sheep than alpacas.

In general, alpaca and llaraa intake were sirailar (P >
0.05) across pastures and seasons.
The greater forage intake by sheep corapared to SAC is
a result of several factors such as a sraaller body size
and a relatively higher energy requireraent for sheep than
for SAC (Schneider et al. 1974, Engelhardt and Schneider
1977), which allows sheep to exhibit higher potentiai for
food consumption (Weston 1982, Blaxter et al. 1966), and
63
the smaller raouth of sheep which allows sheep to be raore
selective of plant parts (leaves) than aniraals with large
raouths (Meyer et al. 1957, Jarraan 1974).
Leaves, which
were selected raore by sheep than SAC (Chapter II), have

been shown to be eaten in greater quantities than steras,
because the leaf fraction is retained for a shorter tirae
in the forestoraach than the stera fraction (Minson 1981).
Also, a higher ruraen volurae and faster particulate passage
rate was observed in sheep corapared to SAC (Chapter IV).
These factors are highly and positively correlated with
intake (Allison 1985, Thornton and Minson 1972).
Bell (1971) has generalized that when forage quantity
is liraited, sraall body size is advantageous, and where
forage quality is liraited, large body size is advantageous.
Also, it has been suggested that sraall rurainants
raeet their relatively high raetabolic requireraent by having

sraall ruraen volurae, short turnover time, and highiy
selective diet (Van Soest 1980) . While some of these
factors raay be true in raany cases, there are sorae contradictions with respect to sheep.
Sheep, a relatively small
rurainant, have a very large ruraen volume (Hanley and

Hanley 1982) , enabling it to exploit the relative abundant
source of fermentable carbohydrates and, therefore, is an
animal well adapted to poor quality rangelands (Hanley
1982).
Thus, these sheep characteristics aiiowed them to
be a species adaptable to areas where not only is forage
64
quantity limited, but also to areas where forage quality
is limited.
On the other hand, SAC seera to be adapted to areas
where forage quantity may be limiting and the nutrients
are highly diluted by structural carbohydrates that are
difficult to digest, characteristics present in the
Altiplano region where there are long periods of a dry
season (4 months of every normal year are dry) and cyclic
years of drought are not uncoraraon.
Under these condi-
tions, SAC have adapted by reducing feed intake and

increasing the transit tirae of the digesta through the
digestive tract to better accoraraodate raicrobial attack of
their high structural carbohydrates diet.
In coraparative
terms, both the lower intake and the better digestion

capacity of the structural carbohydrates in SAC corapared
with sheep were observed in this chapter and Chapter IV,
respectively.
Besides, SAC physiologically are adapted to
the high altitudes (Renafarje et al. 1975) , and raake SAC a

raore appropriate species to utilize the sparce and fibrous
vegetation present on the rangelands of the Altiplano
region.
The energy consuraption across species followed the
sarae general trend as that observed for dry raatter intake
for seasons and pastures.
In all pastures and seasons,
sheep DEI was greater (P < 0.05) than SAC.
Engelhardt and
Schneider (1977) reported an average metabolizable energy
65
requireraent for llaraa maintenance of 61.2 kcal/kg W'
75
This value, if divided by the factor 0.82, represents the

estimated loss of DE by the aniraal in urine and raethane
(Blaxter 1964) and converts the value to digestible energy
(DE) . For SAC, this approxiraates a DE requireraent for
75 . This DE raaintenance
raaintenance of 75 kcal/kg W*
requireraent is 37% lower than that reported for sheep (119
kcal/kg W ^ ^ by NRC (1975).
These DE requireraents corapared with the DEI by the
three species indicates that on the iraproved pasture
across season, the DEI was 200% greater than that required
for maintenance by sheep and SAC. Under range conditions,
the three species grazing the Fedo range site in the rainy
season raet theirraaintenancerequireraent, but on the sarae
range site in the dry season, and on the Feri range site
in both seasons, the aniraals only consuraed about 80% of
that required to cover DE for raaintenance.
CHAPTER IV
FEED DIGESTIBILITY AND PASSAGE
RATES IN LLAMAS AND SHEEP
Introduction
The anatoray of the forestoraach systera of South
Araerican Caraelids (SAC) is quite different frora that of
advanced rurainants (Vallenas et al. 1971).
The SAC
forestomach is divided into three compartraents.
The big
ferraentation charaber is coraposed of a large corapartraent 1

(Cl) and smaller compartraent 2 (C2). Corapartraents C1-C2
are connected by a small canal with compartraent 3 (C3) , a
tubiforra organ which terrainates in the sraall hindstomach
which secretes hydrochloric acid.
Voluraes of Cl, C2, and C3 account for 83, 6 and 11%
of the total volurae, respectively (Vallenas et al. 1971).
In the adult llaraa, the contents of Cl and C2 account for
15% of the body weight, while C3 accounts for 2%
(Engelhardt and Rubsaraen 1979) . Esquerre and Saraaniego
(1980) reported that of the total weight of an alpaca
storaach, Cl, C2, and C3 coraprise two-thirds, one-twelfth,
and one-fourth of this total weight, respectively.
Two types of raucosa are found in the internal surface
of Cl and C2.
Recessed saccules are lined by a glandular

66
67
mucosa in the ventral part and the exposed surfaces are
covered by stratified squaraous epithelium in the dorsal
part.
In llamas, about half of the total surface of the

2
6700 cra raeasured was occupied by glandular raucosa
(Rubsamen cited by Engelhardt and Holler 1982) . Muci-
genous glandular raucosa are present in all corapartraents of

the forestoraach with the exception of theraucosaof the C3
distal fifth (hindstoraach) (Luciano et al. 1980).
It has been postulated that secretion by these
saccules raay provide a substantial contribution to the
buffering of digesta in Cl and C2 (Hansen and SchraidtNielsen 1957, Schraidt-Nielsen 1964, Eckerlin and Stevens
1973).
However, Rubsamen and Engelhardt (1978) indicated
that the bicarbonate secretion observed by Eckerlin and

Stevens (1973) was not observed in their experiment.
Thus, they suggest that, based on the presence of characteristic absorptive cells at the lurainal surface (Curaraings
et al. 1972), the main function of the glandular region in
the forestomach of the SAC is rapid absorption of solutes
and water.
The absorption rats reach about 2 to 3 tiraes
than those found in the rumen of sheep and goats

