Small Ruminant Research 61 (2006) 113–129
Phenotypic and genetic description of fibre traits in South
American domestic camelids (llamas and alpacas)
夽
E.N. Frank
, M.V.H. Hick
, C.D. Gauna
,
H.E. Lamas
, C. Renieri
, M. Antonini
a
SUPPRAD Programme, Catholic University of C´ordoba, Argentina
b
SUPPRAD Programme, Univ. Nac. of La Pampa, Argentina
c
SUPPRAD Programme, INBIAL, Argentina
d
University of Camerino, Italy
e
ENEA C.R. Casaccia Biotec Agro, Italy
Available online 22 September 2005
Abstract
Even though llamas and alpacas are multipurpose animals, fibre production remains the main trait from an international market
point of view. The objectives of this review are to describe the phenotypic traits that determine fibre quality, and to identify
the genetic mechanisms governing them. The finer and lesser prickling effect the fibre has, the higher its value is. All these
characteristics are related to fibre diameter and evenness, and to other traits such as color, type of fleece, fibre length and yield.
Studies on genetic mechanisms for llama and alpaca fleece traits show that the white phenotype is dominant to the pigmented
phenotype and to the spotted phenotype. Black face and extremities phenotypes are dominant to black and wild phenotypes. Lustre
is dominant to non-lustre type and double coated is governed by an additive genetic mechanism. Heritabilities of fleece weight,
staple length and fibre diameter are low to moderate in the high plateau environment and very high outside Altiplano conditions.
© 2005 Elsevier B.V. All rights reserved.
Keywords: Fibre production; Fibre quality; Genetics mechanism; Color; Fleece types
夽
This paper is part of the special issue entitled Special Issue on
Camelids, Guest Edited by Dr. Ahmed Tibary and Dr. Steve Parish.
∗
Corresponding author. Present address: Obispo Trejo 323,
C´ordoba, ZC X5000IYG, Argentina.
E-mail address: frank@uccor.edu.ar (E.N. Frank).
1
SUPPRAD: Sustentabilidad Productiva de Peque˜nos Rumiantes
en ´
Areas desfavorecidas [Small Ruminant Productive Sustainability
in less favoured Areas].
1. Introduction
Alpacas have been historically bred to produce fibre
and meat (
). However, lamas are con-
sidered a multi producer animal (fibre, meat, trans-
portation, skin, fertilizers), especially in the semi arid
areas of the high altitude plateau (‘punas’) (
Camelid fibre value, as in all other fibre producing
species, is highly dependent on mean fibre diameter.
0921-4488/$ – see front matter © 2005 Elsevier B.V. All rights reserved.
doi:10.1016/j.smallrumres.2005.07.003
114
E.N. Frank et al. / Small Ruminant Research 61 (2006) 113–129
However, other traits such as coat colour, and fleece
type play an important role (
Description of the phenotypic and genetic parame-
ters related to fibre production and quality are necessary
for the design of an effective genetic improvement pro-
gram (
While quantitative traits are largely influenced by
environmental effects, the fleece type and coat colour
are governed by Mendelian mechanisms, in which the
phenotypic manifestation is not expected to be influe-
nced by environmental effects (
The objectives of this review are to describe know
information and recent advance on phenotypic traits
that determine fibre quality and the genetic mechanisms
governing those characteristics.
2. Factors affecting commercial value of
camelids fibre
2.1. Textile requirements
Currently, lighter clothes are worn in close contact
with the skin. Thus, it is important to consider the prick-
ling sensation of the fibre. For wool, it has been clearly
established that this particular sensation is related to the
fibre diameters distribution (
In the case of camelid fibre, this aspect is also related
to the type of fleece. The prickling sensation is obvious
in double coat fleece requiring separation of the pri-
mary fibres or dehairing (
; Loro
Piana, P.L. personal communication). This is related to
the frequency of fibres coarser than 30
m (
Phillips, 1995; McGregor, 1997
In a study on North American consumer’s wool fab-
rics preference, commissioned by International Wool
Secretariat (IWS), the most preferred traits were
closely related to the fibre diameter (
It is highly probable that prickle sensation in alpaca
cloth will be related to the incidence of fibre diameter
>30
m (or around 32 m), and to the bending rigidity
of this fibres (
). The International
Alpaca Association also considers as alpaca fibre the
unbristled llama fibre (
) and therefor
it is presumed that llama fibre will be treated similarly
to alpaca fibre by the textile industry.
