Animal (2010), 4:9, pp 1460–1471 & The Animal Consortium 2010

doi:10.1017/S1751731110001035
animal

Hair follicle characteristics and fibre production in South

American camelids
M. Antonini-
ENEA Casaccia Depatment BAS, Secondment at the University of Camerino Industrial Liaison Office, Via Anguillarese 301 S.M. di Galeria, 00060 Rome, Italy

(Received 11 December 2009; Accepted 25 March 2010; First published online 8 June 2010)

Hair follicle and fibre characteristics of Peruvian alpaca and llama and Bolivian llama were analysed in three experimental studies.
The first experiment was designed to determine the age at which all the secondary follicles reach maturity, as well as to compare
the skin follicular structure and activity among these different types of Peruvian camelids. It is concluded that the South American
camelids investigated in this study gained a complete and mature skin follicle apparatus at an early age, and hence producers
should practise an early first shearing. A second Peruvian experiment investigated comparative fibre cuticular structure on twenty
Peruvian domestic camelids comprising huacaya, suri and llama (woolly) ‘chacos’ genotypes. The results showed that the number
of cuticular scales per 100 mm fibre length proved to be strongly affected by both the fleece type and the fibre diameter. The suri
fleece was clearly differentiated from those of both huacaya and llama by possessing the highest percentage of fibres with a
number of scales less than eight, the lowest percentage of fibres with more than nine scales, along with the lowest percentage
of fibres with a diameter of more than 35 mm. It is concluded that, with the exception of the scale height, the cuticular parameters
investigated in this study can be utilised in textile fibre analyses for distinguishing among these three types of fleece, as well as in
selection projects designed to produce homogeneous fibres from Peruvian domestic camelids. A further study was conducted to
determine the age at which the hair follicles in Bolivian llamas reach maturity as well as for comparing the skin follicular structure
and activity between the two distinct genotypes. Thirty-one llama kids were chosen. They were born between January and April
1998 and were of different sex and of ‘Q’aras’ (or Carguera) or ‘T’amphullis’ type. Skin biopsies were taken from the right mid-costal
region at 2, 4, 6, 8,10,12 and 14 months of age in order to monitor four follicular parameters. In this experiment, secondary to
primary (S/P) data show that the Bolivian llama population analysed possessed a complete and mature skin follicle apparatus
at birth that remained essentially constant throughout the investigation period. Due to the variation of these traits inside the
same genetic population, the present results showed that T and Q types could only be subjective on the basis of S/P ratio.

Keywords: alpaca, llama, skin hair follicle, fibre cuticular structure, post-natal follicle development

Implication
This paper aims to provide the latest information on the main
fibre production parameters in domestic South American
camelids. The research focuses on two characters: skin fol-
licular structure, fibre cuticular cell and their distribution in
alpaca and llama. In particular, the information is on the first
few months of life of the animals in order to explain how
fleece and skin change during the first year and when the
fleeces are ready to be collected. New differential data on
suri fibre determination and some important clarification on
the classification of Q’aras and T’amphullis bolivian llama
types was provided.
-
E-mail: marco.antonini@enea.it; marco.antonini@unicam.it
Introduction
Major genotypes of South American camelids (SAC)
The SAC population is dominated by alpaca (47%) and llama
(44%), the former being mainly (91%) distributed in Peru,
and the latter (70%) in Bolivia. Smaller populations are
found in other countries, which include ca 70 000 alpaca in
Australia and 1000 in Italy.
Alpaca (Lama pacos L.) is a domestic mammal specialised in
fibre production (Bustinza Choque, 1985; Flores Ochoa and Mac
Quarry, 1995; Bonavia, 1996). Llama have been traditionally
reared as multi-purpose animals providing both meat and fibre
mainly utilised for handcraft products (Lusky et al., 2006).
For alpaca, two different types of fleece are produced by
genotypes identified as huacaya (Figure 1) and suri (Figure 2;
Calle Escobar, 1984; Novoa and Wilson, 1992). Huacaya

1460

Figure 1 Alpaca huacaya.
Figure 2 Alpaca suri.
represents 90% of alpaca fleece processed in Peru (Hoffman and
Fowler, 1995). It is characterised by compact, soft and highly
crimped fibres with blunt-tipped locks that closely resemble
those of Merino sheep. By contrast, the suri product has locks
with a ‘cork-screw’ shape and straight, less-crimped, lustrous,
silky fibres, very similar to mohair of the Angora goat but not as
bright. Differences between fleece types for properties of cell-
scale frequencysecondary to primary (S/P) follicular ratio have
been described previously (Antonini et al., 2001 and 2004).
The genetic background of huacaya and suri is still not
clearly defined because the segregation analyses carried out
Hair follicle and fibre characteristics camelids
in the past do not show consistent data. This is shown by
reports that indicate that suri is inherited as a single recessive
gene (Calle Escobar, 1984), a single dominant gene (Velasco,
1980), a single dominant gene or a haplotype (Ponzoni et al.,
1997; Baychelier, 2000).
Alpaca are particularly valued for fibre production, being
characterised by homogeneously fine, long and soft fleeces,
very much in demand by the European textile industry.
The main differences in fleece between alpaca and llama
may be related to the nature of fibres present. While alpaca
produce a more homotricous fleece as in huacaya and suri,
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Antonini
Figure 3 T’amphullis (or woolly llama).
llama present as more ‘double-coated’ according to the fol-
lowing classification:
(i) ‘Q’aras’ (or ‘carguera’) (Q), which is the typical double-
coated animal, with more guard hair (outercoat) and
markedly less woolly fibres (undercoat) ranging from
short to very short, and
(ii) ‘Chaku (Peruvian expression)/ T’amphullis (Bolivian expres-
sion)’ (or woolly) (T), which is a predominantly single-
coated animal with soft, crimped secondary fibres, but with
a low-quality fleece because of contamination with primary
fibres present (Figure 3).
SAC fleeces are not, however, limited to the few types illu-
strated above, but are characterised by a great variability
because of extensive interbreeding between and within
llama and alpaca and the absence of selection programmes
in both Peru and Bolivia.
This paper reviews information concerning the main
characteristics of fleece and fibres produced by domesticated
SAC in the context of the hair follicles that produce them.
Particular focus is given to anatomy and dynamic behaviour
of the different types of follicles in skin and to the important
and distinguishing properties of fibres such as structure of
cuticular cells, diameter and medullation. Specific attention
is given to the Bolivian llama types.
Hair follicle characteristics and development
Fibres produced by secondary follicles in the skin are generally
of greatest commercial value. For such commercial purpose,
skin follicle productive potential can be measured either as the
ratio of (S/P follicles or as the density of secondary follicles per
1462
unit area of skin. The S/P ratio, however, is often a more
appropriate measure as no allowance need be made either
for skin expansion during animal growth or for shrinkage of
skin at the processing stage (Henderson and Sabine, 1991).
Hence, the S/P ratio has been extensively used both to com-
pare sheep breeds (Carter and Clarke, 1957) and as a means
of measuring secondary follicle maturation after birth (Fraser,
1954; Schinckel, 1955).
It is now well established that in Merino sheep all the
primary follicles are mature at birth (Fraser, 1954; Lyne,
1961), whereas hardly any of the secondary follicles mature
until after birth (Short, 1955). The age at which all the sec-
ondary follicles are fully developed varies between different
sheep breeds, for example, it occurs at one month in the
Herdwick breed, 3 months in both the Scottish Blackface and
the Romney, and at 17 to 18 weeks in Australian Merinos
(Burns, 1949, 1953, 1954a and 1954b; Fraser, 1952 and
1954; Schinckel, 1955).
Dreyer and Marincowitz (1967) showed that secondary
follicle maturation in Angora goats continued until the age of
6 months, with the greatest increase in S/P ratio occurring
within the first 3 months of life. Wentzel and Vosloo (1975)
confirmed that in Angora goats, at 4 months of age, all
secondary follicles produce mohair and this is also in agreement
with the relatively small variation in fibre diameter observed in
mohair of kid shorn at approximately 6 months of age.
Regarding cashmere production, kids of Australian cash-
mere goats, at the age of 20 weeks, have attained their full
complement of secondary follicles and this is conserved until
about 42 weeks of age, when they begin to shed their fleece
(Henderson and Sabine, 1991).