(Engelhardt and Rubsaraen 19 79).
Engelhardt and Rubsaraen
(1979) suggested that rapid absorption is facilitated by

periodically prolapsing of glandular saccules, thereby
changing the seraifluid contents in this region.
The rate
of absorption in C3 is significantly higher than those
68
values raeasured in the omasura of sheep and goats, after
correction for the differences in body weight.
Ferraentation rate in the SAC forestoraach is similar
to that of the advanced ruminants (Vallenas and Stevens
1971b), Engelhardt and Holler 1982, Dougherty and Vallenas
1968) . Differences inraorphologyand structure, therefore, do not appear to influence the basic function of
volatile fatty acid production and ferraentation rates.
Detailed descriptions of the forestoraachraotilityin
SAC have been reported by raany authors (Vallenas and
Stevens 1971a, Vallenas 1965, Engelhardt and Holler 1982,
Engelhardt and Rubsaraen 1979, Heller et al. 1984, Kay et
al. 1980) . SAC exhibit a raore continuous activity pattern
in the forestoraach with raore frequent cycles of ruraination
than are seen in the advanced rurainants. Kay et al.
(1980) reported that raajor contractions in SAC occur at
about 12 second intervals; brief periods of quiescence
divide the contractions into groups of about six.
This is
dissirailar to the single backward and forward moving

contractions seen at 1-minute intervals in the reticuloruraen of the advanced rurainants.
Florez (1973) studied passage rates of the digesta in
alpacas and sheep using the stained-particle retention
technique (80-5 retention) of Balch (1950)raodifiedby
Castle (1956).
He found a longer retention time in
alpacas (50.3 h) than in sheep (43.2 h ) . Clemens and
69
Stevens (1980), in a comparative study of gastrointestinal
transit tirae in ten species ofraararaals,indicated llaraas
deraonstrated raore rapid transit of fluid and sraall particles than either cattle or horses, but retained larger
particles for an extended period of tirae. On the other
hand, Heller et al. (1986) found that particulate retention time in Cl and C2 was in the same range for llamas in
advanced ruminants of sirailar weight.
Several trials of coraparative _in vivo digestibility
between alpacas and sheep ('Fernandez Baca and Novoa 1966,
Bardales 1969, Oyanguren 1969, Duran 1970, Florez 1973,
Huasasquiche 1974, Itusaca 1983, San Martin et al. 1982a,
San Martin et al. 1982b, Chayna 1983) and between llaraas
and sheep (Riera and Cardozo 1970, Caraargo and Cardozo
1977, Hintz et al. 1973, Engelhardt and Schneider 1977)
have been conducted.
Only a few of these studies observed
greater digestibility in SAC than sheep (Fernandez Baca

and Novoa 1966, Hintz et al. 1973, Caraargo and Cardozo
1971), whereas in others no differences were reported.
Differences in experiraental conditions raay account for the
inconsistency in coraparative digestibility trials. Also,
there usually has been no accounting for possibie differences in diet selectivity as hypothesized by Van Soest
(1982).
Rurainant species often exhibit different grazing
behaviors resulting in large differences in botanical

composition and nutritional values of the diets.
70
Although the structure and physiology of the digestive tract of SAC differs from advanced ruminants, there
is also a lack of consistency in existing data relating
nutritional characteristics of animal diets.
Because SAC
and sheep are sirapatric herbiovres in Peru, they corapete

for or at least share sorae of the sarae forage resources
(Chapter II). The present study was designed to evaluate
differences in digestibility between llaraas and sheep as
affected by level of intake, diet quality, and dietary
fiber content.
In addition, passage rates of the liquid
and particulate phase in the digestive tract of llaraas and

sheep were raeasured in an atterapt to relate these variables to differences in digestion between these two aniraai
species.
Methods
Fluid and Particulate Passage Rate Trial
Experiraent 1 was conducted to compare the effect of
two levels of diet quality [low quality-7% crude protein
(CP) and 2.2 Mcal digestible energy/DM kg (DE), and medium
quality-11% CP and 2.8 DE] (Table 4.1; rations I and II)
on passage rate in llaraas and sheep.
rations were pelleted.
The experiraental
The proportions of all ingredients
and nutritional coraposition are shown in Table 4.1.
Three
ruraen fistulated llaraas and three crossbred wethers of

approxiraately 2 and 2.5 years of age were used.
Weights
of sheep and llaraas were 70+ 7.39 and 145 9.81 kg,
71
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72
respectively.
refusal.
The animals were fed ad libitum up to 15%
Water was offered free choice.
Llamas and sheep
were confined toraetabolismstalls under the sarae environmental conditions and all feeding and sampling were
conducted simultaneously.
Two experiraental periods were utilized during which
aniraals were fed either ration I or ration II. The
treatments were rotated in a Latin square arrangement
within animal species, Experiraental periods consisted of
a 10-d adaptation period during which aniraals were adjusted to diets followed by a 7-d collection period. Aniraals
were fed once daily at 0800 h.
Ruraen liquid passage rate was determined on d 1 and 2
of each collection period.
Each llama and sheep received
an intrarurainal dose of 100 ral of chroraiura ethylenediarainetetraacetate (CrEDTA; Binnerts et al. 1968) , containing 411 rag of Cr.
Ruraen contents were sampled at 4, 8,
12, and 24 h after dosing, strained through two layers of

cheesecloth and frozen.
10000 X g for 15 min.
for later analyses.
Saraples then were centrifuged at
Clear fluid was decanted and saved
Atoraic absorption spectroscopy was
used to raeasure the Cr concentration in standards, fluid

saraples, and a 1:500 dilution of the original CrEDTA
solution.
Particulate passage rate was determined on d 3-7 of
the collection period imraediately after collection of
73
rumen samples for liquid passage rate.
Ytterbium (Yb)-
labeled rations were placed in the rumen of each aniraal.

Rations were labeled by the iramersion and washing procedures described by Teeter et al. (1984).
Fifty grams of
each ration was dosed in the two experiraental periods for

each aniraal.
Rectal grab saraples were collected frora each
aniraal at 0, 4, 8, 12, 16, 20, 24, 30, 36, 48, 60, 72, 82,
94, 106, 118 h post dosing.
Each saraple was placed in a
plastic bag and frozen until analysis.

Fecal saraples were prepared for analyses by drying at
60C for 48 h.
Saraples were ground through a 1-rara screen.
A total of 200 mg was extracted with 20 ml of 0.05 molar

solution of EDTA (pH 6.5 adjusted with concentrated
amraoniura hydroxide).
A 0.5 g KCL/1 EDTA solution was used
to prevent ionization.
The solution was shaken for 1 h
and filtered through Watraan #4 filter paper.
Standards
for Yb analyses were raade using fecal saraples from the 0-h
collection.
All saraples and standards were analyzed for
Yb content by atoraic absorption spectroscopy with a

nitrous oxide-acetylene flarae (Hart and Poland 1984).
Liquid dilution rate (LDR) was calculated (%/h) by
regressing the natural logarithra of Cr concentration on
tirae postdosing (0-h tirae not included).
Ruraen fluid
volume (RV) was estimated (liters) by dividing the dose by

extrapolated concentration at 0-h.
Total turnover tirae
(TTT) was estiraated (hours) by taking the inverse of LDR.
74
The liquid flow rate (LRF) was calculated (liters/hour) by
raultiplying RV tiraes LDR.
Fecal Yb excretion curves were fitted to a two
corapartment model (Grovura and Williaras 1973) which provides estiraates of particulate passage rate (%/h;
K2), and
the transit tirae (TT) of the marker in hours, which was

taken as the calculated interval before first appearance
of the marker in the feces.
The total mean retention time
(TMRT) in hours for the whole gut was obtained as:

TMRT = 1/Kl + 1/K2 + TT.
The experimental design consisted of two Latin
squares (2x2).
One Latin square was used for llamas and
one for sheep, analyzed as single experiments.
The
interaction of square x treatraents was used as the error

terra for testing aniraal species and treatment effect.
Three different experiments (2, 3, and 4) were
conducted to compare the relative digestibility of rations
fed to llaraas and sheep.
All experiraental rations were
pelleted to avoid animal selectivity.
The proportions of
all ingredients and nutritional coraposition of the experimental rations are shown in Table 4.1.
Five Rarabouillet x Suffock wethers were used in
Experiraent 2 and six were used in Experiraents 3 and 4.
Five llamas were used in all three experiments.
At the
initiation of the research, average weights and ages of
75
the sheep and llamas were 40 2.45 kg and 12 rao, and 110
14.42 kg and 18 rao, respectively.
Thus, sheep and
llamas used in this study were relatively similar in terms

of stage of growth and developraent.
In Experiraent 2, the treatraents were level of intake:
(a) ad libitura (AL), (b) 3/4 AL, and (c) 1/2 AL.
Ration
II frora Table 1 was used and contained 11% CP and 2.8 DE.
The AL intake per aniraal was estiraated for 10 days by
feeding llaraas and sheep once daily at araounts sufficient
to allow at least 15% refusal.
Thus, 3/4 AL and 1/2 AL
was calculated after AL was established.