2.2. Market signals
Traditionally the commercial value of South Ameri-
can camelids (SAC) fibre was determined primarily by
colour (
). In a survey of alpacas
fibre prices from 1969 to 1981, pigmented fibres rated
in average about 65.9% of white fibre prices (
) but this difference was lower between
1995 and 2002 (
). In some cases, pig-
mented fibres command higher prices than white fibres
(alpacas and llamas) (
shows the relative importance of each alpaca
fleece characteristics on value for the textile industry.
Table 1
The relative commercial importance of greasy raw specialty animal fibre characteristics
Greasy fibre characteristics
Relative processing significance
Scoured
Top/noil
Yarns
Cloth
Mean fibre diameter
****
****
****
****
Washing yield
****
Vegetable matter contamination (amount and type)
***
***
**
**
Mean fibre length
**
***
*
Staple strength/position of break
**
*
Clean fibre color
*
*
*
Incidence of dark fibres
*
*
*
***
Mean fibre diameter variability
**
**
Proportion of fibres >30
m
*
**
Fibre length variability
**
**
*
*
Resistance to compression (crimp)
*
*
**
**
Incidence of cotts
**
Degree of staple tipiness
*
Style and handle
*
*
References: ****: highly significant; *: some significance. Source:
E.N. Frank et al. / Small Ruminant Research 61 (2006) 113–129
115
Fig. 1. (a) Effect of fibre diameter on the price of raw llama
fibre (1999-2004). Baby (<19.0
m); superfine (19–21.9 m); fine
(22–24.9
m); medium (25–29.9 m); strong (>30 m). Source:
, H.E. Lamas (unpublished). (b) Effect of alpaca fibre
diameter (and type of fleeces) on the relative prices of white alpaca
tops (1985–2003). B SUT: white Suri top (27
m); B BAT: white
baby top (20–22
); B SFT: white superfine top (25.5 m); B ADT:
white adult top (27.5
m). Source: plotted from data supplied in
The commercial characteristics may vary depending on
whether the fibre is knitted by the woven system or the
knitwear system. It is suggested that raw fibre follows
the same tendencies as the top (
As stated above, fibre diameter determines fleece
value. This is clearly illustrated by
a which
shows the relationship between price and fibre in
Argentina altiplano, and in
b which shows the
relationship between prices and mean fibre diameter
of alpacas tops. A great variability in prices is seen
from one year to another within each fibre class. Sim-
ilar findings have been reported in an Australian study
(
2.3. Objective measurements
Fibre diameter determines the potential minimum
mass per unit lengths of yarns, and consequently the
thickness that textile technologists can spin. The qual-
ity of the yarn is strongly correlated with the softness
and prickliness in alpaca, which are both related to
mean fibre diameter and proportion of fibres <30
m
(
). Therefore, fibre diameter rep-
resents the most important factor in the price of mills
products (tops) (
Villarroel Le´on, 1991; Vinella, 1994;
Antonini and Vinella, 1997; Anonymous, 2005
). Nev-
ertheless, only 15% of the larger alpaca farmers receive
differential price for the mean fibre diameter. This sit-
uation led small Peruvian alpaca breeders to increase
fibre diameter (
Velarde Flores, 1988; Anonymous,
) which in turn impacted negatively the value of
domestic SAC fibre in the international stock market
(
Villarroel Le´on, 1991; Vinella, 1994; Anonymous,
In Argentina, about 35% of llama fibre is classi-
fied and sold according to its mean fibre diameter with
a classification system similar to argentine wool and
Peruvian alpaca fibre (
) (SUPPRAD
Programme, Lamas, H.E., unpublished).
The quality of fibre is also determined by the uni-
formity of the diameter. It is generally accepted that a
variation of 5% in fibre diameter implies an increase
or decrease of 1
m in diameter (
Fibre length and length variation also affects fleece
quality because affects the process of yarn production.
Worsted length in alpaca of about 75 mm is considered
ideal. Shorter fibres are more economically processed
by woolen process where final product rate lower value
(
A number of others less important measurable char-
acteristics can affect fibre value, both in alpaca and
llama. The proportion of fibre with diameter greater
than 30
m (prickle factor), washing yield (scourging
yield), vegetable matter contamination, fibre strength
and position of staple break, presence of cotts and
116
E.N. Frank et al. / Small Ruminant Research 61 (2006) 113–129
tipiness, proportion of medullated fibres and resistance
to compression are also reported with some effects on
fibre value in alpacas (
Dehairing yield is another important variable take
in account for llamas fibre. This is because the aspect
of “hairiness” of the yarn and the knitwear is due to
the presence of “cover” fibres inside the staple and
its relative diameter to the “down” fibres (
). Nevertheless, the process
of dehairing is only carried out in extremes cases where
the benefits exceed the costs of processing (
2.4. Subjective measurements
Although white is the most prevalent colour,
camelids fibre has a wide range of colours. This may
be a problem if colourless fibre (i.e. without pigmen-
tation) is used or when a specific colour is desired. In
recent years there has been an increase for the pref-
erence of textile fabric with natural colours in ecola-
bel context (
). The most desirable fibre
colours are: white, black, reddish brown and golden
(
). A problem that often arises is lack
of colour uniformity produced by the mix or dilution
within the fleece, or in some cases, by fibre colour
contamination during harvest or packing of fleeces
(
It is important to note that even though fleece type
affects greatly value, this characteristic is often not con-
sidered important due to the difficulty to appreciate it
in the top (
). Currently, market data do not
show a relationship between type of fleece and value for
Suri alpaca (
). However, the need for dehairing
or not, may become a routine criteria of classification
in the near future.