The development of fully functional hair follicle structures
in skin of SAC has not been adequately investigated. This is
despite the fact that the age at which maximal fibre pro-
duction is reached is of fundamental importance for planning
correct management of shearing of alpaca and llama. It is
therefore important to determine the age at which secondary
follicles reach maturity in the commercially valuable hua-
caya, suri and llama chaku genotypes as well as to define
skin follicular structure and the activity and physical prop-
erties of fibres produced.
Characteristics of fibre cuticle
Generally, to characterise and analyse fibre samples from dif-
ferent animals, the major parameters utilised are fibre diameter
(FD), mean cuticular cell scale frequency (MSF), fibre cuticle cell
scale height (SH), its coefficient of variation and appearance
and pattern of cuticular scales (Phan et al., 1988).
MSF is defined as the number of scales per 100 mm fibre
length. It varies from one type to another, with values ranging
from 10 to 12 for Merino wool (Knott, 1990), 6 to 8 for cash-
mere and 6 to 7 for mohair (Phan et al., 1988). The scanning
electron microscopy method introduced by Deutsches Woll-
forschungsinstitut Aachen (Knott, 1990) to measure MSF is one
of the main methodologies in the textile industry for quantita-
tive analysis of blends of different speciality fibres.
Measurement of SH has been mainly utilised to differentiate
between Merino wool (SH .0.6 mm) and mohair fibres
(SH ,0.4 mm) in industrial analysis. Currently, other fine fibres
(cashmere, camel hair, alpaca and yak) are classified together
with SH ,0.4 mm and with a recognised degree of overlap
(Wortmann and Arns, 1988; Wortmann, 1991).
Cuticular scale pattern in llama/alpaca fibre of medium
fineness was defined as mainly cylindrical with near and ripple-
crenated margins that run perpendicular to the fibre axis. In
comparison with other fine fibres, samples from SAC have
shown very occasional similarities to fine wool and cashmere
with scale structures approaching a mosaic pattern.
Despite the apparent inability of textile laboratories to
distinguish between the (secondary) fibres of llama and
alpaca (Phan et al., 1988), there is logic in the proposal that
variations in MSF and SH could be considered useful objec-
tive parameters in animal selection. This approach could
have merit in differentiating the fibres and consequently
the different types of fleeces of individual animals. It is
supported by data from the preliminary investigation on the
number of cuticular scales detected in fibres of Argentinean
camelids (Antonini et al., 2001).
Application of methodology to llama
The situation for classification in llama is complicated and
different methods have been applied according to breeding
cultures in the different Andean areas. What is known is that
Bolivian llama populations are essentially unselected and
show wide variability. For example, Martinez et al. (1997)
analysed variability in a Bolivian llama population for the
following fleece parameters: percentage of medullated
fibres (fragmented medullation, continuous medullation and
Hair follicle and fibre characteristics camelids
kemp), fibre diameter of different fibre types (coarse, fine
and medullated), average fibre length (undercoat and guard
hair), clean fleece yields and presence of crimp.
The results obtained showed that the Bolivian llama fleece
typically consisted of a mix of heterogeneous types ranging
from unmedullated to kempy fibres (Martinez et al., 1997).
While a ‘suri’ type of fleece seemed to segregate with low
frequency within the population, a ‘suri’ standardised flock
was never discovered. The dishomogeneity and the varia-
bility of llama flocks appear as a clear consequence of lack of
interest in fibre selection by breeders. Until recently, llama
fleece was actually mixed with the alpaca fibre in order
to increase the quantity of product although compromising
the quality of the lots of ‘alpaca’ fleece. In contrast, there is
some organised commercialisation of fleece specifically from
‘woolly’ llama flocks in certain Andean highland areas where
the fibre is directly classified as llama lots.
Characteristics of fleeces and fibres have been studied in
llama populations in the Potosı̀ Department of North and
South Lipez Provinces by Quispe et al. (2004). They defined
the average weight of commercial fleece of T’amphullis (T)
as being greater than that of Q’aras (Q) type, while there
was no difference in weight of fleece in the ventral (belly)
part of the body. They also showed no differences between
the two types of llama in average fibre diameter (21.0 mm
(T) and 21.4 mm (Q)) and in percentage of medullation
(13.9% (T) and 15.4% (Q)), while for staple length, T was
longer than for Q (12.0 v. 10.4 cm).
As indicated above, major parameters affecting the
fleece and fibre production characteristics are the structure
and activity of follicles, These, to date, have scarcely been
investigated in SAC. For Bolivian llama, there is particular
lack of information on the pattern fibre growth and the
age at which maximal fibre production is reached and
their variability. Such information is important in decisions
designed to optimise shearing management and methods for
the early selection of animals in the context of an overall
llama management system.
Material and methods
Study 1. Post-natal development of follicles in skin of alpaca
and llama genotypes
The field experimental work was performed in the ‘Alpaquero’
developing centre of Toccra, located in the Arequipa plateau,
Caylloma province, Peru, at 4350 to 5200 m above sea level
(158520 S, 718260 W). The climate of this area is typical of the
high Andean Puna ecosystem, with a mean annual rainfall of
545 mm, 90% of which falls between January and March, and
a mean annual temperature of 1.78C, with an absolute varia-
tion of 268C (211.78C to 15.18C).
Twenty-five animals comprising 10 huacaya (four females
and six males), five suri (two females and three males) and
10 Chaku (six females and four males) were selected for
investigation of the development and activity of skin follicle
structures. These camelids ‘crias’ (kids) were born in January
1998, and were the offspring of females selected from a
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Antonini
flock of SAC used as a reproductive nucleus in a selection
plan of the local SAC population. Crias remained with their
mothers until weaning at 8 months of age and ran with the
rest of the herd according to the Toccra farm management
strategy (Antonini et al., 2004). Animals were managed in
3217 ha of grazing land, divided into three different plant
environments (Bofedales, Pajonales and Tolares), according
to the seasonal availability of biomass.
Skin biopsies were taken from the right mid-side of each
animal by a 0.8 cm-diameter punch positioned approxi-
mately over the 10th rib and about halfway down the body
(Frank et al., 1989). This body area has been found to give
samples more representative of fleece characteristics than
other fleece regions (Martinez et al., 1997). Sampling started
at March 1998, when the animals were 2 months old, and
was repeated at 4, 6 and 10 months of age. Skin samples
were fixed in Bouin solution and stored in ethanol.
After storing, skin samples were dehydrated in a graded
ethanol series and embedded in paraffin. Transverse sections
of 7 mm were cut with a rotary microtome and stained by
using the Sacpic staining procedure, modified by Nixon
(1993). This dyeing method reveals the follicular inner root
sheath (IRS, which shows activities only in the anagen
phase) and the cellular layer of the outer root sheath, which
are the main histological structures for defining skin folli-
cular activity. The level immediately under the sebaceous
gland was defined as the most suitable depth for carrying
out microscopical observations because it contains the
greatest number of detectable skin follicles and thus max-
imises the possibility of observing the IRS.
With the aid of a calibrated ocular micrometer mounted
on an optical microscope, the histological sections under
observation were subdivided into 16 areas of 1 mm2. The
following parameters were recorded for each area: (i) total
number of primary follicles, (ii) active and inactive primary
follicle number, (iii) total number of secondary follicles, (iv)
active and inactive secondary follicle number and (v) number
of medullated fibres in both primary and secondary follicles.
The recorded data were used to calculate:
(a) the FD, defined as the number of follicles per mm2
multiplied by 0.84, which is the coefficient of correction
measured after skin sample shrinkage, due to fixation
and dehydration procedures;
(b) the ratio of S/P follicles;
(c) the percentage of medullated secondary (PMS) fibres
(calculated from the total number of secondary fibres);
(d) the percentage of active secondary (PAS) follicles
(calculated from the total number of secondary fibres).
Analyses of variance (ANOVA) were performed on FD, S/P,
PMS and PAS transformed data (the square root transfor-
mation was used for density data, whereas the arcsine, or
angular, transformation was used for percentage data). As
animals and age were not randomly chosen, a model I
ANOVA was utilised, with fleece type, age and sex con-
sidered as factors. All the statistical analyses were performed
by means of SAS (version 6.12, 1995) statistical software.
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Study 2. Diameter and physical characteristics of cuticle cells
in fibres of alpaca and llama genotypes
With respect to the Cuticular structure analysis, the fibre
samples were taken from the mid-side of 60 Peruvian
domestic camelids, bred at the ‘Alpaquero’ developing centre
of Toccra, located in the Arequipa Plateau, Caylloma Pro-
vince (Peru), at 4350 to 5200 m above sea level (158520 S,
718260 W). Twenty huacaya, 20 suri and 20 llama ‘chacos’ (or
woolly), all homogeneous for sex (male), age (18 months
old), fleece colour (white) and time of shearing (first shear-
ing), were randomly chosen in order to represent a random
sample of the three fleece types.
The fibres were washed in petroleum ether (pure solution),
oven-dried at 608C for 30 min, coated with gold in a sputter-
coating apparatus (Agar Aids) and then examined with the aid
of a scanning electron microscope (Cambridge Stereoscan 250
MK3 Scanning Electron Microscope with EDX-Detector Oxford
Instruments Mod. 6272), set at an accelerating voltage of
15 kV and a working distance of 23 mm (for further details,
see: International Wood Textiles Organization (IWTO) DTM
–XX, 1997). Measurements of the fibre diameter and com-
putation of the number of cuticular scales per 100 mm fibre
length were performed on a sample of 150 fibres for each
specimen. SH was measured on a different sample of 50
fibres. Statistical analyses were performed using the SPSS
12.0 statistical software package (2003).
Study 3. Comparison of skin follicle characteristics between
Bolivian llama T’amphullis (T) and Q’aras (Q) genotypes
Thirty-one llama kids of different sex (20 male and 11 female)
and type comprising 15 ‘Q’aras’ (10 male and 5 female) and 16
‘T’amphullis’ (10 male and 6 female), born between January
and April 1998, were chosen in the study designed to deter-
mine the age at which the hair follicles reach maturity, as well
as to compare skin follicular structure and activity. The experi-
ment was performed at the Patacamaya Experimental Station
in Bolivia, Department of La Paz, at 3789 m above sea level
(178210 S, 678450 W). The climate of this area is typical of the
high Andean Puna semi-arid ecosystem, with a mean annual
precipitation of 402 mm, coefficient of variation of 31%, and a
mean annual temperature of 10.4 C degrees (Valdivia and
Jetté, 1996). Measurements were made and data analysed
using the same methodology as described for Study 1 above.
Results and discussion
Study 1. Post-natal development of follicles in skin
of alpaca and llama genotypes
For the alpaca (huacaya and suri) and llama (chacos) geno-
types, values for the PMS fibres and PAS follicles were at
their maximum by 2 months of age (Table 1) A similar result
was obtained for density of skin follicles except for chacos
for which the maximum value was reached at 10 months.
The fourth parameter, the ratio of secondary to primary fol-
licles, reached its maximum value by 4 months. It was thus
shown that SAC developed a complete and mature skin
follicle apparatus at an early age.
 Hair follicle and fibre characteristics camelids

