conditions are more favorable in “La Raya”. But also, different management strategies could be responsible for these observed differences.

## Mortality

The average mortality rate during the evaluation period was 5.28 %, 5.45 % and 16.47 % for llamas, alpacas and sheep, respectively. Mortality was found to be higher in sheep than in camelids; however, most of the llama deaths occurred in the period 2013-2014, while in the period 2012-2013 there were more deaths of alpacas (Figure 3). Regarding mortality by age, a mortality of 2.47 %, 0.47 % and 2.34 % was found in offspring, ancutas (llamas between weaning and first mating) and adult llamas, respectively. In the case of alpacas, the mortalities found were 3.7 %, 0.67 % and 1.07 % in offspring, tuis (alpacas between weaning and first mating) and adults; while for sheep, mortalities of 11.28 %, 1.8 % and 3.39 % were calculated for lambs, young sheep and adults, respectively. The mortalities found for llamas in this study were lower than those used in the development of a simulation model of the llama population under environmental and market restrictions in the Bolivian highlands (Treydte et al., 2011) of 29 % and 6 % for juveniles (less than 2 years) and adults (greater than 2 years), respectively. The high rates of llama mortality in Bolivia are due to inadequate management, as well as lack of sanitary controls, such as feeding programs, which result in high incidences of diarrhea and malnutrition (Lopez, 2004, Whitehead and Anderson, 2006). Ruiz et al., (2015) report higher mortalities than those of the present study of 7.47 % and 6.94 % for two associations of alpaca farmers in the Pasco Region, shows that good pasture conditions and proper management of the animals result in lower mortality rates. On the other hand, it was observed that sheep had higher mortality rates than llamas and alpacas, demonstrating that camelids have some resistance to diseases such as pulmonary adenomatosis, pododermatitis and epididymitis, which are diseases with a high incidence in sheep, but are rarely seen in camelids (Sumar and Camino, 1992).

## Harvest

An average harvest of 21.11 %, 23.33 % and 23.64 % was determined for llamas, alpacas and sheep, respectively. Figure 4 shows that the largest amount of camelid harvest occurred in the period 2012-2013 and that the lowest harvest occurred in the period 2014-2015 with 4.99 %. In addition, it was observed that there is no defined month to perform the harvest. The percentage of llama harvest obtained in this study was higher than the 11.4 % found in Bolivia, which is attributed to low fertility and high mortality (FAO, 2005a).
<figure><img src="image_24.png" /></figure>

**Figure 3:** Mortality (%M) of llamas, alpacas and sheep from the Community Cooperative San Pedro de Racco from 2013-2015

<figure><img src="image_25.png" /></figure>

**Figure 4:** Harvest (%H) of llamas, alpacas and sheep from the Community Cooperative San Pedro de Racco from 2013-2015
Gross and real increase

It was found that llamas had an average GI greater (2.09 %), unlike alpacas with -2.05 % and 1.6 % for sheep (Figure 5). During the evaluation years the gross increase rates were the same as the real increase rates in each species. Except for the alpaca period 2012 - 2013, where the % GI was -10.75 and the % RI was -11.35 % since twenty born animals were not reported (Figure 6).

<figure><img src="image_26.png" /><figcaption>**Figure 5:** Gross increase (%GI) of llamas, alpacas and sheep from the Community Cooperative San Pedro de Racco from 2013-2015</figcaption></figure>

<figure><img src="image_27.png" /><figcaption>**Figure 6:** Real increase (%RI) of llamas, alpacas and sheep from the Community Cooperative San Pedro de Racco from 2013-2015</figcaption></figure>
Livestock Efficiency

In the 2012-2013 period, the highest LE was found for camelids, with 26.76 % for llamas and 25.61 % for alpacas, whereas for sheep a LE of 28.16 % was obtained in 2014. Livestock efficiency averages of 23.2 %, 21.08 % and 25.24 % were obtained for llamas, alpacas and sheep in the evaluated years (Figure 7).

<figure><img src="image_28.png" /><figcaption>**Figure 7:** Livestock efficiency (%LE) of llamas, alpacas and sheep from the Community Cooperative San Pedro de Racco from 2013-2015</figcaption></figure>

Table 1 shows all the technical parameters of the Communal Cooperative San Pedro de Racco during the evaluated years, finding that its management system has a medium to high technological level, according to Esponda et al. (2004). This could be a result of keeping records containing the complete information of the animals, such as calving, mating, mortality and harvest records (Condorena, 1987; Cruz, 2012). In addition, studies have shown that communal cooperatives in the Pasco Region exhibit adequate rangeland condition and trend parameters, adequate carrying capacity and forage balance (Barrantes, 2012; Zarria et al., 2012), which would explain the efficient management done by the Communal Cooperative San Pedro de Racco.

Regarding llama management systems in Peru, there have been no recent studies of performance evaluation, however Lopez (2004), reports a birth rate of 34.01 % and a mortality of 2.58 % for llamas in Oruro province, Bolivia.

In the case of alpacas, Palacios (2009) reported technical parameters of 70.94 %, 29.68 %, 18.56 %, 6.33 %, 4.75 % and 23.31 % for %GBR, %RBR, %H, %M, %RI and %LE respectively for Huacaya alpacas from the Mallkini Ranch (Azángaro, Puno) during the period 2001 to 2008.
<table><caption>**Table 1:** Average technical parameters from the Community Cooperative San Pedro de Racco from 2013-2015</caption><thead><tr><th>Technical parameter</th><th>Llamas (2012-2015)</th><th>Alpacas (2012-2015)</th><th>Sheep (2014-2015)</th></tr></thead><tbody><tr><td>%AAC</td><td>507.8</td><td>3086.2</td><td>4683.0</td></tr><tr><td>%GBR</td><td>46.0</td><td>60.0</td><td>66.0</td></tr><tr><td>%RBR</td><td>25.0</td><td>29.0</td><td>41.0</td></tr><tr><td>%M</td><td>5.3</td><td>5.4</td><td>16.5</td></tr><tr><td>%H</td><td>21.1</td><td>23.3</td><td>23.6</td></tr><tr><td>%GR</td><td>2.0</td><td>-2.0</td><td>2.0</td></tr><tr><td>%RI</td><td>2.0</td><td>-2.0</td><td>2.0</td></tr><tr><td>%LE</td><td>23.0</td><td>21.0</td><td>25.0</td></tr></tbody></table>

Likewise, Ruiz et al., (2015) reported average technical parameters of two associations of alpaca farmers in the Pasco Region during the years 2012-2014 of 47.1 %, 26 %, 7.2 %, 18.8 %, 0.2 %, 0.0 % and 19 % for %GBR, %RBR, %M, %H, %GI, %RI and %LE, respectively. This revealed that small community organizations can be as efficient as a company with a medium to high technological level. This can be mainly attributed to the capacity development of human resources.

On the other hand, Cayo (2001) reported 76.76 %, 35.76 %, 10.74 %, 24.37 %, -0.3 %, -0.51 % and 23.87 % for %GBR, %RBR, %M, %H, %GI, %RI and %LE, respectively for sheep Corriedale of Pachacutec Agricultural Society, located in Junin province (Central Peru). This data supports the findings of the present study because both, the farmers' associations and agricultural society use records and perform pasture improvement practices, which is reflected in their technical parameters.

## Conclusion

The Community Cooperative San Pedro de Racco presented values of technical parameters corresponding to a company with good technological level. These results might be attributed to the capacity development of human resources in terms of animal and grassland management.