The crimp of fibre (crimp absence or presence and
type) and the tip of staple have a direct relationship with
the type of fleece, in both alpaca and llamas. In Argen-
tinean llamas, three basic standard types of fleece have
been established: double coat fleeces; non-lustre sin-
gle coated fleeces and lustre single coat fleece (
(
Environmental damage in open pigmented fleeces
(specially the effect of the sunlight) is also taken into
account in the evaluation of the fleece, because it affects
fibre color (pigmented fibre) and fibre lengths (white
and pigmented fibres) (
Fig. 2. Type of fleece from Argentine llamas. DC: double coat fleece;
IC: intermediate coat fleece; SC: single coat fleece; L: lustre fleece;
HL: hemilustre fleece. Source:
3. Fibre production background
3.1. Commercial characteristics
3.1.1. Mean fibre diameter and fibre diameter
variation
The mean diameter of Argentinean lama fibre is
between 24 and 34
m (
). In the Cata-
marca province the non-adjusted mean fibre diameter
was 26.7
± 2.0 m (
When data was pooled from different locations of
the country, the adjusted least square mean of fibre
diameter was 25.2
± 0.2 m (
). A recent study in the NW area of Argentina
altiplano on 3726 lamas, the mean fibre diameter
adjusted for age, interval between shearing, type of
fleece, color and locality was 22.9
m which is sim-
ilar to those obtained in other areas of Argentina. The
frequencies of the different fibres classes was: baby
(<19
m): 4.6%; superfine (19–21.9 m): 43.4%; fine
(22–24.9
m): 35.8%; medium (25–29 m): 13.6%
and strong (>30
m): 2.6% (
In Peru,
reported a mean fibre diam-
eter of 23.8
m for Suri alpacas and 24.02 m for
Huacaya alpacas. The mean diameter for Kcara and
Chaku llamas was 33.8 and 28.06
m, respectively
(
). However, these studies did not adjust
for animal age, sex, color and locality.
shows
E.N. Frank et al. / Small Ruminant Research 61 (2006) 113–129
117
Table 2a
Factors influencing alpaca mean fibre diameter
Factor
Effect
Authority
Age
Increases with age
Del Carpio and Bustinza (1989)
,
Origin
Diverse
Sex
There was no effect
Breed or type of fleeces
Suri coarser than Huacaya
Fibre colors
Black finer than white
Ruiz De Castilla and Olaguibel De Olivera (1991)
Light shade finer than dark
There was no effect
Year of production
Diverse
Ruiz De Castilla et al. (1992)
Nutritional conditions
High level increase diameter
Seasonal variations
Winter decrease diameter
a summary of the factor influencing the fibre diameters
of alpacas.
In Bolivia, the mean fibre diameter, the diameter of
coarse and the diameter fine fibres were 31.6, 40.8 and
25.5
m, respectively (
). Others
have reported a mean fibre diameter of 27.2
m for
all fibres, and 22.3
m for fine fibre, in a herd of ani-
mal bred for meat (
Delgado Santiva˜nez et al., 2001
In young llamas (1st shearing at 21 months),
reported a mean diameter of 22.7
m for fine
fibres and 66.39
m for coarse fibre in a dehairing trial.
In southern Bolivia (near the border with Argentina),
reported a llama herd average fibre
diameter of 21.2
m.
In New Zealand alpacas, the best linear unbiased
mean fibre diameter reported using different measure-
ment methods ranged from 28.0 to 31.9
m (
). In Australia, a mean fibre diameter
of 29.1
m (17.7–46.6 m) was obtained in alpacas
(
shows a summary of factors influencing
the fibre diameters of llamas.
studied fibre diameter variation in different cut
points of the staple, between fibres in the same point of
the staple, between different topographic regions in the
same animal and between animals. Most of the varia-
tion was found between fibres within the staple while
other sources of variations were not very significant.