In Experiraent 3, the treatraents were differences in
quality with three quality levels of CP and DE:
(a) low
quality (7% CP and 2.2 D E ) , (b) mediura quality (11% CP and

2.8 D E ) , and (c) high quality (15% CP and 3.2 DE).
Rations I, II and III (Table 4.1) were used and these
correspond to the low, raedium, and high quality diets,
respectively.
All animals were fed ad libitum.
In Experiraent 4, the treatraents were levels of

neutral detergent fiber (NDF) in the ration:
(a) low
(42%), (b)raediura(58%), and (c) high (68%).
Rations IV,
V, and VI (Table 4.1) were used for low,raediura,and high

NDF, respectively.
The CP level was raaintained near 11%
in all three rations.

libitura.
Again, all aniraals were fed ad
76
Water was offered free choice.
Vitamins A, D, and E
were injected at the start of the trial.
Llaraas and sheep
were confined in raetabolic stalls under the same environmental conditions and all feeding and sample collection
were conducted siraultaneously.
Within each raajor experiraent (levels of intake,
quality, and level of NDF), all animals of each species
were fed one of three sub-level treatments (AL, 3/4 AL,
1/2 AL; low, mediura, high quality; 42%, 58%, 68% NDF) in
one of three periods.
Thus, the treatm.ents were designed
in a Latin Square (3x3) arrangeraent within animal species.

Experiraental periods consisted of a 10-d adaptation
period, during which aniraals were adjusted to diets,
followed by a 7-d collection period.
once daily at 0800 h.
Animals were fed
Orts were weighed and removed
daily.
Total collection of feces for the final 7-d of each
experiraental period was facilitated by harnesses and
collection bags, which were eraptied daily.
Daily aliquots
of feces were frozen, later dried at 60C for 48 H, ground

through a 1-rara screen, and composited for analyses.
Feed
and feces were analyzed for dry and organic matter (OM)
content, nitrogen (expresses as crude protein) (AOAC
1975) , acid detergent fiber (ADF) (Goering and Van Soest
1970), and NDF.
NDF was estimated using the procedures of
77
Robertson and Van Soest (1977), which includes the use of
alfa amylase to eliminate residual starch frora the saraple.
The experiraental design consisted of two Latin
squares (3x3), one with llaraas and one with sheep anaiyzed
as a single experiraent.
Aniraals within species was the
error terra for testing aniraal species effects.
The GLM
procedures of the SAS (1982) coraputer prograra was used for

analyses.
Mean coraparisons were by Least Significant
Difference Test (protected) (Steel and Torrie 1980) .

Results
Liquid and Particulate Passage
Rates Trial
Experiraent 1:
Rates.
Effect of Diet Quality on Passage
Corapartraental analysis as used in this study
assuraes that the pool of ingesta in the larger corapartraents of the gut flows frora each individual corapartment in
the same proportion as the concentration of the marker.
In Grovum and Williaras' (1973) raodel, Kl is the value
assigned to raaterial passing through the larger and slower
raoving corapartraent.
This represents the reticulo-ruraen
particulate passage.
The value for K2 represents
particulate passage to the lower tract of a ruminant.
The
latter value has been validated by comparing excretion

values of markers introduced in both the abomasum and the
rumen (Grovura and Williaras 1973).
78
Organic raatter intake was lower for llamas than for
sheep (P < 0.05) (Table 4.2); however, intake by sheep was
lower than values observed in Experiraents 2, 3, and 4.
There is no explanation for this observed low intake, but
it was not a result of a liraiting nutrient.
Consuraption
by llaraas was sirailar to values observed in the previous

experiraents.
Particulate passage rate in the rumen (Kl) was less
(P < 0.05) for ilamas than for sheep (Table 4.2). Particulate passage rates were lower (P < 0.05) on the low
quality ration than for the mediura quality ration (3.60
vs. 4.48 % / h ) , respectively for both llamas and sheep.
Particulate passage rates through the cecum and colon (K2)
was nearly two times as fast in sheep corapared to llamas,
even though differences were not significant (P > 0.10).
The first appearance of the marker (TT) was earlier (P <
0.05) for sheep than for llamas in both rations.
Total
raean retention tirae (TMRT) also was less in sheep than in

llaraas aithough differences were not significant (P >
0.10) .
Liquid dilution rate was faster in llamas than sheep,
but the difference was not significant (P > 0.10) (Tables
4.3, A.15).
Rumen fluid volurae was sirailar between
species and treatraents, but when based on metabolic weight

75
(per kgW' ) , sheep showed 40% greater (P < 0.10) RV than
llaraas.
Even
though the raeans of LFR and TTT were quite
ro
79
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81
different (lower LFR and higher TTT in llamas than in
sheep), there were no significant differences (P > 0.10)
betv/een species.
Even though digestibility trials under penned conditions limit aniraal selectivity, trials comparing alpacas
and sheep have shown that sheep are raore selective (San
Martin et al. 1982b).
Van Soest (1982) pointed out the
feed refused by selective aniraals usually consisted of

raore lignified parts of the diet.
If this factor is not
quantified and corrected for, it may erroneously lead to

apparently higher digestion coefficients in the animal
practicing greater selection.
On the other hand, diges-
tion trials are coraraonly conducted at restricted intake to

avoid animal variation.
However, this standardization
process has reduced the value of extant digestion data, as

possible inherent differences araong species have been
raasked (Van Soest 1980) .
Thus, all the aniraals in this
study were fed ad, libitura levels, resting pelleted rations.

Experiraent 2;
ibility.
Effect of Level of Intake on Digest-
Intake and digestion coefficients for this trial
are shown in Tables 4.4, A.16, and A.19.
the OM intake
(g/kg w"^^) for the AL treatraent was greater (P < 0.01)

for sheep than for llaraas.
DE consuraed by AL sheep v;as
2.8 tiraes that required for raaintenance (NRC 1975).
Llama
82
(0
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83
DE consumption was 2.4 times that required for maintenance
(Engelhardt and Schneider 1977).
In treatraent 1/2 AL,
both species consuraed enough DE toraeetraaintenance

requireraents by 1.4 and 1.3 tiraes for sheep and llamas,
respectively.
CP intake in AL treatment was 1.9 tiraes the
raaintenance requirement for sheep (Preston 1966), and 1.26

for that required in llamas (Huasasquiche 1974) .
In the
1/2 AL treatraent, CP intake was only 0.96 and 0.63 tiraes

the araount required by sheep and llaraas, respectively.
The organic raatter and NDF digestibilities were
\
greater (P < 0.01) in llaraas than sheep.
No differences
(P < 0.05) between treatraents were found for ADF digestibility.