**Acknowledgments.** This research was supported by KEF Project "Strengthening llama production in the Central Andes of Peru".
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<figure><img src="image_29.png" /></figure>
# The Camelid Registry LAREU: What Are We Breeding In Europe?

## C. Kiesling

Llama and Alpaca Registries Europe (LAREU), P.O. Box 666, CH-3900 Brig-Glis, Switzerland

E-Mail: christian.kiesling@lareu.org

**Abstract.** The Llama and Alpaca Registry Europe ("LAREU") is providing an online registration system for breeders and owners of South American camelids (SACs), free of charge. Founded in the year 2005, over 19,000 animals from more than 10 European countries are stored in LAREU's database by spring 2017, with a yearly growth of about 10 percent. In this article we will present statistical material from the LAREU database with the aim to provide some insight into the breeding goals of the European camelid owners. The main subjects of the data analysis are concerned with the distribution of the different types of alpacas (huacayas and suris) and of llamas (light, medium, heavy wool, and suri), as well as the distribution in the various color classes (for alpacas), according to the color code scheme defined by ARI. Information is also presented about the frequency of parentage for sires and dams, giving some idea about the fraction of "proven" reproducers.

**Resumen.** El Registro de Llamas y Alpacas de Europa ("LAREU") ofrece un sistema de introducción de información gratuito en línea para criadores y propietarios de camélidos sudamericanos (SAC). Tras ser fundado en 2005, la base de datos de LAREU en la primavera de 2017 contenía registros de más de 19.000 animales de más de 10 países europeos en la primavera de 2017, con un crecimiento anual aproximado del 10 por ciento. En este artículo se presentan estadísticas de la base de datos LAREU con el ánimo de sugerir objetivos de mejora a los propietarios de camélidos europeos. La información principal se refiere a la distribución de los diferentes tipos de alpacas (huacayas y suris) y de llamas (ligeras, medianas, pesadas de lana y suri), así como la distribución de los diversos colores de capa (para alpacas) de acuerdo con la clasificación de colores definida por ARI. También se presenta información sobre la el parentesco entre padres y madres, dando idea de la fracción de reproductores "probados".

**Keywords:** Alpacas, llamas, statistics, LAREU registry

## Introduction

With the increasing numbers of alpacas and llamas kept by European owners over the last two decades, also an increase in the number of national camelid associations spread over Europe was encountered. The increase of the latter was fre-
quently motivated by the associations’ diverging breeding goals, but also some-
times because of incompatibility in the views of the associations’ leaders towards
future directions and developments of the associations, let alone the difficulties
related to the different languages spoken in the associations within Europe. Almost
inevitably, every association established its own registration database, with no pos-
sibility of cross-linking to the databases of the other associations. Since neverthe-
less quite often the animals have been transferred between owners of different
associations, also across national borders, most of the “history” of the animals, in
particular the pedigrees, got lost. In addition, the different associations were usually
working with different DNA laboratories using different marker sets for DNA
typing, making a reliable parent verification across association borders impossible.

Given this unsatisfactory situation, LAREU was founded in 2005 as a truly Eu-
ropean registry, open to individual breeders and owners of South American came-
lids. The main idea of LAREU was to provide an internet-based registry without
human intervention from the side of the registry (except providing the registration
page and its software and working on improvements of the informatics part). An
additional requirement of LAREU was to keep the registration process free of
cost. This was made possible by an offer of the animal welfare organization TAS-
SO, based in Germany [TASSO, 2017], to host the LAREU database and to pro-
vide the computing infrastructure for servicing the LAREU registration webpages.
TASSO itself is an organization for registering and localizing run-away pets, with a
database of a few million animals, mostly dogs and cats. The data of the animals
registered with LAREU are entirely separated from the TASSO database, also the
LAREU registration software is being developed independently as a voluntary,
unsalaried project by members of TASSO and LAREU. In contrast to most other
registries, LAREU allows filling only the parents for each animal, which in turn are
(or should be) registered in LAREU as well. With this system unique ancestry trees
are possible, providing reliable statistics over many generations, in principle result-
ing in unlimited (“infinite”) pedigrees.

The transparent borders within the European Union are offering breeders of SACs great opportunities for improving the genetics of their animals, both with respect to fiber quality as well as to conformation and health. LAREU has recognized from the beginning the importance of a Europe-wide registry and of a worldwide standard of the DNA markers used for SACs. Upon initiative of LAREU the “International Society of Animal Genetics (ISAG)” developed in the years from 2008 until 2012 a unique marker panel for alpacas and llamas. LAREU is working together with three laboratories in Germany, France and the Netherlands, who have been part of the early research to establish the unique marker set. In the meantime this set is used by most DNA laboratories worldwide, so that the transfer of DNA data from other laboratories into the common LAREU database and their evaluation for parent verification is now becoming a meaningful routine. Clearly, the DNA typing is a service provided by a qualified laboratory, which is reimbursed for its services directly through the animal owner.
It should be stressed here that all animal data in the LAREU database are en-
tered by the animal owners. Also, there is no requirement for completeness of the
animal data at the time of registration. However, there are a few automatic con-
sistency checks, for example, the transponder number is cross-checked for unique-
ness. All missing information can be entered later, once available. For this reason,
the DNA information is, at this stage, far from being complete for all the animals
registered in LAREU. Nevertheless, the large numbers of animals already in the
database (more than 19,000 in spring 2017, increasing each year by more than
2,000) makes LAREU by now the largest database on SACs in Europe.

In the following, we will first show some statistics on the registration activity
and the numbers of animal registrations for those European countries with the
highest activities. We then turn to some interesting questions concerning the dis-
tribution of types of alpacas and llamas registered. For the case of alpacas, we pre-
sent distributions of the various colors bred in huacayas and suris, and show these
distributions for the four European countries with the largest numbers of regis-
tered animals. Then we turn to the data analysis with respect to possible trends
visible in the breeding program for color. Finally, some data related to possible
breeding strategies is shown, and a "demographic" statistics concludes the article.

Registration Activity and Country Share

<figure><img src="image_30.png" /><figcaption>**Figure 1:** Time development of the animal registrations in the LAREU database. The registry was founded in 2005, registrations were started in 2006. The numbers shown on top of the figure reflect the status of the registration in April 2017.</figcaption></figure>

During the year of foundation in 2005, the initial code of the LAREU registry was
developed. First registrations were possible in the beginning of 2006. During the
first years, the numbers of llamas and alpacas registered were quite similar, the
llamas were dominating until the year 2008. From then on the alpacas became more and more popular and their numbers increased steadily, overtaking the llamas. Today there are more than double the number of alpacas registered compared to the llamas. The registration activities over the years are shown in fig. 1, separated into llamas and alpacas, until April 2017. At this point almost 20,000 SACs were registered, out of them over 14,000 alpacas and more than 5,600 llamas. While the rise of the llama registrations appears almost linear since about 2010, the increase of the alpacas is following a rate of about 10 % per year. Since the beginning of 2013, slightly more than four years ago, the number of registered animals has doubled. If the trend holds, one can expect another doubling by the year 2022.