The coefficients of variation (CV) of fibre diameter
in two studies were 31.0
± 1.55% (
) and 31.7
± 0.25% (
). The
type of fleece is a very important source of diameter
variation. The corrected general mean CV obtained was
26.7
± 0.27% (
reported a mean
fibre diameter variation of 24.33% in Australian
alpacas (range: 15.0–36.7%). The CV was significantly
affected by age of animal (positive correlation), type
of fleece (higher in Suris) and color of fibre (higher in
darker fleece).
Table 2b
Factors influencing llama mean fibre diameter
Factor
Effect
Authority
Age
Increases with age
Rodr´ıguez and I˜niguez, (1977)
et al. (1985a)
,
Origin
Diverse
Breed
Kcara and Intermediate coarser than Wooly
There was no effect
Fibre colours
Black finer than white
Interval of shearing
Increase with annual shearing
Year of production
Diverse
Ruiz De Castilla et al. (1992)
Type of fleece
There was no effect
118
E.N. Frank et al. / Small Ruminant Research 61 (2006) 113–129
3.1.2. Colour
The need for fleece classification by colour has led
to the development of different types of charts similar
to those used for natural fibres (
Calle Escobar, 1982; Frank et al., 1991; Patthey Salas,
1994
). Other researchers have used colorimetrics stan-
dard charts (i.e. Munsel Soil Chart) (
and Maman´ı, 1990; Renieri et al., 1991
). The first stud-
ies on coat colour distribution in Argentinean herds
were conducted in Catamarca by
. These authors observed a predomi-
nance of brown coat color.
In a detailed study of 3200 animals throughout
Argentina,
reported a wide
variation of colour frequency by region. Most areas
studied showed a high frequency of white (B) except
for Catamarca, where many variants of brown (T: pure
tan; Mr: pure reddish brown; Mrc: combined reddish
brown) were found (
In a recent study on 3736 llamas from NW regions
of the Argentine altiplano, white colour was the most
commonly found (38%) but there was an increase in
brown or tan colours (30%). Frequencies were similar
for other commercial colours (
These findings are similar to those reported in Peru-
vian alpacas, where the frequency of white animals is
high (50–56%) and the brown or coffee color repre-
sent about 30% (
). In Bolivia, both alpacas
and llamas show a lower frequency of white animals
(8–15.8%), and a higher frequency of brown or coffee
(32–33%). Nearly 45% of Bolivian llamas are spotted
(
3.1.3. Type of fleeces
Originally, llama fibre was considered in scientific
literature as a double-coated fleece (
Riera, 1969; Calle Escobar, 1982
). However, research
in Argentina showed variability in type of fleeces
(
). Similar findings were
reported in llamas from southern Bolivia (
Also, regional and individual differences in the
distribution of frequencies of type of fleeces were
observed with a predominance of single coated fleece
and lower frequencies of double coated and lustre
fleeces in Argentine llamas (
Type of fleeces distribution in the NW area of
Argentina altiplano show a high frequency of single-
coated fleeces (46.6%). The frequency for lustre fleeces
is very low (3.3%). The frequency for double-coated is
intermediate (22.4%). The frequency of intermediated-
coated and hemilustre fleeces were respectively 9.6%
and 18.2% (
). This distribution of llama
type of fleeces confirms archaeological studies that
suggested the existence of llamas with different type
of fleeces. In the old literature these differences were
attributed to species differences (llamas versus alpacas
versus llama–alpaca crosses) (
The existence of different type of fleece has also
been described in Peruvian alpacas (
1959, 1991; Calle Escobar, 1982
) and is similar to
variations seen in Angora goats (
). Older
studies on alpacas reported a frequency distribution
of 70.6% Huacaya and 29.3% Suri (
however recent observation place these ratio at 95%
Huacaya and 5% Suri (
). In Peruvian
llamas there is a predominance of Kcara or dou-
ble coated type (80%) (
). In Bolivia, the
Huacaya and Kcara types have a similar frequency
(
3.1.4. Fibre length
The staple length is estimated from fibre length.
reported a correlation of 0.98
between average fibre length and staple length in llama
fleeces. Studies of llama fibre from the Altiplano of
Argentina found that the staple length of 80% of sam-
ples was of sufficient length to enable worsted pro-
cessing with the remainder used for woolen spinning
(
). The mean staple length from
typical harvesting systems includes fibre with more
than 12 months growth. Fbre from Jujuy province has
a non-adjusted mean staple length of 19.1 cm and an
adjusted 12-month growth period mean staple length
of 15.5 cm (
Frank et al., 1999; Frank, 2001a
).
reported a mean staple length of 13.00 cm
in 1-year-old Bolivian llamas and a mean length
of 16.75 cm for sheared animals from different age
classes.