Experiraent 3;
ibility.
Effect of Diet Quality on DigestOrganic raatter intake (g/kg W* 75 ) was greater (P
< 0.01) for sheep than for llaraas.
Differences between
treatraents (P < 0.05) also were observed (Tables 4.5,

A.19).
Consuraption increased in both species frora the low
quality toraediuraquality diet, and then declined when fed

the high quality ration.
There was an interaction (P <
0.05) between species x treatraents for OM and NDF digestibilities (Tables 4.5, A.17).
The digestion coefficients
for OM and NDF were greater (P < 0.05) for liaraas than for
sheep for the low andraediuraquality treatraents, but were
sirailar (P > 0.05) between species on the high quality
diet.
ADF digestibility was greater (P < 0.05) in llamas
co
84
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(A
 0.10), but different between treatments (P <

0.01); digestibility coefficients were greater for both
llaraas and sheep on the high quality ration.
Experiraent 4;
bility.
Effect of Fiber Content on Digesti-
Although rations were forraulated to supply the
sarae CP level at 11%, the actual CP content was 13, 11 and

12% for low,raediuraand high fiber content rations,
respectively.
The digestion coefficients and intake are
shown in Tables 4.6, A.18, and A.19.

75
OM intake (g/kg W
) was greater for sheep than for
llaraas (P < 0.01).
No differences (P > 0.05) were observ-
ed in intake between treatraents, although both species

showed a slight reduction in intake when fed the high
fiber content ration.
Differences between species for OM
digestibility (P < 0.05), NDF digestibility (P < 0.01),

ADF digestibility (P < 0.05), and CP digestibility (P <
0.05) were observed; llaraas had greater digestion coef- ,
ficients than did sheep in all cases.
Across aniraal
species, differences were observed in OM digestibility (P

< 0.01), NDF digestibility (P < 0.05), and CP digestibility (P < 0.01) between treatraents; digestibility of NDF
was lower when aniraals were fed the lower fiber ration.
co
86
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87
Discussion
Liquid and Particulate Passage Rates
The longer retention time of the particle matter in
llaraas (62.3 h) corapared to sheep (40.9 h) agrees with
results of Florez (1973) who observed a longer retention
tirae in alpacas than in sheep.
Clemens and Stevens (1980)
also found a longer retention tirae of particulate raatter

in llaraas corapared to that in cattle and horses,
The liquid passage rate between treatraents (low and
raediura quality rations) in llaraas and sheep were sirailar.
This raay be related to the fact that animals of both
species had the same intake on both treatments.
Level of
intake has been positively related to change in fluid

turnover tirae (Galyean et al. 1979, Evans 1981).
The
faster LDR, LFR, and lower TTT across treatraents in llamas

than sheep agree with the results reported by Clemens and
Stevens (1980) , who corapared the gastrointestinal transit
tirae in ten species of mararaals.
Llaraas demonstrated more
rapid transit of fluid and small particles than either

cattle or horses.
Maloy (1972) reported a higher fluid
outflow frora the forestoraach in caraels than in zebu

steers.
No data are available on RV estimation for SAC.
Esquerre and Samaniego (1980) reported that the relative
weight of the wet SAC forestomach tissues corapared with
those of sheep and cattle are similar.
The RV obtained in
88
sheep are in the range of those given by Puiser and Moir
(1966), but lower than those reported by Hanley and Hanley
(1982).
The low sheep intake values (Table 4.2) and
pelleted rations used in this experiraent raay explain, in

part, the relatively low RV found in sheep (Teeter and
Owens 1983, Merchen et al. 1986).
The greater RV/kg W ^ ^
observed in sheep than llaraa would be related to the fact

that sheep had greater intake than llaraa (Van Soest 1982).
The raore rapid LDR in llaraas than in sheep may be a
result of higher rates of salivary flow and a higher ratio
of salivary flow to forestoraach size (Ortiz 1971).
Owens
and Isaacson (1977) indicated that prime deterrainants of

forestoraach fluid dilution rate appeared to be fluid or
salivary input.
Harrison et al. (1975) reported an
iraproved efficiency of raicrobial protein synthesis of up

to 25% as a result of increased LDR because of the application of artificial saliva.
The relatively shorter
retention of rurainal fluid in llaraas than in sheep allows

llaraas to iraprove the microbial growth present in C1-C2
compartraents, assuring that arainirauraaraount of energy is
available to maintain the microbial population (Isaacson
et al. 1975, Hespel and Bryant 1979, Orskov 1982).
The reasons for differences in retention time of
liquid and particle raatter in the forestoraach of ruminants
is not well understood.
It is assuraed that in llamas and
sheep, the function of the canal between C2 and C3
89
(reticulura and oraasum) and the raotility of C^ (reticulura)
are raainly responsible for this selective transport of
liquid and particles.
Heller et al. (1984) pointed out
that both species have coraparable pressure-suction mechanisms at the oraasal canal.
Coraparative studies have shown
that raorphology (Vallenas et al. (1971) and histology

(Engelhardt and HoIIer 1982) of the forestoraach differ
raarkedly between the two species, and further that the
basic pattern of raotility of the storaach compartraent
appears to be quite different (Vallenas and Stevens 1971a,
Heller et al. 1984, Kay et al. 1980).
Thus, differences
in passage rates between these species is a reflection of

the striking differences in their digestive anatomy and
physiology.
Digestibility
It has been shown that increases in feed level
usually result in reduced digestibility coefficients
(Schneider and Flatt 1975).
This effect was not observed
in Experiraent 2 where the effect of level of intake on

digestibility in Ilaraas and sheep was raeasured. On the
contrary, digestibility of ADF was lower (P < 0.05) for
both species on the 1/2 AL intake than on the 3/4 AL and
AL intakes.
Further, values for digestibility of OM and
NDF were the sarae on 1/2 AL corapared to AL, but values for
1/2 AL were lower than 3/4 AL.
This may be due partially
to lower CP intake corapared to energy intake on the 1/2 AL
90
treatraent.
On the 1/2 AL treatraent, CP intake was 96 and
63% of raaintenance requirements for sheep and llamas,

respectively.
Thus, the ratio of nitrogen to energy raay
have been so low that raicrobial activity was depressed and

resulting in a reduction in digestibility values.
Other
factors, such as the deficiency of other specific nutrients (trace rainerals, etc.) required by raicrobes, raay also
have affected the low digestibility values in the 1/2 AL
treatraent (Orskov 19 82).
The greater NDF digestibility by liaraas than by sheep
on the 1/2 AL treatraent (27% vs. 39%) may be a result of
higher concentrations of NH3 in Ilaraas in compartment Cl
and C2 compared to sheep (Engelhardt and Schneider 1977,
Hinderer and Engelhardt 1975).
Engeihardt and Heller
(1985) indicated that caraelids recycle and utilize body

urea for raicrobial protein synthesis extreraely efficiently
on low protein diets.
This would provide Ilaraas with raore
nitrogen available for protein synthesis, thus iraproving

digestibility.
When different quality rations were fed to sheep and
Ilaraas (Experiraent 3 ) , Ilaraas digested the low and medium
quality feedstuffs better than sheep.
When a high quality
ration was fed, both species had sirailar digestibilities.