<table><caption>**Table 1:** Breakdown of animal registrations in the various European countries, ordered by the numbers of animals. Also given are the numbers of registered owners and the average numbers of animals registered per owner.</caption><thead><tr><th>Country</th><th>Animals</th><th>Fraction (%)</th><th>Alpakas</th><th>Llamas</th><th>Owners</th><th>Animal/Owner</th></tr></thead><tbody><tr><td>DE</td><td>5967</td><td>30.0</td><td>4872</td><td>1095</td><td>774</td><td>7.7</td></tr><tr><td>FR</td><td>5812</td><td>29.2</td><td>3362</td><td>2450</td><td>1096</td><td>5.3</td></tr><tr><td>CH</td><td>1330</td><td>6.7</td><td>1110</td><td>220</td><td>151</td><td>8.8</td></tr><tr><td>IT</td><td>1008</td><td>5.1</td><td>479</td><td>529</td><td>109</td><td>9.2</td></tr><tr><td>AT</td><td>668</td><td>3.4</td><td>470</td><td>198</td><td>138</td><td>4.8</td></tr><tr><td>NL</td><td>545</td><td>2.7</td><td>520</td><td>25</td><td>73</td><td>7.5</td></tr><tr><td>BE</td><td>358</td><td>1.8</td><td>334</td><td>24</td><td>73</td><td>4.9</td></tr><tr><td>FI</td><td>188</td><td>0.9</td><td>179</td><td>9</td><td>43</td><td>4.4</td></tr><tr><td>SE</td><td>140</td><td>0.7</td><td>66</td><td>74</td><td>23</td><td>6.1</td></tr><tr><td>CZ</td><td>90</td><td>0.5</td><td>35</td><td>55</td><td>18</td><td>5.0</td></tr><tr><td>others</td><td>417</td><td>2.1</td><td>350</td><td>67</td><td>23</td><td>18.1</td></tr><tr><td>no owners</td><td>3369</td><td>16.9</td><td>2426</td><td>943</td><td></td><td></td></tr><tr><td>Sum</td><td>19892</td><td></td><td>14203</td><td>5689</td><td>2583</td><td>7.7</td></tr></tbody></table>
<figure><img src="image_31.png" /><figcaption>**Figure 2:** Breakdown of types for alpacas and llamas, taking into account all registered animals since the start of the LAREU registry. The type “suri” exists for both species, both alpacas and llamas show similar fractions of suris.</figcaption></figure>

A breakdown of the registrations (status of April 2017) into the individual European countries is shown in Table 1, ordered in descending numbers of registered animals. The most active countries are Germany and France, who have registered about 30% each of the total number of animals in the LAREU database, followed by Switzerland, Italy and Austria. The Netherlands and Belgium have also registered substantial numbers. There is also a relatively large fraction of animals not attributed to any country (“no owners”). These animals have been transferred from their original breeders to other owners who do not (yet) register their animals with LAREU. The transfer of an animal to a new owner is initiated by the previous owner: After the transfer all animal data, including the pedigree, are visible in the account of the new owner and have disappeared from the list of animals of the previous owner. If the new owner is not registered with LAREU, the animal data are transferred to a special “no owner” area. Once the new owner registers with LAREU, the animal data can be retrieved from the “no owner” area.

## Gender and Type Distributions

The different types of wool for alpacas and llamas are shown in fig. 2. While for the alpacas only two types exist (huacaya and suri), the llamas are traditionally separated into four classes. In recent years, the “classic llama” has become more rare and is being overtaken by the wooly type, at least one can observe this in the registration activity, but also at shows. A similar situation can be found for the suris, which have seen a relative increase in numbers with respect to the total number of SACs registered with LAREU.
<figure><img src="image_32.png" /><figcaption>**Figure 3:** Breakdown of color classes for huacayas and suris. The similarity of the relative fractions within huacayas and suris is remarkable.</figcaption></figure>

While the fraction of suri is pushing close to 10% for the llamas in recent years, the fraction of suri alpacas remained pretty constant over time, as shown in Fig. 2.

Another interesting piece of data is the breakdown into gender, namely males, castrated males ("geldings"), and females. The fraction of geldings is small, but with a significant difference between llamas and alpaca. While only 3.9% of geldings are registered for alpacas, a much larger fraction of 8.4% is registered for llamas. Possible reasons for the larger fractions of geldings in llamas might lie in their use as companion animals in various recreational activities, such as trekking, or also in animal-assisted activities and therapy. Correspondingly, there is a difference in the registrations of males: 37.9% for llamas and 40.3% for alpacas, while the fraction of females is 55.8% for the alpacas and 53.7% for the llamas.

We now turn to the statistics concerned with the fiber which is an important marketing subject for the alpaca owners. Since wool characteristics related to the fiber diameter and comfort factor are usually dependent on the animal age and may vary substantially during the animal's lifetime, these quantities are not stored in the LAREU database and cannot therefore be used here in a statistical analysis. Instead, we concentrate on the aspects of fleece color, which can be more reliably judged by the animal owner and is therefore a mandatory item in the LAREU database. For the various color shades we use the classification of ARI [AOA, 2014], which exhibits 16 natural colors and 3 additional multi-color classes. From these 19 classes for the alpacas we have summed up the main color traits, as shown in Fig. 3 (the fiber of llamas is not of so much importance as quality indicator, mainly due to the higher average fiber diameter, but also due to the large variation of color shades in the llamas). While the colors white and beige are unique, the colors
brown and fawn are divided into three further classes each: light, medium and dark. The grey color is divided into two main classes, rose grey and silver grey. These three are again subdivided into light, medium and dark. Finally, the color black is divided into true black and bay black. All subdivisions are summed up into the respective color classes shown in this and the following figures.

Comparing huacaya and suris shows a remarkable similarity in the relative fractions of colors. In both types the color white is dominating, for obvious reasons, since the early European alpaca population was imported from wool producing countries where emphasis is put on easy-to-stain fiber. Fawn and brown follow, black is still around 13-15 %, the grey shades, multi-color and beige are very similar, adding up to less than 20 % in total.

While the data in Fig. 3 are summed over all countries, it may be interesting to see whether there are any national differences in the color fractions. This is shown in Figure 4 for the huacaya types, displaying the four countries with the largest number of registrations, namely Germany, France, Switzerland and Italy (see also Table 1). From these data one finds that there are indeed differences among the four nations. The color white dominates in Italy, but is least prominent among the four in Switzerland. Switzerland, on the other hand, has the largest fraction of grey huacayas, whereas Germany and France have much less. Switzerland dominates in the black colors, and Italy has the largest fraction of brown animals. The largest fawn community one finds in Germany, Italy has the smallest fraction of fawn animals of the 4 countries. Since more detailed information is missing, it is not possible to reliably explain these regional differences. But it seems plausible that the differences have to do with the regional market situation and with the preferences of the animal owners and the intended use (wool, companionship etc.) of the animals.

The national differences observed in the color distributions of Fig. 3 may point to some possible trends in the selection of alpacas, or in other words, reflect the “taste” and the “preferences” of the animal owners. In order to address this interesting question, the various colors were counted starting on January 1, 2014, and were compared with the distribution before that date.
<figure><img src="image_33.png" /><figcaption>**Figure 4:** Breakdown of color classes for huacayas, comparing the four European countries with the largest numbers of registered alpacas. Significant differences are observed in the relative fractions between the countries.</figcaption></figure>
<figure><img src="image_34.png" /><figcaption>**Figure 5:** Breakdown of color classes for huacayas, comparing the number of registrations before the year 2014 with those afterwards. January 1, 2014 marks the date when the half of the present animals in the database have been registered.</figcaption></figure>

The significance of the date (January 1, 2014) comes from the fact that the same number of animals was registered before and after this date. This means that this date defines the median in the temporal frequency of registrations (see also Fig. 1, where the median is indicated). The result of the color count is shown in Fig. 5, where all European countries are summed up in order to see a general trend in the data. Here one observes that the color white is becoming less popular in the recent years, while the “colored” animals are increasing in fractions, most significantly black, grey, multi-colored and beige. This trend may indicate that the emphasis is slowly shifting from “finest fiber” (mostly found in white animals due to selection) towards “companion animals”, which are pleasing the eye and are more attractive.