In alpaca studies, staple lengths of 12.6 cm for Hua-
cayas and 16.8 cm for Suris was reported in Peru
(
). The staple lengths in alpacas from
New Zealand was 9.9
± 0.2 cm (
Australian alpaca staple length was 9.4
± 0.5 cm and
7.7
± 0.7 cm for two different consecutive produc-
E.N. Frank et al. / Small Ruminant Research 61 (2006) 113–129
119
tion years (1996 and 1997), respectively (
In wool, a variation (decrease or increase) of 10 mm
in average fibre length in the top equates to the same
spinning performance as a variation of 1
m in diame-
ter (
3.1.5. Dehairing yield
Dehairing is the removal of coarse fibres (hair
or “guard hair”) which grow from the primary skin
follicles. Depending on the type of fleece, these fibres
may be longer than the finer secondary fibres (
(
). Dehairing is also being developed
in alpaca fibre, but there is no published data on this
process.
Dehairing trials of llama fleece, developed in
Argentina, give a 35–50% yield of very good quality
down. The process is accomplished with a depuration
machine, which uses air to separate the finer fibre,
coarser fibre and contaminants (
).
In Bolivia, hand-dehaired llama fibre yields lower
results (
). Machine dehairing is
carried out in Bolivia but date is not yet available in
literature.
3.2. Fibre production
Fibre production is expressed as the weight of
raw fibre harvested per animal sheared and rep-
resents a certain value. The value of the fleece
is established based on the characteristics previ-
ously described. Domestic South American Camelid
fleece is lighter compared to those obtained from
selected sheep, but is similar to that of other fibre-
producing animals.
shows a summary of greasy
fleece weight of llamas and alpacas from different
countries.
4. Genetics of fibre characteristics
From a genetic point of view, fibre production traits
can be subdivided into quantitative traits, and factorial
or Mendelian traits. The most important commercial
Table 3
Means and ES of greasy fleeces weight measured in alpacas and llamas in different countries
Species
Country
Fleece weight
Authority
Llama
Argentina
1.39
± 0.2 kg (adult)
1.89
± 0.7 kg (hogget)
1.54
± 0.6 kg
1.61
± 0.2 kg
Bolivia
Wooly: 1.13 kg (hogget); 1.4 kg (adult); 1.5 kg (old)
Kara: 0.9 kg (hogget); 1.1 kg (adult); 1.1 kg (old)
At peasants level: 0.7 kg
At exp. station: 1.1 kg
Flock average: 1.2 kg
Peru
Wooly: 1.3 kg (hogget)
Intermediate: 1.02 kg (hogget)
Karas: 0.97 kg (hogget)
Flock average: 1.01 kg
Alpaca
Peru
0–2 Years: Huacaya: 1.4 kg, Suri: 1.4 kg
3–8 Years: Huacaya: 1.8 kg, Suri: 2.0 kg
>8 Years: Huacaya: 1.7 kg, Suri: 1.9 kg
New Zealand
Crias: 1.97
± 0.07 kg
Tuis: 3.02
± 0.2 kg
Adult: 2.16
± 0.06 kg
Australia
1996: 3.01
± 0.6 kg
1997: 2.09
± 0.4 kg
120
E.N. Frank et al. / Small Ruminant Research 61 (2006) 113–129
traits (fibre diameter, fleece weights and staple length)
are quantitative traits.
Coat colour and types of fleece are factorial traits,
they are less important for fibre production but have
some visual importance. The study of colour genetics
starts with a definition and description of phenotypic
patterns and their variations (especially of the white
spotted animals) (
). The study of
fleece types is more difficult because phenotypic vari-
ations are sometimes less obvious (
4.1. Phenotypical description of coat colours
patterns
Color phenotype description for llamas and alpacas
has been adapted from other species (
). Direct visual appraisal of color phenotypes is
carried out with the help of a colour chart such as the
Munsell’s chart. But this description is no very accu-
rate and must be supported by objectives biochemical
laboratory methods that determines the concentration
of eumelanins and pheomelanins (
Spectrophotometric assay of alcali-soluble melanin,
also allows the analysis of the brown eumelanins con-
tent (
Ito et al., 1993; Frank, 2001a
). Transmitting
electronic microscopic (TEM) also provides informa-
tion since it can differentiate eumelanosomes from the
pheomelanosomes and from intermediate types. TEM
can help in the description of colour phenotypes which
are difficult to appraise visually in alpacas (
The four-dimension coat colour pattern description
method (
) allows the description
of the phenotype shown in
. The major difficulty
in the description of the colour phenotype is to be able to
distinguish between predominantly pheomelanics and
brown eumelanic coats. This problem was solved by
the development of alkali-soluble technique (
The presence of brown eumelanins in Argentinean
llamas was verified by the detection of visible alter-
ations in the eumelanosomes using the TEM (
). These alterations were similar to those present
in alpacas skin (
). Molecular genet-
ics techniques may help refine these techniques. Some
genes related to the tyrosinase have already been
sequenced in alpacas (
patterns distribution (
) as well as white spotting
Fig. 3. Color pattern (a) and white spotting (b) schemes in Argentine
llamas. Dark: eumelanic; Light: pheomelanic or mixes; uncolored:
white. (a) Coat color patterns: (1) eumelanic (Eu); (2) black and tan
(B&T); (3) black face and extremities (BL); (4) mule strip (MS); (5)
badger face or black belly (BF); (6) wild (vicu˜na left and guanaco
right) (W); (7) pheomelanic (Ph). (b) White spotting design: (1) mark
(M); (2) irregular spotting I (IS); (3) irregular spotting II (IS); (4)
full white (FullW); (5) regular spotting I (RS); (6) regular spotting II
(RS); (7) spotted design (S). Sources:
. (For interpretation of the references to color in this figure
legend, the reader is referred to the web version of this article.)