This agrees with results reported by San Martin et al.
(1985b) who indicated greater coefficients of digestion
for dry raatter, crude fiber, and CP in alpacas than in
91
sheep when the feed offered was of lower quality (less
than 7.5% of C P ) . No differences were found between
species when the feed offered was above this threshold
level of CP.
These results support the conclusions of
Engelhardt and Heller (1985) that caraelids show a greater

adaptability to extrerae dietary conditions than do the
advanced rurainants.
When different levels of NDF were fed to sheep and
llaraas, the digestion coefficients for OM, NDF, ADF, and
CP were greater in llaraas than in sheep at all levels of
dietary fiber.
Even the low NDF level produced this
effect, and the NDF level in this ration raay not have been
low enough at 42% to corapensate for the low digestibility
in the reticulo-ruraen of sheep by compensatory postrurainal digestion (Schneider and Flatt 1975, Van Soest
1982) .
The observed lower digestibility coefficients of NDF
and ADF when both sheep and Ilaraas were fed low fiber
rations agrees with results reported by several authors
(Chapell and Fontenot 1968, Chou and Walker 1964, El
Shazly et al. 1961, Slyter et al. 1971, Slyter et al.
1970).
Low fiber feedstuffs have raore starch and soluble
carbohydrates thus raore energy (Table 4.1). When digestion occurs in the rumen, there is an initial rapid
fermentation that may produce sufficient acid to cause a
decline in rumen pH, causing a delay in fiber digestion.
92
This drop in fiber digestion is due to changes in number
and kinds of rumen microbes, with a predominance of
araylolytic bacteria over cellulolytics, thus inhibiting
cellulolitic enzyrae synthesis (Leatherwood 1973).
In general, the greater digestibilities for OM, NDF,
and ADF in Ilaraas than sheep in all experiraents, with the
exception of the high quality ration in Experiment 2,
agree with the results obtained in Ilaraas and sheep by
Hintz et al. (1973), Caraargo and Cardozo (1970) (Table
A.20), and in alpacas and sheep by Fernandez Baca and
Novoa (1966) (Table A.21).
However, differences found
here are not as large as obtained by Fernandez Baca and

Novoa's (1966) study, who reported that alpacas were about
50% more efficient in the digestion of dry matter and
crude fiber than were sheep when they were given oat hay
containing 25% crude fiber (Table A.22).
Hintz et al. (1973) suggested that the greater
efficiency of digestion of the SAC raay be accoraplished
because of their raore frequent forestoraach contractions
and ruraination cycles which provide for raore efficient
raaceration,raixing,and absorption.
AIso, Heller et al.
(1984) indicated that the greater digestive efficiency of

SAC than the advanced ruminants may be due to more frequent contractions per motility cycle in the forestomach
of SAC, allowing maxiraum mixing of food with bicarbonate
buffer and with microorganisras v/ithout increasing the
93
rate of forestomach eraptying.
Thus, the greater digest-
ibility in Ilaraas than sheep observed in this study is

related to the longer retention time of the digesta
associated with the peculiar raorphology, histology and
raotility of the forestoraach in Ilaraas.
A longer retention
of particles in the forestoraach of Ilaraas than sheep

enables Ilamas an extensive fermentation, particularly for
potentially digestible, fibrous feed.
Blaxter (1963) states that maximal digestion occurs
only if the passage of food is delayed so that the food is
exposed to those sites where raicrobial actions takes
place.
In this study, the digesta retention time of both
low andraediuraquality rations in C1-C2 of Ilamas was 29 h

(1/Kl), whereas the retention time in the reticulo-rumen
of sheep was 22 h (1/Kl).
In the case of cecura-colon
retention time (1/K2), the values were 11 and 6 h in

Ilamas and sheep, respectively.
Because of these longer
retention tiraes for Ilaraas, digestion is greater for

Ilaraas.
This is especially true when aniraals eat low
quality rations.
Blaxter (1963) pointed out that when
rate of passage is increased, there is an apparent depression in the digestibility of poorer quality foods than for
those of high quality.
Digestibility of feed of high
quality is relatively unaffected by retention time in the

reticulo-rumen, and it also undergoes much compensatory
94
degradation post-ruminally (Schneider and Flatt 1975, Van
Soest 1982).
On the other hand, several studies (Tables A.20,
A.2I, A.22) have reported no differences in the digestion
coefficients between SAC and sheep.
These discrepancies
could be attributed to selectivity factors which have not

been taken into account in those experiments.
ntake
The level of OM intake by sheep frora all three
experiments in this study agrees with other published data
(NRC 1987).
The OM intake of llamas, which was lower (P <
0.05) than for sheep in every experiment, also agrees with

several coraparative studies between llaraas and sheep
(Table A . 8 ) .
This lower intake in Ilamas than in sheep is
due in part to lower basal metabolisra (Schneider et al.

1974) and energy requireraent for raaintenance (Engelhardt
and Schneider 1977) than for sheep.
Weston (1982) pointed
out that intake differences between species are clearly

related to the aniraals'raetabolisraactivity.
Therefore,
aniraals with higher basal raetabolisra (as is the case of

sheep with respect to Ilaraas) will exhibit high potential
for feed consuraption.
In addition, severai studies have
negatively and positively related the retention time and

rumen volume with intake, respectively (Thornton and
Minson 1972, Campling 1970, Ailison 1985).
Thus, the
lower intake in llamas than sheep may aiso be explained by
95
longer retention time of digesta in Ilamas, and the
smaller Ilama ruraen volume in terms of metabolic weight
than in sheep (Experiraent 1 ) .
Conclusion
Because of the lower energy requireraent and ruraen
75
volurae, per kg W' , the slower transit tirae for particle
raatter and the faster liquid transit tirae of the digesta
in Ilaraas than in sheep, it is evident that behavioral and
nutritional strategies differ between these species.
Apparently Ilaraas, with a lower voluntary intake, coupled
with a greater digestive efficiency, can subsist on poorer
quality food which requires a longer retention tirae in the
digestive tract to extract theraaxiraumavailable energy
from it.
Furtherraore, the ferraentation process in the
Ilaraa is enhanced through the reduction of raicrobial

raaintenance requireraent because of higher liquid turnover
rates.
Sheep, with a greater daily intake rate, appear to
have less capacity to extract energy frora fiber than do
Ilaraas and suggests that this species would use different
nutritional strategies to balance quality of food intake
and digestive efficiency (Van Soest 1980) .
This greater
intake in sheep requires a raore rapid passage rate and

consequent reduction in digestibility.
CHAPTER V
SUMMARY AND CONCLUSIONS
The rangelands of Southern Peru support large populations of sheep and South Araerican Camelids (SAC).
Studies
focused on raultiple aniraal use of these rangelands' need

to be eraphasized to obtain an ecological balance araong
plant coraraunities and aniraal species.
The objectives of this study were to seasonally
describe and contrast dietary botanical selection, dietary
nutrient content, and intake of Ilamas, alpacas, and sheep
grazing an improved pasture and two range sites.
A second
objective was to determine and contrast the digestion

efficiency and kinetics of SAC and sheep.
On the iraproved pasture, sheep consuraed 2.6 tiraes
raore leguraes than SAC.
On the range sites, Ilaraas select-
ed greater proportions of tall grasses than alpacas and

sheep.
Alpacas and sheep showed a high selection for
short grasses and forbs.
With respect to plant parts,
sheep consuraed raore leaves than did alpacas and Ilaraas.