Table 1 Mean parameters for skin follicles in three types of South although decreasing in month 10 in early spring. Nixon
American camelids (SAC), recorded at four different ages (Antonini (1993) reported that in animals selected for fibre production,
et al., 2004) such as New Zealand predominantly single-coated Angora
Age (months after birth) goats, the activity of secondary follicles remained high from
winter to spring, thus suggesting that the growth of fine, gen-
SAC 2 4 6 10 erally non-medullated fibres, was continuous and not affected
Follicular density (n/mm2) by such seasonal change. Allain and Renieri (2010) have
Huacaya 26.74 a,A
18.27b,A 18.66b,A 22.30c,A recently concluded that predominantly single (secondary-
Suri 25.24a,A 20.75 b,B
15.15c,B 19.90b,B follicle)-coated animals tended to be less affected by changes
Chacos 14.41a,B 12.49b,C 11.43b,C 17.98c,B in photoperiod than those exhibiting active growth by both
Secondary to primary follicular ratio
primary and secondary hair follicles. Thus, it may be assumed
Huacaya 7.33a,A 9.39b,A 8.81b,c,A 8.08a,c,A that the simple, non-scientific selection, employed by farmers
Suri 8.77a,B 8.83 a,A
7.79 a,b,B
6.89b,B (‘campesiños’) for improving fibre quality in huacaya alpaca,
Chacos 4.41a,C 5.87b,B 4.62a,C 4.66a,C could have produced animals possessing secondary skin folli-
cles with activity maintained throughout the winter period and
Percentage of medullated secondary fibres
Huacaya 58.20a,A 46.28a,A 33.57b,A 49.47a,A
addressed mainly to the production of non-medullated fibres.
Suri 82.38 a,B
75.73 a,B
34.47 b,A
79.23a,B A slightly different trend was shown by the ratio of sec-
Chacos 39.38 a,C
44.42 a,A
34.64 a,A
35.39a,C ondary to primary follicle, which, having peaked 2 months
later than the other parameters at month 4, underwent
Percentage of active secondary follicles
a progressive decrease until the end of the observations
Huacaya 96.19a,A 88.33b,A 98.71a,A 77.82c,A
Suri 97.26 a,A
99.71 a,B
84.17 b,B
99.55a,B
(12 months). This indicates that the growth of the new fibres
Chacos 93.89 a,A
81.99 b,C
54.58 c,C
79.90b,A in early spring does not involve the formation of new
secondary follicles, but rather the renewal of activity from
Superscript letters a, b and c are used within the same type of SAC for quiescence. In any case, detection has been performed
comparisons between different ages.
Superscript letters A, B and C are used for comparisons between different referring to follicular activities.
SAC, at the same age. Finally, with reference to the magnitude of the values of
In both comparisons, different letters indicate significantly different values at these four parameters, the huacaya and suri alpaca types
P , 0.01.
generally showed similar maximum values for follicular skin
density and ratio of secondary to primary follicles. By contrast,
The density of skin follicles, PMS fibres and PAS follicles, the Chaku llama type appeared well differentiated from the
with the exception of those of huacaya, were shown to alpaca ones, exhibiting the lowest values throughout for the
decrease until month 6. This was followed by an increase in parameters investigated.
values, which, in the majority of cases, were not significantly It is important that the above findings on follicle numeracy
lower than those recorded at the beginning of the observations and activity be exploited for a rational management of, and
at month 2. In terms of environmental conditions, month 6 fibre production from, the SAC crias (kids). This is because
occurred in July, which is characterised by the coldest tem- the fleece of kids is the most requested in the market and
perature, shortest day length and the lowest pasture produc- llama and alpaca have the potential to produce valuable
tion of the Peruvian Highlands during the Andean dry season. fibre from an early age. Farmer-producers may thus be
Thus, unfavourable environmental conditions are asso- encouraged to practise an early first shearing. This may
ciated with negative effects on both the number and phy- increase their revenues by a value equivalent to one addi-
siological activity of the skin secondary follicles and also tional shearing over the productive life of an animal.
with decreases in medullation. Seasonal effects, possibly It is also evident that the effect of environmental factors on
including photoperiod, may be expected to influence skin follicular skin activities will require particular investigation, to
follicle activity. In addition, a positive correlation between determine the effects of the harsh conditions (environmental
this parameter and the PMS fibres has been previously temperature, raining period, feed availability, effect of photo-
reported for a range of mammalian species and breeds, period, etc.), of the main traditional production areas of the
including goats (Nixon et al., 1991a and 1991b; Nixon, Peruvian and Bolivian Andean plateau.
1993), while extrinsic melatonin/prolactin regulation has
been identified as affecting growth and activity or quies- Study 2. Diameter and physical characteristics of cuticle cells
cence of hair shaft production during the hair follicle cycle in fibres of alpaca and llama genotypes
mechanisms for induction of the medullary cell lineage, Mean values for fibre diameter, cuticular cell scale frequency
including those of intrinsic origin (Galbraith, 2010), which (MSF) and cuticle cell SH are shown in Tables 2 to 4. A con-
appears to be unexplained. The possible association with sistent feature of the data is the wide variability, as shown
reductions in nutrition requires further study. The only by differences between minimum and maximum values and
exception to the general trend towards winter-associated large coefficients of variation.
reductions occurred in the PAS follicles of huacaya, which Table 2 shows the data related to the fibre diameter in the
were essentially maintained during the winter period three different types of Peruvian domestic camelid fleece.