pattern (
) can be determined with precision using
these above described techniques.
4.2. Colour phenotype segregations
Segregation studies of full white (non-albino)
phenotypes, irregular spotting and some pigmentary
patterns (black face, eumelanic black and wild) were
carried out in controlled llama flocks from Argentina.
Two hypotheses of recessive-dominance were put
E.N. Frank et al. / Small Ruminant Research 61 (2006) 113–129
121
Fig. 4. Coat color patterns frequencies distribution in the NW area of
Argentina altiplano. Eu: eumelanic pattern; B&T: black and tan pat-
tern; BL: black face and extremities pattern; MS: mule strip pattern;
BF: badger face pattern; W: wild pattern; Ph: pheomelanic pattern;
NN: non-identifiable patterns. Source:
forward for each analyzed phenotype (
These studies showed that the absence of pigmen-
tation (full white) is a dominant trait with incomplete
penetrancy to all pigmented patterns and white spot-
ting. Some colour patterns (i.e. brown with black face
and extremities) are dominant to other color patterns;
the eumelanic pattern (black or brown) is recessive to
all the others pigmentary patterns. These inheritance
patterns are summarize in
. Pigmented pheno-
types are segregated by the locus Agouti (
Frank et al., 2001b, 2002; Renieri et al., 2002
). There
were no consistent results for black and brown eume-
lanins segregation as well as some pigmentary patterns
named mule strip, badger face and black and tan. Like-
wise, the dominance patterns between wild guanaco
and wild vicu˜na phenotypes are unclear.
Fig. 5. White-spotting distribution in the NW area of Argentina alti-
plano. M: small white spotting; IS: irregular spotting; FW: full white
spotting; RS: regular spotting; S: spotted design (paint); NW: without
white spotting. Source:
4.3. Phenotypical description of type of fleece
From the phenotypical point of view, three types of
fleeces are identified: pattern types; double coat, sin-
gle coat and lustre. Two intermediate types are also
described; the hemi lustre which is an intermediate type
between single coat and lustre type and the intermedi-
ate coat which results from the double coat and single
coat (
shows
different criteria used to describe type of fibres. The
fibre types are described based on fibre length, fibre
thickness, types of waves or crimps (sinusoidal, heli-
coids, twisted sinusoidal) and presence or absence of
lustre. These macroscopic observations are validated
by microscopic evaluations of the types of cuticular
scales, fibre and medulla shapes. These variables are
related to follicle types from which the fibres emerge
(
The relationship between the fibre diameter pro-
duced by the primary and secondary follicles deter-
mines the presence or absence of double-coated fleeces
(
). The shape and number of cuticular
scales allow a distinction between lustre fleece and non-
lustre fleece when fibre casts are examined (
). The thickening of the follicle inner root
sheet allows a differentiation between the lustre and
non-lustre fleece (
). Lustre fibres are gen-
erated by follicles with thicker inner sheet as described
in sheep with “dogginess” wool (
The Freney effect (
), which
cause a marked crimp under the bilateral cortex effect
helps to differentiate the lustre (absence of bilateral
symmetry) from the non-lustre (presence of bilateral
symmetry) fibre in llamas and alpacas (
Fleece types present differences in productivity and
textile behavior. In alpacas, Suri (lustre) type has longer
fleece (
). In Argentinean llamas, lustre and
semi-lustre fleece types have longer staple length than
the double coated fleece type (
). No differences
in fibre diameter and fleece weight were found among
the type of fleeces.
The textile quality is affected by the type of fleeces.