Alpacas had an intermediate selection, between Ilamas and
sheep, for leaf and stera raaterial, emphasizing the fact
that alpacas were more selective grazers than the llaraas.
96
97
The indices of sirailarity between species were high
between llaraas and alpacas, but lower between SAC and
sheep on iraproved pasture.
On the range sites, high diet
sirailarity indices were found in the dry season.
On the
Fedo range site, sheep and alpacas showed the highest

similarity index, whereas on Feri range site, Ilama and
alpaca had the highest index.
These changes of sirailarity
indices araong species are attributed to alpaca's adaptability to shift plant species preferences according to
Thus, in a corapleraentary grazing
system, llamas and sheep offer the most efficient way for
harvesting the forage available, while alpacas seera to be
adequate for single-species use of these rangelands.
Nutritional quality in the diets of the three species
across pastures and seasons are a confirraation of the
aniraal species capability to select.
Llaraas showed the
lowest dietary quality, sheep the highest, while alpacas

were interraediate between both species.
The dietary
quality across species on the improved pasture was lower

in the rainy than in the dry season.
On the range sites,
the lowest dietary quality values were obtained in the dry

season.
This study confirmed the greater consuraption in sheep
than in SAC reported in previous studies.
Under penned
conditions (digestibility trials), the average consuraption

. 75
by sheep in terras of metabolic weight (kg W* ) was 36%
98
greater than SAC.
The same difference was found when
comparisons were made under grazing conditions on an

improved pasture.
On the range sites under grazing
condition, the average sheep consumption was also greater

in sheep than in SAC by a factor of 26%.
Considering adult body weights for sheep, Ilaraas and
alpacas of 40, 65, and 108 kg, which for coraparative
purposes, corresponds to 15.9, 22.9, and 33.5 kg of
raetabolic weight, respectively, alpacas and Ilaraas raetabolic weights are 1.4 and 2.1 tiraes that of sheep (1.0).
Further, a Ilaraa'sraetabolicweight is 1.5 tiraes that of
alpacas.
Thus, considering a 30% lower intake in SAC than
in sheep, the conversion factor between alpacas and Ilaraas

with sheep would be 1.0 for alpacas (1.4 x 0.7) and 1.5
for llamas (2.1 x 0.7), respectively.
Those ratios,
alpaca:sheep and Ilamarsheep, are quite different frora

stocking exchange ratios reported in the literature which,
without a differentiation between alpacas and Ilaraas,
indicate ratios frora 1.5 to 1.8 per sheep unit, respectively.
Finally, it is important to keep in raind that
these ratios do not include herbage and environmental

characteristics.
Thus, they can only be used as a tool in
the determination of stocking rates under range conditions.

Greater digestibilities of organic matter, neutral
detergent fiber, and acid detergent fiber were obtained in
99
Ilamas compared to sheep.
On the other hand, when high
quality ration was offered, no differences were found

between both species.
This finding reinforces the theory
that SAC are species well adapted to raake a better utilization of poor quality food than sheep.
The better
utilization of this poor quality food in SAC is possible

because the longer retention tirae of the particulate
matter in their digestive tract than sheep, favoring a
prolonged period of exposure to the action of microbial
population located in compartraents 1 and 2 of the SAC
forestoraach.
South Araerican Caraelids' dietary selection characteristics, low intake, and high efficiency of the utilization of structural carbohydrates, besides their
physiological adaptation to the high altitudes, make these
species raore appropriate to utilize the sparce and fibrous
vegetation present in the high altitude range of the
Altiplano region.
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110
Pfister, J. A., and J. C. Malechek. 1986. Dietary
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APPENDIX
115
116
Table A.l.
Forage availability during the dry and rainy

seasons on an iraproved pasture.
Plant Group
and Species
Grasses
Dry Season
(June-July)
DM kg/ha (%)
Rainy Season
(Deceraber-January)
DM kg/ha (%)
1663 (94)
1520 (95)
Festuca rubra
1203 (68)
912 (57)
Loliura perenne
460 (26)
608 (38)
106 ( 6 )
80 ( 5)
Trifoliura repens
88 ( 5 )
80 ( 5)
AlcheraiIIa pinnata
18 ( 1 )
__
Forbs
TOTAL
1770 (100)
1600 (100)
117
Table A.2.

on a Festuca dolicophylla range site.
Plant Group
and Species
Dry Season
(June-July)
DM kg/ha (%)
Rainy Season
(December-January)
DM kg/ha (%)
Total Grasses
2958 (87)
2046 (93)
Tall Grasses
2516 (74)
1914 (87)
2516 (74)
*
1804 (82)
88 ( 4)
Festuca dolichopylla
Festuca ortophylla
Stipa Brachiphylla
22 ( 1)
Short Grasses
442 (13)
132 ( 6)
Muhlerabergia fastigiata
Hordeura
rauticura
Poa candaraoana
Broraus unioloides
Pasalura pigraaeun
Calaraagrostis heterophylla
Grass-Like Plant
Eleocharis albibracteata
Luzula peruviana
Juncus sp.
Carex sp.
204 ( 6 )
T
T
170 ( 5 )
*
68 ( 2)
340 (10)
T
T
*
306 ( 9)
102 ( 3)
66 ( 3)
T
T
22 ( 1)
22 ( 1)
T
44 ( 2)
T
T
44 ( 2)
110 ( 5)
*
*
T
T
Forbs
Hypericura caespitosura
Oenothera s p .
AlcheraiIIa p i n n a t a
Lepechinea raeyeni
Gnaphaliura s p .
Peperonia sp.
Oxalis sp.
Notothriche sp.
T r i f o l i u r a araabile
Hipsella sp.
Geraniura sessiliflorura
Teraxacura officionale
TOTAL
68^(
* 2)
*
*
*
*
*
*
22
44 (( 1)
2)
T
T
T
T
44 ( 2)
T
*
T
T
3400 (100)
T Represents values less than 0.5%

* Species were not found.
2200 (100)
118
Table A.3.

on a Festuca rigida range site.
Plant Group
and Species
Dry Season
(June-July)
DM kg/ha (%)
Rainy Season
(Deceraber-January)
DM kg/ha (%)
Total-Grasses
3230 (85)
3128 (92)
Tall Grasses
2584 (68)
2788 (82)
1824
23
456
76
190
(48)
(.6)
(12)
( 2)
( 5)
1326 (39)
*
1292 (38)
646 ( 5)
340 (10)
Festuca rigida
Festuca dolichopylla
Stipa obstusa
Stipa ichu
Stipa brachiphylla
Short Grasses
Muhlerabergia peruviana
Muhlerabergia fastigiata
Vulpia
raegalura
P a s p a l u r a pigraaeura
Broraus u n i o l o i d e s
Poa candaraoana
Forbs
Peperonia sp.
Lepechinea
raeyeni
Liabura ovatura
Alcherailla pinnata
Plantago
raonticola
Notothriche sp.
Gnaphaliura sp.
Hipsella sp.
Oenothera sp.
Oxalis sp
Trifoliura araabile
Astraqalus garbancillo
Bidens andicola
Hipochoeris taraxacoides
Cardioneraea raraosisima
TOTAL
170 ( 5)
190 ( 5)
*
418 (11)
23 ( . 6 )
T
~
T
*
306 ( 9)
570 (15)
272 ( 8)
T
102 ( 3)
T
T
31 (.9)
T
T
T
T
"^
T
68 ( 2)
T
456 (12)
*
27 (.7)
T
*
*
76 ( 2)
*
T
*
T
T
T
23 (.6)
T
T
3800 (100)
T Represents values less than 0.5%