## Sires, Dams and Offspring

As mentioned in the beginning, the possibility to construct an “infinite pedigree” is one of the attractive features of the LAREU registry. The principle is shown again in Fig. 6: Each animal is linked to its parents (and only to its parents), and the animal information including the color type and the DNA typing (if existing) is displayed. Parent verification is indicated as well by adding the affixes “FV” for “father verified”, or “MV” for “mother verified” (see arrow in Fig. 6). This information is filled into the database by the DNA lab and cannot be changed by the animal owner. The ancestry tree can be followed (on the web-browser) by
clicking on an animal within the tree, which then lists the three ancestry generation of the animal clicked.

The DNA information is not mandatory at the time of registration, but can be added afterwards by ordering DNA typing and parent verification from one of the DNA laboratories working with LAREU at any convenient time (import of previous DNA results into the database, launched by the DNA lab, is also possible). Despite all this, the number of DNA-verified animals is still small in the LAREU database. This is shown in Table 2: A total of 858 (2,643) alpaca sires (dams) and 474 (1,298) llama sires (dams) are entered in the database. The alpaca (llama) sires have 4,215 (2,939) offspring, the corresponding numbers for the dams can be found in the table. The table also shows clear indications for the frequent use of “proven” sires and dams: For example, in the alpaca sires 50 % of the offspring are fathered by only 127 sires, 33 % by only 62 sires (similar numbers for the llamas, see also the corresponding numbers for the dams of both types). The maximum number of offspring for the “most productive” sires is 54 for the alpaca and 84 for the llama. The most productive dams have 11 (13) offspring for the alpacas (llamas). One should note, however, that these numbers are entered by the owners, there is no unique proof about the parentage. The last row of each of the tables gives the truly verified parentage: Here one finds that only about 10 % of the sires, dams and their offspring are verified by DNA at present.

<table><caption>**Table 2:** Statistics of DNA-typed sires and dams and their verified offspring in the LAREU database in alpacas and llamas</caption><thead><tr><th></th><th>Offspring</th><th>Sires</th><th>Offspring</th><th>Dams</th></tr></thead><tbody><tr><td rowspan="3">Alpacas</td><td>4215 (100%)</td><td>858</td><td>5384 (100%)</td><td>2643</td></tr><tr><td>50%</td><td>127</td><td></td><td>669</td></tr><tr><td>33%</td><td>62</td><td></td><td>370</td></tr><tr><td>Max. offsprings</td><td></td><td>54</td><td></td><td>11</td></tr><tr><td>Average</td><td></td><td>4.9</td><td></td><td></td></tr><tr><td>DNA verified</td><td>389</td><td>129</td><td>456</td><td>321</td></tr><tr><td rowspan="3">Llamas</td><td>2939 (100%)</td><td>474</td><td>3227 (100%)</td><td>1298.0</td></tr><tr><td>50%</td><td>54</td><td></td><td>318.0</td></tr><tr><td>33%</td><td>27</td><td></td><td>174.0</td></tr><tr><td>Max. offsprings</td><td></td><td>84</td><td></td><td>13.0</td></tr><tr><td>Average</td><td></td><td>6.2</td><td></td><td>2.5</td></tr><tr><td>DNA verified</td><td>138</td><td>52</td><td>147</td><td>101.0</td></tr></tbody></table>
<figure><img src="image_35.png" /><figcaption>**Figure 6:** Example of part of the ancestry tree. Three ancestry generations are shown on the web browser</figcaption></figure>

*Age Distribution*

The registry logs the time of registration and demands the date of birth. Furthermore, the owner can mark an animal as deceased. With these informations one can generate an age distribution of the animals. This is shown, separately for alpacas and llamas, in Fig. 7. One observes that the alpaca population is peaking at about 4 years, while the llama population shows a broader peak, with a maximum of about 5-6 years. One also can see that there are apparently llamas and alpacas at a very advanced age (beyond 25 years). These extremely long lifetimes are most likely not real, but are the result of forgetting to mark the animal as deceased.
a) Alpacas

<figure><img src="image_36.png" /></figure>

b) Llamas

<figure><img src="image_37.png" /><figcaption>**Figure 7:** Age distribution of the SACs in the LAREU database. Top: alpacas, bottom: llamas</figcaption></figure>

**Summary and Conclusion**

The European Camelid Registry LAREU is steadily growing with an increase of more than 2,000 animals per year. It is now the largest registry for camelids in Europe. With a total number of close to 20,000 animals (spring 2017), a number of significant statistics can be provided, giving some insight into the breeding program and the preferences for keeping these animals. In the recent years, alpacas
have clearly overtaken the llamas in numbers, which contribute now only one third of the total number of registered animals.

Using the large statistics of the LAREU registry it was shown that there are clear differences between the European countries concerning the distribution of colors in alpacas. Lacking further detailed information, no clear reasons for these differences can be given. In addition, one observes a significant trend in recent years towards “colorful” alpacas (beige, fawn, black, grey) which are becoming more popular at the cost of the white types.

The data also show that there is clear (and understandable) trend to use “prov-
en” sires and dams in the breeding program. However, the data relies dominantly
on the input of the animal owners, only a small fraction (about 10 % of the sires,
dams and offspring) are DNA tested with verified parentage.

In the pursuit of a meaningful breeding program, breeders should be encour-
aged to clarify the parentage by taking DNA typing of their animals more serious.

References

Alpaca Owners Association Inc. (previously Alpaca registry Inc. ARI), 2014, 8300 Cody
Dr, Ste A, Lincoln, NE 68512, www.alpacainfo.com

TASSO-Haustierzentralregister für die Bundesrepublik Deutschland e.V., 2017,
Otto-Volger-Str. 15, 65843 Sulzbach/Ts., Germany, www.tasso.net
<figure><img src="image_38.png" /></figure>
# Comparación de los Criterios de Selección de los Productores con el Reglamento Oficial para Llamas en el Perú

D.Y. Calderon<fnref n="1" />, M. Wurzinger<fnref n="2" />, J.G. Mendoza<fnref n="1" /> and G.A. Gutiérrez<fnref n="1" />

E-Mail: gustavogr@lamolina.edu.pe

E-Mail: maria.wurzinger@boku.ac.at

**Abstract.** The objectives of this study are: 1) To know the selection criteria for K'ara llama breeders 2) To identify the relationship between the selection criteria of the breeders with the descriptors of the K'ara racial standard existing in the Regulation of Genealogical Records of Alpacas and Llamas of Peru – RGALLP and 3) Associate the selection criteria of the breeders and the descriptors of the racial standard with the corporal measurements of K'ara llamas. The study was carried out in four zones of the Pasco Region, collecting information from 19 breeders with the evaluation of 21 llamas. A ranking of animals (from first to sixth place) was based on visual evaluation of breeders and technicians who used the guidelines of the official national registration system (RGALLP). Body weight and different body measurements were taken of each animal. The three main selection criteria for the breeders were: large cannon bone, chest width and height at the withers. The correlation between the ranking of the breeders versus the ranking of the technicians ranged from 0.76 to 0.99, being significant (p<0.05). However, this association does not mean that the selection criteria are the same. The average measurements of the circumference of cannon bone, chest width, height at the withers and live weight, are consistent with the ranking of the breeders, but this does not occur in the ranking of the technicians. The ranking of llamas by farmers and as well by technicians shows a negative relationship with body measurements. This means that llamas with larger corporal dimensions are usually better ranked. In general, the selection criteria of the breeders lead to larger animals in their body measurements and these in turn are oriented to a greater production of meat and its use as pack animals.