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Antonini

Table 2 Fibre diameter (mm) in three types of Peruvian domestic camelid fleeces (Valbonesi et al., 2010)
95% CI for mean
Type of fleece Number of fibres Mean s.e. Lower boundary Upper boundary Minimum Maximum CV (%)

Llama 3000 29.56 0.24 28.77 30.35 10 92 43.85

Huacaya 3000 27.41 0.18 26.80 28.01 8 72 36.65
Suri 3000 24.42 0.14 23.95 24.89 7 67 31.85

Table 3 Cuticular scales counted on a fibre length of 100 mm in three types of Peruvian domestic camelid fleeces (Valbonesi et al.,
2010)
95% CI for mean
Type of fleece Number of fibres Mean s.e. Lower bound Upper bound Minimum Maximum CV (%)

Llama 3000 9.72a 0.04 9.60 9.84 4 18 20.00

Huacaya 3000 9.11a 0.03 9.02 9.20 5 15 17.78
Suri 3000 7.57b 0.02 7.50 7.65 4 13 17.12
Different superscript letters indicate significantly different values at P < 0.001.

Table 4 Cuticular scale height (mm) measured in three types of Peruvian domestic camelid fleeces (Valbonesi et al., 2010)
95% CI for mean
Type of fleece Number of fibres Mean s.e. Lower bound Upper bound Minimum Maximum CV (%)

Llama 1000 0.40 0.005 0.38 0.42 0.20 1.20 42.09

Huacaya 1000 0.52 0.005 0.50 0.54 0.20 1.10 28.78
Suri 1000 0.47 0.006 0.45 0.49 0.10 1.50 39.80