Th double coat presents a greater variation of fibre
diameter in the different body parts (
Dehairing is more noticeable in double-coated fleeces
than in the lustre type, and it is intermediate in sin-
gle coated and intermediate coated. This mechanical
122
E.N. Frank et al. / Small Ruminant Research 61 (2006) 113–129
Table 4
Summary of mating scheme tested in relation to color genetics hypothesis
Phenotips
Phenotypic and Genetic hypotheses
Segregation scheme
Uniform white non-albino (UW)
White dominant
Uniform white (UW) vs.
pigmented non-spotting (P)
UW
× UW
UW
× P
P
× P
White spot (S)
Irregular spot extension
Pigmented non-spotting (P)
vs. irregular spotting (IS)
Presences as recessive vs. Spot absence
P
× P
P
× IS
IS
× IS
Pigmentation models
Black face and extremities (BF) vs. eumelanic (Eu)
Eumelanic as recessive vs.
BF
× BF
Black face and extremities (BF)
BF
× Eu
Eu
× Eu
Black face and extremities (BF) vs. wild (W)
Unknown
W
× W
W
× Eu
Eu
× Eu
Results of the Mendelian hypothesis tested
Segregations
Tested relations
Mating schemes
Tested hypothesis
Statistic significance
P > 0.05
Results of genetic
relations
UW vs. IS
UW dominant to IS
UW
× UW
3:1
ns
Accepted
UW
× IS
1:1
ns
UW vs. P
UW dominant to P
UW
× UW
3:1
ns
Accepted
UW
× P
1:1
ns
BF vs. W
BF dominant to W
BF
× BF
3:1
ns
Accepted
BF
× UW
1:1
ns
BF vs. Eu
BF dominant to Eu
BF
× Eu
1:1
ns
Accepted
Source:
a
If penetrancy is considered in heterozygote.
Fig. 6. Effect of type of fleeces on staple length in a llama flock.
DC: double coat fleece; IC: intermediate coat fleece; SC: single coat
fleece; L: lustre fleece; HL: hemilustre fleece. Source:
operation (dehairing) reduces the prickling effect in the
double coat and its effect is less noticeable on the single
coated and lustre types (
Lustre types shows less bulk in the fibre and fabric
than the non-lustre type. The tactile attributes evaluated
in the fabrics show that lustre types are softer, with a
more regular or smooth surface and less tactile warmth.
Dehaired double-coated types have similar (low) prick-
ling effect than lustre types (
4.4. Type of fleeces segregation
The double coat trait has been characterized as
partially dominant to the single coat in Peruvians
llamas (
). In alpacas, studies in
E.N. Frank et al. / Small Ruminant Research 61 (2006) 113–129
123
Table 5
Simplified schemes for type of fleece classification by visual assessments
No. type
Intermediate coated
Double coated
Hemi Lustre
Lustre
Single coated
1
1163
2
1253
3
2163
4
2253
2253
2253
5
2343
6
2352
7
2353
2353
8
2411
9
2412
10
2413
11
2452
12
2453
13
2511
14
3243
15
3323
16
3331
17
3333
3333
18
3343
19
3412
20
3413
21
3421
3421
3421
22
3422
3422
3422
23
3423
3423
3423
24
3432
Criteria of classification of type of fibre
Intermediate coated
Double coated
Hemi Lustre
Lustre
Single coated
Length: 1
◦
1. Short fibre
X
2. Medium fibre
X
X
X
X
3. Long fibre
X
X
X
X
Thickness: 2
◦
1. Very fine fibre
X
X
2. Fine fibre
X
X
X
X
X
3. Medium coarse fibre
X
X
X
X
X
4. Coarse fibre
X
X
X
X
X
5. Very coarse fibre
X
Type of waves: 3
◦
1. Completely straight fibre
X
X
2. Straight-big waved fibre
X
X
X
3. Big waved and helicoidally fibre
X
X
4. Big waved helicoidally combined with
some small sinusoidal waves
X
X
5. Small sinusoidal or helicoidally wave
with some big sinusoidal wave fibres
X
X
X
X
6. Small sinusoidal and closed wave fibres
X
X
Brightness: 4
◦
1. Opaque or chalky fibre
X
X
X
X
X
2. Opaque-glassy fibre
X
X
X
X
3. Glassy fibre
X
X
X
X
X
An example:
Type of fibre: 1163
1. Short fibre; 1. very fine fibre; 6. small sinusoidal and
closed wave fibres; 3. glassy fibre
Source:
a
Type of fibre more frequent within each type of fleece.