* Species were not found.
24
(.7)
*
3400 (100)
119
Table A.4.
B o t a n i c a l c o m p o s i t i o n (%) of Ilaraa, a l p a c a ,
and sheep d i e t s d u r i n g t h e dry and r a i n y
s e a s o n s on a F e s t u c a d o l i c o p h y l l a range s i t e
Plant Group
Dry^ Season
and Species Llaraa Alpaca Sheep
Overall Mean
Rainy Season
?^paca Sheep
Llaraa Alpaca Sheep Llaraa .
1
Grass
89^
Fedo^
Feri
Feor
Stic
Stob
Mufa
Poca
Brun
Cahe
Mupe
Honu
26^
4^
3''
4^
2^
9
9
3^'
29
l^
62^
61'=
14d
1?
..ab
1
10
6
2^
19
l^
î
.3^
15
5
l^
20
3^
87^
56^
86^
88
59
74
45
28^
20^
23
1
.2
.5
17
.5
.1
.5
11
6
4^
20
11
3,
ll^
3
21
5
29^
6
lO^
4^
18^
5^
36
2
2
2
1
lO^
8^
4
24^
.5
1
ll^
.5
.5
^^b
13^
2
3
2^
l^
~b
6
Grass-Like
Plant
6^
Elal
Jusp
Lupe
2^
2
3^
..bc
^bc
4^
2
l^
.3
3
â
2
2
1
.1
1
.5
Forbs
4"=
38
24
Alpi
PIsp
Gese
Rasp
Hita
Gnsp
Leme
Tram
Oesp
8
14
8
2
6
,a
1
3
2
2
.4
.2
âb
35^
.7
.1^
>
35^
-^bc
6
3
..bc
ll^
2
JDC
3^
l'^
42^
13^
8
21^
3^
3
3
5
l^
l^
.3
4
,a
1
2
1
.a
.4
1
.5
.5
-a
.3
2^
3
â
2
2
2
.2
1
"Sleans with the sarae superscripts on row are not significantly

-different (P < 0.05).
Scientific narae and faraily name are reported in Table A.6.
120
Table A.5.
B o t a n i c a l c o m p o s i t i o n (%) of I l a m a , a l p a c a ,
and s h e e p d i e t s d u r i n g t h e d r y and r a i n y
s e a s o n s on a F e s t u c a r i g i d a r a n g e s i t e .
Plant Group
D]cy Season
and Species Llaraa Alpaca Sheep
Rainy Season
Overall Mean
Llaraa Alpaca Sheep Llaraa Alpaca Sheep
1
Grass
87^
83^
54^
Feri
Fedo
Feor
Stic
Stob
Stbr
Cahe
Mufa
Poca
Brun
Mupe
Papi
18
17
2^
9
O^
.1"
.1^
.1^
^b
O^
3.
is''
19^
24^
23^
.bc
i^
2^
15^
.1^
O
13,^
13^
1
84^
44^
65^
15
10
2
20^
23^
2
14^
8^
1?
27^
.5
gb
0,
13^
12^
O^
86
64
16^
.05
.05
5
14^
.1
.05
.2
l^
12
16
2^
3
8
8
.05
?^
18
23
â
10
4
4
.5
60
7^
13
12
2^
4
14
8
Grass-Like
Plant
2"
l^
.5^
.5
.2
Lupe
2^
l^
.5^
.5
.2
12<^
15^
16-^
58^
35"
14
36
40
2^
6^
2
2
.5
3
1
2
.5
.5
4
2
2
4
.5
4
.5
4
2
6
8
4
3
4
6
4
4
1
.5
.1
Forbs
AIpi
PIsp
Gese
Hita
Leme
Gnsp
Rasp
Hyca
Oxsp
Oesp
Trara
3
..bc
3
.,bc
45
6^
6^
l^
i^
8
7^
8^
3
2^
3:
3
..bc
"b
^b
%
0
8=
,-ab
5
0=
6^
_a
7
â
5
- -a
15^
l''
..bc
Ib
3
7b
>
.2
0
28^
"Sfeans with the same superscripts on row are not s i g n i f i c a n t l y

d i f f e r e n t (P < 0.05) .
S c i e n t i f i c name and faraily narae are reported in Table A.6.
15
121
Table A.6.
Scientific naraes, faraily and identification

codes of plants consuraed by Ilaraas, alpacas
and sheep.
Code
Species
Faraily
Fedo
Festuca dolicophylla
Festuca rigida
Graraineae
Feri
Feor
II
Stic
Stob
Festuca orthophylla
Stipa ichu
S. obstusa
Stbr
S. brachiphylla
II
Cahe
Muhleraberqia peruviana
Muhlemberqia fastigiata
Poa candamoana
II
Mupe
Mufa
Poca
Papi
Brun
Horau
Elal
Lupe
Jusp
AIpi
PIsp
Gese
Gnsp
Hita
Lerae
Rasp
Paspalura pigraae
Broraus unioloides
Hordeura rauticura
Eleocharis albibracteata
Luzula peruviana
Juncus sp.
AlcheraiIIa pinnata
II
II
II
II
II
II
II
II
II
Cyperaceae
Juncaceae
II
Rosacea
Plantiganacea
Geraniacea
Corapositae
Plantago sp.
Gnaphaliura sp.
Hipochoeris taraxacoides
Lepechinea raeyeni
Libiatae
Trara
Ranunculus sp.
Trifoliura araabile
Ranunculaceae
Legurainosae
Oesp
Oenothera sp.
Oxsp
Oxalis sp.
Oenotheraceae
Oxalidaceae
Hyca
II
Hypericaceae
04
Q)
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132
Table A.13.
Average body weight of fecal collector