A nested analysis of variance was carried out to test whether
the means recorded are significantly different from each
other. This analysis indicates no evidence of a significant var-
iance component among the types of fleece (F(2; 57) 5 1.51,
P 5 0.247), yet there is a highly significant (F(57; 8940) 5 42.83,
P , 0.001) added variance component among specimens
within the type of fleece. The percentage contribution to the
total variation, owing to variation among types of fleece,
among specimens within each fleece type and among mea-
surements within each specimen was 1.6%, 21.4% and
76.9%, respectively (Valbonesi et al., 2010).
Table 3 illustrates the data related to the number of cuti-
cular scales counted on a fibre length of 100 mm. The nested
analysis of variance indicates that there is a significant
added variance component among the types of fleece
(F(2; 57) 5 19.61, P , 0.001) as well as among specimens
within the same type of fleece (F(57; 8940) 5 29.14, P , 0.001;
Valbonesi et al., 2010).
Table 4 illustrates the data related to the height of the
cuticular scales. The nested analysis of variance indicates
that there is no evidence for a significant variance compo-
nent among the types of fleece (F(2; 57) 5 2.37, P 5 0.124).
There is a highly significant added variance component
among specimens within the type of fleece (F(57; 2940) 5
39.06, P , 0.001). The percentage contribution to the total
variation owing to variation among types of fleece, among
specimens within each fleece type and among measure-
ments within each specimen was 8.3%, 39.6% and 52.0%,
respectively. (Valbonesi et al., 2010).
The fitting of a linear regression model (with R2 5 0.29)
for describing this relationship is illustrated in Table 5. These
results were confirmed by regression analyses carried out on
each fleece type. The huacaya type of fleece showed no
significant regression between the number of scales and the
diameter of the fibres (b 5 0.004, P 5 0.386), whereas a
significant positive regression was present in both llama
(b 5 0.054, P , 0.001) and suri types of fleece (b 5 0.029,
P , 0.001).
By exploiting a different statistical approach, the data
related to the two discrete variables, fibre diameter and
number of cuticular scales were arranged into four frequency
classes according to the criterion reported in the previous
section. The data related to the frequency distribution of
these classes were arranged into a contingency table to test
whether they were independent of fleece type (Table 6). The
results of this analysis (x2(6) 5 113.68; P , 0.001) show
clearly that the frequency of the different classes of fibre
diameter is dependent on fleece type. In particular, the fre-
quencies of the class comprising the thickest fibres (diameter
above 35 mm) in llama, huacaya and suri significantly differ
from each other (llama v. suri 1 huacaya: x2(1) 5 68.51,
P , 0.001); huacaya v. suri: x2(1) 5 29.44, P , 0.001).

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 Hair follicle and fibre characteristics camelids

Table 5 Estimate of parameters related to the dependent variable of number of cuticular scales* (Valbonesi et al., 2010)
95% confidence interval
Parameters Coefficients s.e. t P Lower boundary Upper boundary

Intercept 6.877 0.159 43.307 0.000 6.565 7.188

Huacaya fleece 2.121 0.212 9.988 0.000 1.705 2.537
Llama fleece 1.262 0.198 6.363 0.000 0.873 1.651
Fibre diameter 0.029 0.006 4.611 0.000 0.016 0.041
Huacaya fleece fibre diameter* 20.025 0.008 23.124 0.002 20.040 20.009
Llama fleece fibre diameter* 0.025 0.007 3.429 0.001 0.011 0.039
*The suri type is used as reference fleece.

Table 6 Frequencies of four classes of fibre diameter in three different types of Peruvian domestic camelid fleeces (Valbonesi
et al., 2010)
Classes of fibre diameter (mm)
,20 20 to 27 28 to 35 .35
Types of fleece n (%) n (%) n (%) n (%) Total

Llama 631 (21.0) 1011 (33.7) 586 (19.5) 771a (25.7) 3000
Huacaya 620 (20.7) 1146 (38.2) 686 (22.9) 549b (18.3) 3000
Suri 763 (25.4) 1391 (46.4) 569 (19.0) 277c (9.2) 3000
Total 2014 (22.4) 3548 (39.4) 1841 (20.4) 1597 (17.7) 9000
Different superscript letters indicate significantly different values at P , 0.001.

Table 7 Frequencies of four classes of cuticular scale number, counted on a fibre length of 100 mm in three different types of
Peruvian domestic camelid fleeces (Valbonesi et al., 2010)
Classes of cuticular scale number
,8 8 9 .9
Types of fleece n (%) n (%) n (%) n (%) Total

Llama 343a (11.4) 477 (15.9) 649 (21.6) 1531a (51.0) 3000
Huacaya 394a (13.1) 637 (21.2) 909 (30.3) 1060b (35.3) 3000
Suri 1586b (52.9) 791 (26.4) 380 (12.7) 243c (8.1) 3000
Total 2323 (25.8) 1905 (21.2) 1938 (21.5) 2834 (31.5) 9000
Different superscript letters indicate significantly different values at P , 0.001.

In addition, the variable cuticular scale number, monitored
on 100 mm of fibre length, was distributed into four con-
tiguous classes (Table 7). In testing their independence of
the fleece type, a significant (P , 0.001) value for x(6)2 5
864.92 was obtained that rejects the null hypothesis of
independence. A further investigation on subsets of the data
showed that (i) the frequency of the class comprising the
lowest number of scales (below eight) in suri significantly
differed from those of llama and huacaya (suri v. huacaya 1
llama: x(1)2 5 601.74, P , 0.001), whereas the last two
frequencies did not differ from each other (huacaya v. llama:
x(1)2 5 0.81, P 5 0.369), and (ii) the frequencies of the class
comprising the largest number of scales (above nine) in
llama, huacaya and suri significantly differed from each
other (suri v. huacaya 1 llama: x(1)2 5 399.62, P , 0.001;
huacaya v. llama: x(1)2 5 60.09, P , 0.001).
Finally, the frequency classes of these two variables were
jointly analysed in all of the pairwise combinations, and the
resulting 16 subgroups of data were used as variables to
carry out a stepwise discriminant analysis. The original set of
variables was reduced to three variables: variable 1 (which
corresponds to the number of fibres with a diameter smaller
than 20 mm and bearing less than eight scales), variable 5
(number of fibres with a diameter ranging from 20 to 27 mm
and bearing less than eight scales) and variable 16 (number
of fibres with a diameter larger than 35 mm and bearing
more than nine scales). The data relating to these variables
are reported in Table 8. Of the two canonical discriminant
functions used in the analysis, the first accounts for 89.5% of
the variance (Table 9). Furthermore, as the Wilks’ lambda
value, associated with the first canonical discriminant func-
tion, is close to zero, it can be assumed that the differences

1467
Antonini

Table 8 Frequencies of the three variables, selected by the stepwise discriminant analysis in three different types of Peruvian
domestic camelid fleeces (Valbonesi et al., 2010)
Variable 1 (ø , 20; Ns , 8)* Variable 5 (ø 5 20 to 27; Ns , 8) Variable 16 (ø . 35; Ns . 9)
Types of fleece n (%) n (%) n (%) Total

Llama 134 (21.9) 97 (10.3) 500 (65.3) 731

Huacaya 57 (8.9) 157 (15.0) 206 (26.9) 420
Suri 452 (70.2) 792 (75.7) 60 (7.8) 1304
Total 643 1046 766 2455
*ø 5 fibre diameter (mm); Ns 5 number of cuticular scales.