**Resumen.** Los objetivos del estudio fueron: 1) Conocer los criterios de selección de los criadores en llamas K'ara del sexo macho, 2) Identificar la relación entre los criterios de selección de los criadores con los descriptores del estándar racial de llamas K'ara existente en el reglamento de Registros Genealógicos de Alpacas y Llamas del Perú (RGALLP) y 3) Asociar los criterios de selección de los criadores y los descriptores del estándar racial con

<footnote n="1">¹Animal Breeding Program (PMA). Universidad Nacional Agraria La Molina – UNALM. Av. La Molina S/N. La Molina, Lima 12, Perú.</footnote>

<footnote n="2">²Division of Livestock Sciences. Department of Sustainable Agricultural Systems. University of Natural Resources and Life Sciences – BOKU. Gregor-Mendel-Str. 33, A-1180 Vienna, Austria.</footnote>
las medidas corporales de llamas K'ara. El estudio se realizó en cuatro zonas de la Región Pasco, recopilando información de 19 criadores con la evaluación de 21 llamas. La información de rankings (De primero a sexto lugar) de los animales fue por evaluación visual tanto de criadores como de técnicos (conocedores del RGALLP); Para determinar el peso corporal se utilizó una balanza tipo plataforma y para las medidas biométricas se utilizó un bastón zoométrico y una cinta inextensible graduada en centímetros. Los tres principales criterios de selección de los criadores fueron: cañas gruesas, ancho de pecho y altura a la cruz. La correlación entre el ranking de los criadores versus el ranking de los técnicos tuvo un rango de 0.76 a 0.99, siendo una asociación significativa (p<0.05), sin embargo, esta asociación no significa que los criterios de selección sean iguales. Las medidas promedio del perímetro de cañas, ancho de pecho, altura a la cruz y el peso vivo, son congruentes con el ranking de los criadores, más esto no ocurre en el ranking de los técnicos. La asociación entre el ranking de llamas, realizado por los criadores y los técnicos, con las medidas corporales son negativas, es decir las llamas con mayor dimensión corporal ocupan generalmente los primeros puestos en el ranking. En general, los criterios de selección de los criadores conducen a animales de mayores dimensiones en sus medidas corporales y estos a su vez están orientados a una mayor producción de carne y su adaptación a ser animales de carga.

**Keywords:** selection criteria, llamas, local knowledge

## Introducción

La mayor parte de la crianza de llamas a nivel alto andino en el Perú es realizada por pequeños productores bajo un sistema familiar. En general en el Perú hay escasa investigación realizada en llamas, ya que se da más importancia a la alpaca. La mayor parte de la investigación en llamas se ha realizado en la región sur del Perú. Las áreas de investigación son reproducción, salud animal, producción de fibra y carne, transformación y comercialización (Mendoza 2014). Lamentablemente se ha prestado poca atención al desarrollo de estrategias de selección, y al establecimiento de objetivos de mejoramiento genético en llamas. Por otro lado, la llama viene perdiendo importancia como animal de fibra y como animal de carga de bienes, producto del desarrollo de la alpaca para producción de fibra y el incremento de vías de comunicación en las zonas rurales. Como resultado de esta orientación la población de llamas está en peligro de extinción, a menos que se hagan esfuerzos concertados urgentes para seleccionar, caracterizar y manejar adecuadamente los rebaños (Maquera 1991).

La llama K'ara en gran medida está dirigida a la producción de carne por lo que presenta un potencial genético para este tipo de producción. Sin embargo, para poder seleccionar a los mejores animales se requiere definir los criterios de selección más adecuados. En la región de Pasco existen llamas fenotípicamente diferentes (mayor altura a la cruz y mayor longitud corporal) respecto a la región sur. Para el diseño de un programa de mejoramiento genético se requiere conocer los criterios de selección e indicadores fenotípicos corporales que utilizan para la selección de sus animales dirigidos a la producción de carne y/o carga.
Los objetivos del estudio fueron: 1) Conocer los criterios de selección de los criadores en llamas K'ara del sexo macho, 2) Identificar la relación entre los criterios de selección de los criadores con los descriptores del estándar racial de llamas K'ara existente en el reglamento de Registros Genealógicos de Alpacas y Llamas del Perú (RGALLP) y 3) Asociar los criterios de selección de los criadores y los descriptores del estándar racial con las medidas corporales de llamas K'ara.

# Materiales y Methodos

## Localización

El presente estudio se realizó en las zonas de Iscaycocha, Tunacancha, Los Andes y Pucunan, ubicadas en la Región Pasco, Perú.

## De los animales

De una tropa de llamas, se preseleccionaron animales machos en tres categorías (Categorías: A= bueno, B=regular y C=malo) de 1 a 2 años de edad, que fueron identificados con aretes de plástico codificados con numeración correlativa, y para la evaluación se les coloco bretes numerados. La preselección se realizó semanas antes de realizar el trabajo de investigación para cada zona (Cuadro 1).

<table><caption>Cuadro 1: Distribución de llamas K'ara según zona geográfica</caption><thead><tr><td>Zona</td><td>Sexo</td><td>Edad</td><td>n</td></tr></thead><tbody><tr><td>Iscaycocha</td><td>Macho</td><td>1 – 2 años</td><td>3</td></tr><tr><td>Tunacancha</td><td>Macho</td><td>1 – 2 años</td><td>6</td></tr><tr><td>Los Andes</td><td>Macho</td><td>1 – 2 años</td><td>6</td></tr><tr><td>Pucunan</td><td>Macho</td><td>1 – 2 años</td><td>6</td></tr><tr><td>Total</td><td></td><td></td><td>21</td></tr></tbody></table>

## Evaluación de los animales

La evaluación visual de las llamas por los criadores se realizó de una manera participativa. Un grupo de cuatro a cinco criadores por zona, realizaron individualmente un ranking (del mejor al peor animal) de los animales de acuerdo al fenotipo del animal y a sus propios criterios de selección. Además, informaron de las principales razones para la ubicación de las llamas en el ranking que cada uno elaboró independientemente, estas fueron registrados en fichas individuales. Luego los criadores discutieron sus resultados y llegaron a un consenso para establecer las características fenotípicas de mayor y menor importancia.

La evaluación visual de llamas por los técnicos, se realizó en cada zona, participando dos a tres técnicos. La evaluación se realizó utilizando las llamas que fueron
previamente evaluados por los criadores, pero de manera independiente. Los téc-
nicos realizaron la evaluación utilizando los descriptores del estándar racial de llam-
mas K'ara del Reglamento de los Registros Genealógicos de Alpacas y Llamas del
Perú (Minagri, 2011).

Mediadas corporals

Las medidas corporales ancho de pecho, altura a la cruz, y perímetro de cañas ante-
riores fueron medidas utilizando una cinta métrica de 1.5 metros y una regla de
madera de una longitud de 2 metros con una escuadra incorporada móvil. Para
medir el peso vivo de cada uno de los animales se utilizó una balanza ganadera
digital electrónica tipo plataforma.