124
E.N. Frank et al. / Small Ruminant Research 61 (2006) 113–129
Peru (
et al., 1997
) have shown that the lustre type (Suri) was
dominant to the non-lustre type (Huacaya). It is impor-
tant to note that precise distinction between fleece
types requires dissection of staples, fibre analysis
and skin analysis to identify follicles characteristics
(
). This is probably why results
obtained from studies on phenotypic evaluation of
fleece types are generally confusing. Segregation
studies in Argentine llamas demonstrate that Lustre
type is dominant to non-lustre types with a penetrancy
effect that allow that some heterozygotes show an
intermediate type of fleece (between single coated and
lustre), and double coated is probably governed by an
additive genetic mechanism (
4.5. Genetic parameters of quantitative traits
The fleece structure is basically determined by fibre
length, fibre diameters and their inter relationship. A
great part of the fleece structure is determined by
the type of staple but it is also influenced by quan-
titative traits. The genetic determinism of these traits
corresponds to additive multi genetic inheritance types
described in llamas and alpacas.
Preliminary data on alpacas in Australia and New
Zealand indicate a high to very high heritability of
fleece weight, fibre diameter and staple length (
et al., 1999; Wuliji et al., 2000
). Data from Peru
place these heritabilities to relatively high or medium
Table 7
Bibliographic heritabilities estimated of llamas
Characteristics
Parameter
References
Fleece weight
0.48
± 0.02
0.27
Staple length
0.34
± 0.08
0.28
± 0.37
Fibre diameter
0.28
± 0.10
Frank, E.N. (unpublished)
(
) (
). Our preliminary data
obtained on Argentinean llamas suggest that the value
of heritability of most of these traits is medium.
Available phenotypic correlations indicate that there
is a significant negative correlation between fleece
weight and fibre diameter as described in sheep
(
Ponzoni et al., 1999; Wuliji et al. 2000
4.6. Non-genetic factors that affects production
Environmental effects are the most important non-
genetic factors which affect fibre production in domes-
tic South American camelids.
classified them as permanent or internal and temporary
or external effects.
In alpacas, fleece weight is influenced by sex
(2.4%), locality (32.6%) and age (6.5%) (
). Mean fibre diameter is affected by
age, sex, origin, breed, fibre colours, year of produc-
Table 6
Bibliographic heritabilities estimated of Huacaya alpacas:
Traits
Age
Parameter
References
Fleece weight
1
◦
Shearing
0.35
± 0.02
0.22
0.21 + 0.07
0.38 + 0.34
0.31 + 0.17
Ruiz De Castilla et al. (1992)
All ages
0.79
± 0.36
0.63
± 0.22
Fibre diameter
0.18
Le´on-Velarde and Guerrero (2001)
(averaged)
0.67
± 0.30
0.73
± 0.19
Staple length
1
◦
Shearing
0.43 + 0.39
0.21 + 0.07
0.31
Le´on-Velarde and Guerrero (2001)
(averaged)
All ages
0.63
± 0.48
0.57
± 0.18
E.N. Frank et al. / Small Ruminant Research 61 (2006) 113–129
125
tion, nutritional conditions and variation of seasonal
conditions (
In Argentina,
found significant
effect of age on average diameter, coefficient varia-
tion of the diameter, staple length and medullation.
Fleece weight is significantly affected by geograph-
ical location (60.9%) and age (15.9%) (
When values were adjusted to annual shearing no effect
was observed on fleece weight. However, the inter-
val between shearing explains most of the variation
(47%) of mean fibre diameter. There was a highly sig-
nificant interaction between age and interval of shear-
ing (24%). In llamas, diameter of fibre increases for
the first 5–6 years of age then decreases thereafter.
Staple length is primarily affected by the interaction
between geographical location and frequency of shear-
ing (70.6%). It decreases with age without showing
a biphasic behavior as seen for the average diameter
and fleece weight (
). Similar effects
were studied in Bolivian llamas with same results
(
5. Conclusions
Traits that determine fibre quality in South Ameri-
can domestic camelids are mean fibre diameter, color,
type of fleeces, fibre length and uniformity of diameter
and length. The amount of fibre production per head is
low; however there are alpacas and llamas populations
that show excellent fibre quality.
The quality of alpacas and llamas flocks may be
improved by increasing the frequency of fleece that
would bring premium price in the textile industry. The
fleece should be white or even colored, fine, soft han-
dled and have low prickling. Luster (Suri alpacas) and
double coated fleeces after efficient dehairing process
naturally possess these attribute. Genetic improvement
may be achieved by selection program for specific traits
such as color phenotype and fleece type. More research
is needed to determine the relationship between quan-
titative traits (fleece weights, staple length and fibre
diameter) and environmental effects.
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Document Outline
- Phenotypic and genetic description of fibre traits in South American domestic camelids (llamas and alpacas)
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