Table 9 Parameters related to the canonical discriminant functions (Valbonesi et al., 2010)
Standardised coefficients
Functions Eigen value Percentage of variance Canonical correlation Wilks’ l P* Variable 1 Variable 5 Variable 16

1 7.445 89.5 0.939 0.092 ,0.001 0.469 0.921 20.375

2 0.872 10.5 0.589 0.682 0.005 0.601 0.339 0.938
*x2 transformation of Wilks’ l is used along with the degree of freedom to determine significance.
Different superscript letters indicate significantly different values (P , 0.001).

among the three types of fleece are adequately described by
this function alone. By inspection of the standardised coef-
ficients associated with this function, it can be seen that
large values of this function correspond to suri fleece that
exhibits a large number of fibres with a small diameter
(<27 mm) and a small number (less than eight) of scales (i.e.
variables 1 and 5) along with a reduced number of fibres
with a large diameter (.35 mm) and a large number (more
than nine) of scales (variable 16). By using these three
variables in a classification analysis, 86.7% of the specimens
investigated are assigned to their original type of fleece. In
particular, the suri fleece appears clearly differentiated from
the other types, as 100% of cases are correctly classified v.
the 85.0% and 75.0% of cases correctly classified in huacaya
and llama, respectively.
All this evidence leads to the conclusion that, in contrast
to what has already been reported in the literature (Phan
et al., 1988), it is possible to distinguish between the llama,
huacaya and suri fleeces. In short, rather than considering
mean values of single cuticular parameters, an objective
identification of these three types of camelid fleece may be
better achieved by a multifactorial approach. This could
involve the combination of the two parameters, such as scale
frequency and fibre diameter, in the form of a bivariate dis-
tribution coupled with the frequency distributions of fibre
classes (identified here on the base of these two parameters
and either separately or jointly analysed).
On the basis of these results and analysis, it may be con-
cluded that the suri fleece, which exhibits the highest per-
centage (90.8%) of fibres with a diameter ,35 mm and the
lowest mean value (7.57/100 mm) of cuticular scales and
very close to those reported for both mohair and cashmere
(Phan et al., 1988), could be differentiated by huacaya and
llama fibre on the basis of fibre diameters and lowest number
of cuticular scales. Hence, in selection projects addressed to the
production of homogeneous fibres from Peruvian domestic
camelid fleeces, the main target should be directed towards
the breeding of the suri genotype of alpaca.
Study 3. Comparison of skin follicle characteristics between
Bolivian llama T’amphullis (T) and Q’aras (Q) genotypes
The parameter affecting the fleece and fibre production
characteristics is the skin follicular structure and activity, but
until now it has been scarcely investigated, despite the fact
that the age at which llama maximal fibre production is
reached is of fundamental importance for understanding
the variability of the Bolivian llama population, the correct
shearing management and the early animal selection. The
aim of this work is to fill this gap by determining the age at
which all the secondary follicles reach maturity in T and Q
types, as well as to compare the skin follicular structure and
activity among these different types of Bolivian llamas.
Differences between T’amphullis (T) and Q’aras (Q) llama
types in S/P ratio, secondary follicular activities and medul-
lated primary fibre (%) were not significant (Table 10) and
results are expressed as means combined for both types.
However, T types presented a consistently higher percentage
of active primary follicles and lower medullated secondary
fibre, apart from the April sample collections, than the Q type
llama. No significant effect of sex was shown in any of the
parameters analysed between the two types of llama. There
was also very limited change in the S/P ratio averaged for
both genotypes during the time period of sampling, which
reflects increases in age of the llama studied. The differences
between Q and T in active primary follicle and in medullated
secondary fibres show at first analysis a general reduction
of activities in Q with respect to T type. On the other hand,
no consistent behaviour was noted with respect to the

1468
 Hair follicle and fibre characteristics camelids

Table 10 Means parameters for skin follicle in Bolivian llama types recorded at seven different ages (Lusky et al., 2006)
Age (months after birth)
Llama type 2 (April) 4 (June) 6 (August) 8 (October) 10 (December) 12 (February) 14 (April)

S/P
T and Q 5.49 5.46 5.34 5.35 5.33 5.48 5.45
Active secondary follicle (%)
T and Q 51.49c,d 44.25c 44.50c 62.69b 26.49a 26.50a 57.93b,d
Active primary follicle (%)
T and Q 71.31 80.60 75.83 84.58 77.70 54.95 74.80
T 100a 86.25a,b 80.82a,b,c 83.98a,b 88.02a,b 72.41b,c 71.42c
Q 64.28c 78.46a,b 71.21 b,c
84.89a 70.83b,c 38.88d 77.76a,b,c
Medullated secondary fibre (%)
T and Q 19.02 23.24 17.67 17.98 7.72 27.08 15.33
T 42.43a 17.83b 15.06b 14.82b 3.77c 5.98c 20.76b
Q 13.29c,d 24.38b 20.26b,c 19.62c 10.35d 46.50a 11.05d
Medullated primary fibre (%)
T and Q 84.42a 89.78a,b 80.37b 76.12b 57.91c 75.61b 74.54b
Different superscripts are significantly different. a,b,c and d P < 0.05.

secondary fibre medullation between the two types. For
example, the Q type produces less medullated fibre in April,
but had greater values than the T types in the other periods
studied. There also appears to be no homogeneous beha-
viour of the follicular activities during all of the experimental
period. In addition, the environmental and photoperiod
effects may be expected to affect the expression of the skin
follicular characters and therefore deeper investigation is
needed to verify their effects.
The great genetic variability of the Bolivian llama
(Lauvergne 1994 and Lauvergne et al., 2001) provides a
basis for explaining the morphological differences especially
in fleece characters between the T and Q genotypes. These
also need to be considered in the context of influences
arising from environmental effects of the Bolivian husbandry
environment as indicated above and by the absence of a
defined genetic selection plan.
Frank (2001) supported this theory because he defined the
genetic heredity of the double (Q) and single coated (T)
Argentinean llama as being obtained by an additive effect
where the character of simple and double coat bearing in the
llama population seems to be more a quantitative than quali-
tative parameter. Moreover, the results of Quispe et al. (2004)
describing the fibre and fleece characteristics of the Bolivian
llama population studied on the South Lipez Provinces seem to
confirm the quantitative characteristics of the double and simple
fleece structure because there are no parameters in the fleece
that can characterise specifically the two type of llama (Q and T).
In fact, Quispe et al.’s (2004) results did not show any significant
differences in belly fleece weight, average fibre diameter and
percentage of medullated fibres between T and Q llama types.
Lusky et al. (2006) also showed, from analysis of the
Bolivian llama population, that a complete and mature skin
follicle apparatus is present at birth and remains constant
until the age of 14 months. For this reason, T and Q types
could be considered only a subjective classification due to
the variation of the different characters composing the fleece
within the same population. As the llama types are generally
considered double-coated animals, we can assume that the
relation between the under coat and the outer coat in this
population seems to be governed by quantitative characters
that are difficult to classify in specific categories.
In both the T and Q Bolivian llama types, the ratio of
secondary to primary follicles reached maximum value by
2 months of age. This result contrasts with those of Antonini
et al. (2004) for Peruvian llama in which the S/P Ratio achieved
its maximum value after a further 2 months. It was thus con-
firmed that Domestic SAC (alpaca and llama) gained complete
and mature skin follicle development at an early age. Moreover,
the variation seems to be associated with seasonal changes, in
which unfavourable environmental conditions may negatively
affect the physiological activity of skin follicles.
The results obtained confirm that Bolivian llama fleece typi-
cally consists of a mix of heterogeneous types of fibre. The
dishomogeneity and variability of llama flocks appear as a clear
consequence of lack of interest in fibre selection by breeders.
The follicle parameters affecting the fleece and fibre production
characteristics show no differences between T and Q.
Such information is important in decisions designed to
optimise shearing management and methods for the early
selection of animals in the context of an overall llama
management system.
The results that define the production patterns of juvenile
llama, when taken in addition to those obtained by Lusky et
al. (2006), may be exploited for an improved management of
the ‘crias category’. This is particularly so since llama kid
fleece is the most requested in the market, and fibre pro-
duction is present in llama from an early age. Producers
should thus be advised to practise early selection of breeding
animals based on fibre production potential, as well as an