Análisis de datos

Los criterios de selección de los criadores, se obtuvo del consenso y discusión
grupal que se realizó entre los criadores después de finalizada su evaluación,
además se usó la información de las fichas de calificación donde cada criador regis-
tró sus rankings y las razones de la evaluación visual por cada animal en cada una
de las zonas. Para realizar un ranking general de todos los criterios de selección de
los criadores se determinó mediante un índice. Los índices se calcularon como: la
suma de (8x para el ranking 1 + 7x para el ranking 2 + 6x para el ranking 3 + 5x
para el ranking 4 + 4x para el ranking 5 + 3x para el ranking 6 + 2x para el ranking
7 + 1x para el ranking 8) multiplicado por 0.01 (Abegaz 2014).

Para determinar el grado de asociación entre los rankings de los criadores y los rankings de los técnicos se utilizó la correlación de Spearman y para la asociación de los criterios de selección de los criadores y los descriptores del estándar racial con las medidas corporales se utilizó la correlación de Pearson.

Resultados y Discussion

Criterios de selección

Los criterios de selección utilizados por los criadores para la selección de llamas K'ara, se presentan en orden decreciente de acuerdo al índice obtenido (Cuadro 2). La característica “cañas gruesas” alcanza el índice más alto de 0.29 ocupando el primer lugar en orden de importancia; y la característica “presencia” alcanza un índice de 0.02 ocupando la última posición según el orden de importancia (Cuadro 2).
<table><caption>**Cuadro 2:** Ranking de criterios de selección de criadores según zonas</caption><thead><tr><th rowspan="2">Características</th><th colspan="4">Zona</th><th rowspan="2">Índice</th></tr><tr><th>Iscaycocha<br>(n = 5)</th><th>Tunacancha<br>(n = 5)</th><th>Los Andes<br>(n = 5)</th><th>Pucunan<br>(n = 4)</th></tr></thead><tbody><tr><td>Cañas gruesas</td><td>1</td><td>1</td><td>1</td><td>1</td><td>0.29</td></tr><tr><td>Ancho de pecho</td><td>3</td><td>2</td><td>2</td><td>2</td><td>0.24</td></tr><tr><td>Alzada</td><td>2</td><td>3</td><td>3</td><td>3</td><td>0.22</td></tr><tr><td>Cuello largo</td><td>5</td><td>5</td><td>4</td><td>5</td><td>0.14</td></tr><tr><td>Longitud corporal</td><td>-</td><td>4</td><td>-</td><td>4</td><td>0.10</td></tr><tr><td>Conformación testicular</td><td>-</td><td>6</td><td>-</td><td>6</td><td>0.06</td></tr><tr><td>Tipo</td><td>6</td><td>8</td><td>7</td><td>8</td><td>0.04</td></tr><tr><td>Conformación de cabeza</td><td>-</td><td>7</td><td>-</td><td>7</td><td>0.04</td></tr><tr><td>Ancho de anca</td><td>4</td><td>-</td><td>-</td><td>-</td><td>0.03</td></tr><tr><td>Fortaleza</td><td>-</td><td>-</td><td>5</td><td>-</td><td>0.03</td></tr><tr><td>Presencia y otros</td><td>-</td><td>-</td><td>6</td><td>-</td><td>0.02</td></tr></tbody></table>

1 = Primer lugar, 2 = Segundo lugar, 3 = Tercer lugar, etc. (Ranking de las características); n = Numero de criadores.

Las características cañas gruesas, ancho de pecho y altura a la cruz, ocupan el 1er, 2do y 3er lugar, respectivamente; siendo este orden consistente en todas las zonas de estudio. Estas características fueron determinadas por los criadores de llamas K'ara como criterios de selección para la evaluación visual de reproductores orientados mayormente a la producción de carne. En este contexto, los criadores consideran que animales más gruesos (con pecho amplio, cañas gruesas, cuello ancho) y altos tendrán mayor producción cárnica, no tomando en cuenta ningún indicador zoométrico que tenga relación con una mayor masa muscular; ni la orientación de la selección en función de características de importancia económica como es el peso vivo para las llamas K'ara (Mamani et al., 2011).

Estos resultados concuerdan con los criterios de selección empleados por criadores de las zonas alto andinas de Arequipa (Torres, 2014), quienes mencionan criterios como llamas gruesas y altas, al igual que lo reportado para Pasco (Mendoza, 2014) como la alzada, conformación y tipo. En Bolivia además de la alzada se consideran otras características como largo de cuerpo, largo de cuello y ausencia de defectos congénitos (Rodríguez y Quispe, 2007, Saavedra et al., 2012, Sepúlveda, 2011), y también la conformación corporal, conformación testicular y salud (Markemann y Valle Zárate, 2009), además, Wurzinger et al. (2007), también reporto como único criterio el tamaño corporal.
## Correlación del ranking de los criadores vs el ranking de los técnicos

Los valores de los coeficientes de correlación de Spearman entre los rankings de criadores versus el ranking de los técnicos vario de 0.76 a 0.99 (Cuadro 3). Estos valores relativamente altos, nos indicarían la existencia de una significativa asociación entre los rankings elaborados a partir de información de los criadores y de los técnicos. Si bien el ranking de criadores y técnicos tienen similar orden de los animales con fines de selección, ello no significa que los criterios de ambos sean similares. Los criadores utilizan criterios de selección diferentes a los que se describen en el estándar racial del RGALLP.

<table><caption>**Cuadro 3:** Coeficientes de correlación de Spearman entre los rankings de criadores con el de los técnicos, según zonas</caption><thead><tr><th></th><th colspan="4">Zonas</th></tr><tr><th></th><th>Iscaycocha</th><th>Tunacancha</th><th>Los Andes</th><th>Pucunan</th></tr></thead><tbody><tr><th>Número de criadores</th><td>5</td><td>5</td><td>5</td><td>4</td></tr><tr><th>Número de técnicos</th><td>3</td><td>3</td><td>3</td><td>2</td></tr><tr><th>Número de animales</th><td>3</td><td>6</td><td>6</td><td>6</td></tr><tr><th>Coef. de Spearman</th><td>0.99<sup>**</sup></td><td>0.88<sup>*</sup></td><td>0.89<sup>*</sup></td><td>0.76</td></tr></tbody></table>

## Medidas corporales y el peso vivo

### Según ranking de criadores

Las llamas que obtuvieron el primer lugar en el ranking de los criadores, son aquellas que registraron los mayores valores promedio en las tres principales características y en el peso vivo (Cuadro 4). Estos resultados nos indicarían que las evaluaciones visuales realizadas por los criadores son congruentes con mayores valores de las medidas corporales y peso vivo.

Varios autores reportan medidas corporales para llamas K'ara de uno y dos años, así en la región de Puno: Maquera (1991) reporta para altura a la cruz 91.65 y 98.60 cm y pesos vivos de 60.05 y 87.35 kg.; García y Franco (2006) también mencionan medidas en altura a la cruz 94.60 y 102.30 cm y en ancho de pecho 23.10 y 25.70 cm.; y en Oruro – Bolivia Cortez et al. (2006) para altura a la cruz 98.3 y 101.1 cm y peso vivo para llamas de 2 años de edad de 96.70 kg. Todos estos valores son inferiores a los reportados en el presente estudio, confirmando que las llamas de la región Pasco presentarían mayor altura, conformación y peso corporal, en comparación con llamas de otras regiones del sur del Perú y Bolivia

<footnote>* = Significativo (p<0.05), ** = Significativo (p<0.01).</footnote>
(Gutiérrez et al. 2012). Sin embargo, Cano et al. (2012) en Marcapomacocha – Junín, reporta valores para altura a la cruz de 110.70 y 115.10 cm., para ancho de pecho 33.80 y 36.10 cm y pesos vivos de 122.60 y 137.70 kg, en llamas K'ara, de uno y dos años de edad, respectivamente, los cuales son superiores a los valores hallados. Tales medidas, reportadas por Gutiérrez et al. (2012) y por Cano et al. (2012), nos indicarían que las llamas del Centro del País (Pasco y Junin) son mucho más desarrolladas, mostrándonos una mayor altura y un mayor peso vivo en relación a las llamas del sur, indicándonos que el estándar racial del RGALLP no concuerda con las características de las llamas K'ara de la Región Pasco. Nuestros resultados, y los reportes mencionados, podrían explicar la causa del porque los criadores eligen estas características como los principales criterios de selección los cuales están asociados directamente a una mayor producción de carne.