Ilamas, alpacas, and sheep used during
dry and rainy seasons on three pastures.
Body Weight Metabolic Weight
Species
Pasture
Season
(kg)
(kg w"^^)
Llama
Improved
Dry
Rainy
84.9
85.8
28..0
27.
Alpaca
Dry
Rainy
68.6
65.8
23..8
23..1
Sheep
Dry
Rainy
48.3
51.3
18..3
19..2
Dry
Rainy
85.8
97.5
27.,2
31..0
Alpaca
Dry
Rainy
68.8
70.5
23..9
24..3
Sheep
Dry
Rainy
50.2
54.0
18..8
19..9
Dry
Rainy
86.7
91.9
28..4
29..7
Alpaca
Dry
Rainy
68.9
68.2
23..9
23..7
Sheep
Dry
Rainy
52.1
52.7
19,.4
19,.5
Llama
Llama
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Table A.19.
75
Organic matter intake (g/kg W ' ^ ) by sheep
and Ilamas in Expe:riments 1,, 2, 3, and 4.
Experiment
2
Animal
Sheep
Llamas
Ad libi-tum
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1
2
3
4
5
Means
SD
CV, %
93.48
113.70
148.42
98.97
95.45
110.00
22.89
20.81
57.24
62.76
72.30
82.86
50.78
65.19
12.64
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1
2
3
4
5
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CV, %
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86.27
116.31
80.44
77.39
87.11
16.85
19.34
41.70
45.51
50.31
66.13
41.86
49.10
10.14
20.66
1/2 AL
1
2
3
4
5
Means
SD
CV, %
46.69
56.82
74.14
50.71
50.24
55.72
10.92
19.60
28.60
31.37
34.93
44.18
29.34
33.68
6.36
18.88
Low QuaLlity
1
2
3
4
5
6
Means
SD
CV, %
104.46
107.79
99.10
80.75
86.33
90.14
94.76
10.69
11.28
49.54
57.41
61.58
61.94
44.38
TreatmeiQt
54.97
7.74
14.09
139
Table A.19.
(cont.)
Experiment
3
Treatment
Medium Quality
High Quality
High Fiber
Medium Fiber
Sheep
Llamas
1
2
3
4
5
6
Means
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CV, %
106.00
100.75
101.95
104.69
100.29
106.33
103.34
2.68
2.59
49.63
64.83
70.28
47.65
61.65
1
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3
4
5
6
Means
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CV, %
72.20
72.48
84.79
89.12
74.48
78.35
78.57
6.99
8.90
45.26
63.36
45.10
73.94
41.95
1
2
3
4
5
6
Means
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CV, %
70.44
52.25
74.08
91.30
66.33
63.46
69.64
12.97
18.63
41.94
60.73
52.34
40.76
38.81
75.71
82.73
75.96
65.84
73.04
82.22
75.92
6.26
8.25
53.04
52.39
43.03
53.66
50.08
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3
4
5
6
Means
SD
CV, %
58.81
9.81
16.68
53.92
14.02
26.00
46.92
9.34
19.90
50.44
4.36
8.64
140
Table A.19.
(cont.)
Experiment
4
Treatment
Animal
Sheep
Llamas
Low Fiber
1
2
3
4
5
6
Means
SD
CV, %
82.65
90.02
65.66
68.04
83.64
70.34
76.72
9.98
13.01
59.35
51.76
37.47
61.55
55.48
Low Diet
1
2
3
Means
SD
CV, %
53.44
76.26
45.99
58.60
15.43
26.34
40.80
58.80
51.03
48.44
6.45
13.31
Medium Diet
1
2
3
Means
SD
CV, %
47.42
52.17
76.20
58.57
15.77
26.93
55.26
42.44
47.61
50.21
9.03
17.98
53.12
9.52
17.91
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Table A.22
Summary of comparative digestibility estimations in alpaca and sheep.

Magnitude of Difference
of digestibility*
Diet
DM-
CF'
Oat Hay
25.1
27.9
Totora (Scirpus sp.)
17.2
12.7
Oat Hay
0.5
6.4
Alfalfa Hay
5.2
6.3
Oat Silage
2.8
1.6
Totora (Scirpus totora)
1.4
1.0
Alfalfa Hay
0.6
-2.0
Alfalfa Hay (chopped)
-1.0
-2.0
Alfalfa Hay + conc.
-4.8
-7.1
Alfalfa Hay (chopped)

+ conc.
1.9
0.9
Maize Forage +
conc. (6.5% CP)
1.1
1.8
Maize Forage +
conc. (10.5% CP)
-1.0
-0.5
Maize Forage +
conc. (14% CP)
-1.7
-2.9
Maize Forage +
conc. (17% CP)
-2.7
-0.8
Native Pasture
(4.4% CP)
0.2
0.2
Native Pasture
(3.2% CP)
5.2
3.1
Cultivated Pasture
(10.9% CP)
0.1
0.9
Reference
Fernandez Baca &
Novoa (1966)
Bardales
(1969)
Oyanguren
(1969)
Florez
(1973)
Huasasquiche (1974)
San Martin, et al.

(1982a)
San Martin, et al.

1982b)
146
Table A.22.
(cont.)
Magnitude of Difference
of digestibility*
Diet
CF'
Reference
-0.8
4.5
San Martin, et al.

(1982b)
0.0
1.1
Itusaca
(1983)
Oat Hay, Preflowered
-4.7
-14.6
Chayna
(1983)
Oat Hay, Post

Flowered
-0.9
-1.7
Cultivated Pasture
(14% CP)
Alfalfa +
Dactylis Hay
DM-
Alfalfa Hay
2.3
Totora (Scirpus sp.)
4.3
*[Alpaca digestion coefficient - sheep digestion

-coefficient].
^Dry Matter.
Crude Protein.

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COMPARATIVE FORAGE SELECTIYITY AND NUTRITION OF

SOUTH AMERICAN CAMELIDS AND SHEEP

I am especially grateful to committee members

Dr. Jim Pfister, Dr. Bill E. Dahl, Dr. B. Frank Craddock,

The Universidad Nacional

Mayor de San Marcos also provided support for my graduate

the La Raya Research Station also deserve special

My sincere gratitude goes to Dr. Enrique

Franco, Dr. Julio Sumar, Ing. Juan Alpaca, Dr. Victor

to thank Ms. Gretchen Scott for her valuable assistance in

INTRODUCTION AND OBJECTIVES

DIETARY SELECTION BY LLAMAS, ALPACAS, AND

FEED INTAKE AND DIET QUALITY IN LLAMAS,

FEED DIGESTIBILITY AND PASSAGE RATES

Fluid and Particulate Passage

Liquid and Particulate Passage

Liquid and Particulate Passage Rates . . .

SUMMARY AND CONCLUSIONS

tions focused on (1) contrasting the seasonal dietary

On the range sites, llamas ate

more (P < 0.05) tall grasses than alpacas or sheep.

Sheep always select-

ed more (P < 0.05) leaf material than llamas and alpacas,

When ranked from

most selective to least selective, the order was sheep,

Alpacas were the most opportunistic.

Diet similarity between species on improved pasture was

highest between llamas and alpacas.

In the dry season on

a Festuca dolicophylla range site, alpacas and sheep diets

Liquid passage rate in the

first two compartments of the forestomach was faster in

Only when animals ate high quality rations

were the digestibility values compared (P > 0.05) between

Estimated population and distribution of

Average botanical composition (%) of llama,

Botanical composition (%) by plant groups

Botanical composition by plant groups of

Plant parts (%) in the diet of llama,

Similarity indices (%) between llama,

Similarity indices (%) between llama,

Plant species important in llama, alpaca,

Nutritive composition of esophageal

Nutritive composition of esophageal

Nutritive composition of esophageal samples

Ingredients and chemical composition

Particulate passage rate and mean

Rumen fluid volume and dilution rate from

Digestibility coefficients (%) and intake

Digestibility coefficients (%) and intake

Digestibility coefficients (%) and intake

Forage availability during the dry and

Forage availability during the study seasons

Forage availability during the study seasons

Botanical composition (%) of llama, alpaca,

Botanical composition (%) of llama, alpaca,

Scientific names, family and identification

Dry matter fecal output (g/100 kg BW) in

Comparative daily dry matter intake as a

Comparative daily dry matter intake as a

Daily intake values for llamas, alpacas,

Dry matter intake (DMI) and digestible

Average body weight of fecal collector