1469
Antonini
anticipated earlier first shearing. One suggestion that may
be advanced to increase revenues for llama producers would
be to breed llama for meat and exploit the maximum
potentiality of the baby fibre production before the slaugh-
ter. This would be designed to increase the number of
shearings by one to take advantage of the finer quality of
juvenile fleece that is available until the age of 14 months.
Conclusion
This research carried on alpaca and llama fibre basically
shows the three main results.
(a) Maturity of the Domestic SAC. Alpaca and llama gained
a complete and mature skin follicle apparatus at an early
age as in Peruvian and Bolivian camelids. In Peruvian
alpacas and llamas, skin follicular maturity was achieved
in a 4-month-old, while in Bolivian llamas it was a little
anticipated in the 2-month-old. This apparatus, however,
did not remain constant throughout the period of the
investigation. The unfavourable environmental condi-
tions might negatively affect both the number and the
physiological activity of the skin secondary follicles.
The data presented here may be exploited for a rational
management of the ‘crias category’. As kid fleece is the
most requested in the market, and fibre production
potential is present in llama and alpaca from an early age,
producers should practise an anticipated first shearing.
This would increase producers’ revenues by a value
equivalent to one shearing over the productive life of an
animal. In particular, llama kid fleece is the most
requested in the market, and fibre production is present
in llama from an early age. Producers should practise early
selection based on fibre production potential, as well as
anticipated first shearing. This would increase producers’
revenues by a value equivalent to one shearing over the
productive life of an animal especially for llama meat
production purposes.
In order to substantiate the present findings, further
studies are required using larger numbers of animals and
for periods of at least 3 years. This would also enable the
effect of any environmental factors on follicular skin
activities (which are responsible for fibre production) to
be investigated, potentially a very important considera-
tion in the harsh conditions of the Andean plateau.
(b) Possible differentiation between suri and hucaya alpaca
and llama fibre. All of the evidence carried on in study 2
leads to the conclusion that it is possible to distinguish suri
fleece from both llama and huacaya fleeces on the basis of
the average number of cuticular scales. However, rather
than considering only this single, discriminant cuticular
parameter, the results suggest a combination of this
parameter with the fibre diameter.
It can thus be stated that suri that exhibits the highest
percentage (90.8%) of fibres with a diameter ,35 mm and
the lowest mean value (7.57) of the number of cuticular
scales, very close to those reported for both mohair and
1470
cashmere, is the best fleece for textile fibre production. A
further development of this work will be to verify whether
the parameters utilised for distinguishing among the three
fleece types are suitable for more heterogeneous condi-
tions, that is, for animals exhibiting different sex, age,
fleece colour and different time of shearing.
(c) Classification of Bolivian llama types of fleece. Study 3
shows that since S/P is the main parameter to define the
follicular structure, T and Q types could be only a
subjective classification due to the variation of the
characters inside the same genetic population. Fibre
analyses on fleece samples drawn at the same time of
the skin sample collection will have to be carried out in
order to provide further and defined information on
Bolivian llama types and to confirm the skin follicular
structure data obtained in this paper.
Acknowledgements
This review is based on an invited presentation at the 59th
Annual Meeting of the European Association for Animal Pro-
duction held in Vilnius, Lithuania during 24–27 August 2008.
References
Allain D and Renieri C 2010. Genetics of fibre production and fleece
characteristics in small ruminants, Angora rabbit and South American camelids.
Animal 4, 1472–1481.
Antonini M, Gonzales M and Valbonesi A 2004. Relationship between age and
postnatal skin follicular development in three types of South American domestic
camelids. Livestock Production Science 90, 241–246.
Antonini M, Gonzales M, Frank E, Hick M, Pierdominici F, Catalano S and
Castrignanò F 2001. Cuticle cell mean scale frequency in different type of fleece
of domestic South American camelids. In Progress in South American camelids
research (ed. M Gerken and C Renieri), EAAP no. 105, pp. 110–116.
Wageningen Pers, Wageningen, The Netherlands.
Baychelier P 2000. Suri and huacaya: two alleles or two genes? In Proceeding
of Australian Alpaca Association. National Confonference Camberra, Australia,
pp. 79–85.
Bonavia D 1996. Los Camelidos Sudamericanos (South American Camelids) Una
introducción a su estudio. IFEA-UPCH, Conservation International, Lima, Peru.
Burns M 1949. Study on follicle population in relation to fleece change in lambs
of the English Leicester and Romney breeds. The Journal of Agricultural Science
39, 64–79.
Burns M 1953. Observation on the follicle population of Blackface sheep. The
Journal of Agricultural Science 43, 422–431.
Burns M 1954a. The development of the fleece and follicle population in
Herdwick sheep. The Journal of Agricultural Science 44, 443–464.
Burns M 1954b. Observations on the development of the fleece and follicle
population in Suffolk sheep. The Journal of Agricultural Science 44, 86–99.
Bustinza Choque V 1985. Razas de Alpacas del Altiplano: Suri y Wakaya.
Universidad Nacional del Altiplano, Instituto de Investigaciones para el
Desarrollo Social del Altiplano, Puno, Peru.
Calle Escobar R 1984. Animal breeding and production of American camelids.
Ron Hennig, Patience, Lima, Peru.
Carter HB and Clarke WH 1957. The hair follicle group and skin follicle
population of some non-merino breeds of sheep. Australian Journal of
Agricultural Research 8, 109–119.
Dreyer JH and Marincowitz G 1967. Some observations on the skin histology
and fibre characteristics of the Angora goat (Capra hircus angoraensisi). South
African Journal of Agricultural Science 10, 477–500.
Flores Ochoa J and Mac Quarry K 1995. I Camelidi andini (Andean Camelids).
In L’oro delle Ande (The gold of Andes) (ed. FO Patthey and Sons), vol. 1,
pp. 23–35. Soto Impresor, Madrid, Espana.