<table><caption>**Cuadro 4:** Medidas promedio de perímetro de cañas anteriores, ancho de pecho, altura a la cruz y peso vivo según el ranking de los criadores</caption><thead><tr><th>Ranking general</th><th>n</th><th>Perímetro de cañas anteriores (cm)</th><th>Ancho de pecho (cm)</th><th>Altura de la cruz (cm)</th><th>Peso vivo (kg)</th></tr></thead><tbody><tr><td>Primero</td><td>4</td><td>14.89</td><td>28.42</td><td>109.42</td><td>122.53</td></tr><tr><td>Segundo</td><td>4</td><td>14.05</td><td>26.16</td><td>107.68</td><td>112.56</td></tr><tr><td>Tercero</td><td>4</td><td>13.26</td><td>25.84</td><td>106.63</td><td>101.26</td></tr><tr><td>Cuarto</td><td>3</td><td>13.14</td><td>25.00</td><td>103.43</td><td>99.04</td></tr><tr><td>Quinto</td><td>3</td><td>13.57</td><td>24.79</td><td>101.64</td><td>94.09</td></tr><tr><td>Sexto</td><td>3</td><td>12.79</td><td>23.29</td><td>100.21</td><td>89.30</td></tr></tbody></table>

### Según ranking de técnicos

Las llamas que obtuvieron el primer lugar en el ranking de los técnicos (Cuadro 5), no siempre son aquellas que registraron los mayores valores promedio, a excepción de la característica “ancho de pecho”. Consecuentemente, se aprecia valores desordenados que no están de acuerdo al ranking general, claramente se aprecia que no hay un patrón en forma descendente, diferente a los observado para el caso de los criadores. Probablemente estos resultados se deban a que los descriptores del estándar racial del RGALLP se basa en criterios de selección distintos y abstractos, cuyos puntajes no reflejan las características corporales y peso vivo de los animales evaluados (Cuadro 5).
<table><caption>**Cuadro 5:** Medidas promedio de perímetro de cañas anteriores, ancho de pecho, altura a la cruz y peso vivo según el ranking de los técnicos</caption><thead><tr><th>Ranking general</th><th>n</th><th>Perímetro de cañas anteriores (cm)</th><th>Ancho de pecho (cm)</th><th>Altura de la cruz (cm)</th><th>Peso vivo (kg)</th></tr></thead><tbody><tr><td>Primero</td><td>4</td><td>14.25</td><td>27.50</td><td>108.00</td><td>112.15</td></tr><tr><td>Segundo</td><td>4</td><td>14.75</td><td>26.50</td><td>110.25</td><td>121.20</td></tr><tr><td>Tercero</td><td>4</td><td>13.00</td><td>26.50</td><td>106.50</td><td>100.60</td></tr><tr><td>Cuarto</td><td>3</td><td>13.33</td><td>25.67</td><td>106.33</td><td>108.17</td></tr><tr><td>Quinto</td><td>3</td><td>12.33</td><td>21.67</td><td>101.33</td><td>87.63</td></tr><tr><td>Sexto</td><td>3</td><td>13.00</td><td>24.00</td><td>98.33</td><td>83.93</td></tr></tbody></table>

### Asociación de los criterios de selección de los criadores con las medidas corporales

Los coeficientes de correlación entre el ranking general de los criadores con las medidas corporales (perímetro de caña anterior, ancho de pecho y altura a la cruz) y el peso vivo, resultaron ser negativos (Cuadro 6). Los coeficientes de correlación indican que si el animal esta mejor posicionado en el ranking (es decir más próximos al primer lugar) las medidas corporales y el peso vivo serán de mayor magnitud.

<table><caption>**Cuadro 6:** Coeficientes de correlación de Pearson del ranking de los criadores versus medidas corporales según zona</caption><thead><tr><th>Ranking de la zona</th><th>n</th><th>Perímetro de caña anterior</th><th>Ancho de pecho</th><th>Altura a la cruz</th><th>Peso vivo</th></tr></thead><tbody><tr><td>Iscaycocha</td><td>3</td><td>-0.87</td><td>-0.87</td><td>-0.72</td><td>-0.87</td></tr><tr><td>Tunacancha</td><td>6</td><td>-0.53</td><td>-0.63</td><td>-0.71</td><td>-0.23</td></tr><tr><td>Los Andes</td><td>6</td><td>-0.47</td><td>-0.40</td><td>-0.66</td><td>-0.60</td></tr><tr><td>Pucunan</td><td>6</td><td>-0.56</td><td>-0.54</td><td>-0.76</td><td>-0.75</td></tr><tr><td>Promedio</td><td></td><td>-0.61</td><td>-0.61</td><td>-0.71</td><td>-0.61</td></tr></tbody></table>

### Asociación de los descriptores del estándar racial con las medidas corporales

Los promedios de coeficientes de correlación entre el ranking general de los técnicos basados en los descriptores del estándar racial de llamas K'ara con las mé-
didas corporales (perímetro de caña anterior, ancho de pecho y altura a la cruz) y el peso vivo, también son negativas (Cuadro 7).

<table><caption>**Cuadro 7:** Coeficientes de correlación de Pearson del ranking de los técnicos versus medidas corporales según zona</caption><thead><tr><th>Ranking de la zona</th><th>n</th><th>Perímetro de caña anterior</th><th>Ancho de pecho</th><th>Altura a la cruz</th><th>Peso vivo</th></tr></thead><tbody><tr><td>Iscaycocha</td><td>3</td><td>- 0.87</td><td>- 0.87</td><td>- 0.76</td><td>- 0.87</td></tr><tr><td>Tunacancha</td><td>6</td><td>- 0.68</td><td>- 0.62</td><td>- 0.60</td><td>- 0.49</td></tr><tr><td>Los Andes</td><td>6</td><td>- 0.82</td><td>-0.76</td><td>- 0.84</td><td>- 0.85</td></tr><tr><td>Pucunan</td><td>6</td><td>- 0.75</td><td>- 0.45</td><td>- 0.46</td><td>- 0.48</td></tr><tr><td>Promedio</td><td></td><td>- 0.78</td><td>- 0.68</td><td>- 0.67</td><td>- 0.67</td></tr></tbody></table>

Al igual que en el caso de los criadores estos coeficientes nos indican que si el animal esta mejor posicionado en el ranking (es decir más próximos al primer lugar) las medidas corporales tanto perímetro de caña anterior, ancho de pecho, altura a la cruz y peso vivo serán de mayor magnitud (es decir mayor perímetro de caña anterior, ancho de pecho y altura a la cruz en centímetros y mayor peso vivo en kg).