Frank EN 2001. Description y analisis de segregacion de fenotipos de color y
tipos de vellon en llamas argentinas. (Segregation analysis and description of
colour and fleece types phenotypes in Argentinean llamas). PhD, Buenos Aires
University, pp. 151–153.
Frank EN, Parisi de Fabro SG and Mendez T 1989. Determinacion de variables
foliculares en cortes de piel de camelidos sudamericanos y su relacion con las
caracteristicas de vellon. Revista Argentina de Producción Animal 9, 379–386.
Fraser AS 1952. Growth of the N-type fleece. Australian Journal of Agricultural
Research 3, 435–444.
Fraser AS 1954. Development of the skin follicle population in Merino sheep.
Australian Journal of Agricultural Research 5, 737–744.
Galbraith H 2010. Fundamental hair follicle biology and fine fibre production in
animals. Animal 4, 1490–1509.
Henderson M and Sabine JR 1991. Secondary follicle development in Australian
cashmere goats. Small Ruminant Research 4, 349–363.
Hoffman E and Fowler ME 1995. The alpaca book: management, medicine,
biology, and fiber, 1st edition. Clay Press, Inc., Herald, CA, USA.
International Wood Textiles Organization (IWTO DTM–XX) 1997. Quantitative
analysis of blend of wool with speciality fibres by scanning electron microscopy.
International Wool Textiles Organization, Brussels, Belgium.
Knott J 1990. Fine animal fibres and their depigmentation process. EEC Comett
Eurotex, Universidad do Minho, Guimaraes, Portugal.
Lauvergne JJ 1994. Characterization of domesticated genetic resources of
American camelids: a new approach. In Proceedings of the European
Symposium of South American Cemelids (ed. M Gerken and C Renieri),
pp. 59–63. University of Camerino Publishing, Camerino, Italy.
Lauvergne JJ, Martinez Z, Ayala C and Rodriguez T 2001. SUPREME project:
identification of a primary population of South American domestic camelids in
the provinces of Antonio Quijarro and Enrique Baldivieso (Department of Potosı̀,
Bolivia) using phenotypic variation of coat colour. In Progress in South American
camelids research (ed. M Gerken and C Renieri ), EAAP no 105, pp. 64–74.
Wageningen Pers, Wageningen, The Netherlands.
Lusky T, Valbonesi A, Rodriguez T, Ayala C, Weimin L and Antonini M 2006. Skin
follicular structure in Bolivian llamas. In South American camelids research.
Proceedings of the 4th European Symposium on South American Camelids and
DECAMA European Seminar, Vol. 1 (ed. M Gerken and C Renieri), EAAP
publication, pp. 207–216. Wageningen Pers, Wageningen, The Netherlands.
Lyne AG 1961. The post natal development of wool follicles, shedding, and skin
thickness in inbred Merino and southdown- Merino crossbred sheep. Australian
Journal of Agricultural Research 14, 141–156.
Martinez Z, Iniguez LC and Rodriguez T 1997. Influence of effects on quality
traits and relationships between traits of the llama fleece. Small Ruminant
Research 24, 203–212.
Nixon AJ 1993. A method for determining the activity state of hair follicle.
Biotechnology and Histochemistry 60, 316–325.
Hair follicle and fibre characteristics camelids
Nixon AJ, Gurnsey MP, Betteridge K, Mitchell RJ and Welch RAS 1991a.
Seasonal hair follicle activity and fibre growth in some New Zealand cashmere
bearing goats (Capra hirus). Journal of Zoology of London 22a, 589–598.
Nixon AJ, Saywell DP and Bown MD 1991b. Fibre growth cycles and medullation
in Angora goats. Proceeding of New Zealand Society of Animal Production 51,
359–364.
Novoa C and Wilson T 1992. The management of global animal genetic
resources. FAO, Animal Production and Health 104, pp. 189–203, Rome, Italy.
Phan KH, Wortmann FJ, Wortmann G and Arns W 1988. Characterization of
specialty fibres by scanning electron microscopy. Shriftenz Deutsches
Wollforschungsinstitut 103, 137–162.
Ponzoni RW, Hubbard DJ, Kenyon RV, Tuckwell CD, McGregor BA, Howse A,
Carmichael I and Judson GJ 1997. Phenotypes resulting from huacaya
by huacaya, suri by huacaya and suri by sury alpaca crossings. In Proceeding
of the International Alpaca Industry Seminar. NSW, Sydney, Australia,
pp. 11–13.
Quispe JL, Antonini M, Rodriguez T and Martinez Z 2001. Clasificacion y
caracterizacion de fibra de Llamas criadas en el altiplano sur de Bolivia
(Classification and characterization of llama fibre raised in the south of Bolivian
altiplano) (ed. M Gerken and C Renieri). In Progress in S.A. Camelids Res. EAAP.
Pub. No. 105, pp. 286–294.
SAS 1995. SAS user’s guide: statistics, version 6.12. SAS Inst. Inc, Cary, NC, USA.
SPSS 2003. SPSS 12.0 ‘‘user’s guide’’. SPSS Inc., Chicago, IL, USA.
Schinckel PG 1955. The post-natal development of the skin follicle population
of a strain of Merino sheep. Australian Journal of Agricultural Research 6,
68–76.
Short BF 1955. Development of the secondary follicle population in sheep.
Australian Journal of Agricultural Research 6, 62–67.
Valbonesi A, Cristofanelli S, Pierdominici F, Gonzales M and Antonini M 2010.
Comparison of fiber and cuticular attributes of alpaca and llama fleeces. Textile
Research Journal 80, 344–353.
Valdivia C and Jetté C 1996. ‘‘Peasant households in semi-arid San José:
confronting risk through diversification strategies.’’ IBTA 181/Technical Report
49/SR-CRSP 47. La Paz, Bolivia.
Velasco J 1980. Mejoramiento Genético de Alpacas. Anales III Reunion
Cientifica Animal. Sociedad Peruana de Produccion Animal, Lima, Peru.
Wentzel D and Vosloo LP 1975. Dimensional changes of follicles and fibres
during pre- and postnatal development in the angora goat. Agroanimalia 7,
61–64.
Wortmann FJ 1991. Quantitative fibre mixture analysis by scanning electron
microscopy, part 3: round trial results on mohair/wool blends. Textile Research
Journal 61, 371–374.
Wortmann FJ and Arns W 1988. Untersuchungen zu den Schuppenkantenho-
henverteilungen von wolle und edlend Tierhaaren. Schriftenr Deutsches
Wollforschungsinstitut Institue 102, 89–97.
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