## Conclusiones

En la región Pasco, los criadores que participaron en el estudio identificaron los siguientes criterios de selección según su importancia: 1º) perímetro de cañas, 2º) amplitud de pecho y 3º) la altura a la cruz (talla). La correlación entre el ranking de los criadores versus el ranking de los técnicos varió de 0.76 a 0.99, indicando diferencias menores entre ellos. El valor promedio del perímetro de cañas, ancho de pecho, altura a la cruz y el peso vivo, son congruentes con el ranking de los criadores, pero no con el ranking de los técnicos. En general, los criterios de selección de los criadores conducen a animales de mayores dimensiones en sus medidas corporales y estos a su vez están orientados a una mayor producción de carne y su adaptación a ser animales de carga.

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# Selection and Evaluation of Fiber Characteristics of an Extreme Fine Alpaca Strain at Victory Farm in Missouri

T. Wuliji

Lincoln University of Missouri, Jefferson City, Missouri, 65102, USA

E-Mail: wuliji@lincolnU.edu

**Abstract.** Data recordings of animal breeding, fleece production, and fiber characteristics in an alpaca herd were analyzed and presented. A selective mating strategy was applied while establishing an extremely fine fiber alpaca strain at Victory Farm over 10 years. The herd size was expanded from one dozen breeding females and five males to 200 head in 2016. All animals were recorded for sires, dams, and registered pedigree, birth weight, weaning weight, shearing weight, and fleece weight. Mid-side flank fleece samples were measured for Optical-based Fibre Diameter Analyser (OFDA) fiber characteristics, including mean fiber diameter, length, and curvature. Live weight, fleece weights, and fiber characteristics were analyzed using SAS® GLM procedures. Mean birth weight, weaning weight, mixed age shearing weight, fleece weight, fiber diameter, coefficient of fiber diameter variation, fiber length, and fiber curvature were 6.82 kg, 25.27 kg, 48.84 kg, 0.85 kg, 16.67 µm, 25.2 %, 64.6 mm, and 56.9 deg/mm, respectively. In addition, the finest 25 percent of the herd's fleeces were tested and found to have an average fiber diameter of 14.17 µm, which was within vicuna fiber diameter ranges. These characteristics did not significantly increase with age. Heritability estimates for the fleece weight and fiber diameter, length, and curvature were 0.40, 0.65, 0.29, and 0.50, respectively. The average fiber diameter measurements of the herd at Victory Farm were significantly (p<0.001) finer than the comparison herds. This study found that strict selective breeding is effective for genetic gains in ultralow fiber diameter and fleece quality in an extremely fine fiber alpaca strain.

**Resumen.** Se analizó y se presentó información sobre cía de animales, producción de vellones y características de fibra en un rebaño de alpacas. Se ha creado una variante de alpacas extremadamente finas mediante estrategias de apareamiento selectivo durante 10 años en la Granja Victory. El rebaño partió de una docena de hembras y cinco machos reproductores, llegando en 2016 a 200 cabezas. Se registró el padre y la madre y el pedigrí completo de todos los animals, así como el peso al nacimiento, al destete y los pesos de esquila y del vellón. Se analizaron muestras de vellón del costillar medio para obtener el diámetro, la longitud y la curvatura medios de la fibra mediante un Analizador de Diámetro de Fibra Óptico (OFDA). El peso vivo, los pesos del vellón y las características de la fibra se analizaron con el procedimiento GLM del software SAS®. Las medias del peso al nacimiento, al destete, peso a edad mixta de esquila, peso del vellón, diámetro de fibra, coeficiente de variación, longitud de fibra y curvatura de la fibra, fueron 6,82 kg, 25,27 kg,
48,84 kg, 0,85 kg, 16,67 µm, 25,2 %, 64,6 mm y 56,9 °/mm respectivamente. Además, se analizó únicamente el 25 % mejor de los vellones del rebaño y se obtuvo un diámetro de fibra medio de 14.17 µm, valor dentro del rango de la fibra de la vicuña. Estas características no aumentaron significativamente con la edad. Las heredabilidades estimadas para peso del vellón, diámetro, longitud y curvatura de la fibra fueron 0,40, 0,65, 0,29 y 0,50 respectivamente. El diámetro medio de fibra de la Granja Victory fueron significativamente (p<0,001) más finas que los rebaños de comparación. Este estudio demostró que la mejora selectiva estricta es efectiva para obtener ganancias genéticas en ultrabajo diámetro de y de calidad del vellón en una variante de alpaca de fibra extremadamente fina.

**Keywords:** Vicugna pacos, alpaca strain, extremely fine, selective breeding, heritability

**Introduction**

Although camelids are indigenous to the Andean highlands of South America, now they are distributed in many parts of the world. The alpaca (Vicugna pacos) is the most numerous fiber-producing member of the four South American camelid species: llama, guanaco, alpaca, and vicuna (Kadwell et al., 2001). Two types of alpacas have been introduced into the United States, namely, huacaya and suri; however, the majority of alpacas (80 %) are huacaya. Whereas Peru owns 75 percent of the alpaca population and produced about 90 percent of the world's camelid fiber in the past (Morante et al., 2009), the alpaca population introduced into nonnative regions has increased steadily in the last 30 years, especially in Australia, Canada, the United Kingdom, New Zealand, and the US. Alpacas are becoming an important specialty fiber production livestock species in these regions beyond the South American Andes. The alpacas registered to the Alpaca Registry, Inc. (ARI) in the US from 1986 to 2017 have increased to 255,128 head and the total number of fiber-growing camelids has been estimated at passing the 300,000 population mark in the US (Alpaca Consulting Services USA, 2017). Alpacas can be found in every state of the US and are farmed in various geographical environments, ranging from the desert to high mountain ranges (Wuliji, 2012). Coat color varies widely in camelids, ranging from white to black and various shades, including 22 natural color categories (Hoffman, 2003). Fleece production and fiber traits have been reported in some studies in the alpacas' native land (Pumayalla & Leyva, 1988; Frank et al., 2006; Montes et al., 2008; Morante et al., 2009; Cervantes et al., 2010). Animal husbandry and reproduction, especially fleece and fiber characteristics, were studied in these nonnative environments (Wuliji et al., 2000; McGregor, 2004; McColl et al., 2004). Profiles of the fiber characteristics of both huacaya and suri breeds farmed in the US have been reported (Lupton et al., 2006; Wuliji, 2012). Some of the literature and production records indicate that the introduced camelids had significantly improved body weight and fleece weight performance, whereas the fiber diameter had become coarser when grown on more nutritive grazing lands than the high Andes. The owners of Victory Farm
initiated an alpaca breeding venture in Braymer, Missouri, with just one dozen females and five males 10 years ago. By persistently acquiring breeding stock from fellow breeders and through restrictive selection within their animals, their herd size increased to 200 head in 2016. The goal was to establish an extremely fine alpaca herd by corrective breeding and intensive selection within the US camelid population. Therefore, this study was conducted to record, evaluate, and summarize the progress of breeding, production, and management practices established at Victory Farm. Animal production performance was presented for birth weight (BW); weaning weight (WW); shearing live weight (SW); fleece weight (FW); staple length (SL); and fiber characteristics, including fiber diameter (FD), fiber diameter variation (FDsd and FDcv), curvature (CUR), and medullation rate (MD).

## Materials and Methods

### Animal husbandry, welfare, and health management

Victory Farm, owned by the Smith family, is located in Missouri at 39.5870° N, 93.7960° W, at an elevation of 233 m above sea level, with a landscape of rolling hills, with an annual average precipitation of 914 mm as rain and 406 mm as snowfall. Monthly average temperature ranges were recorded at -8 °C to 3 °C (January), 6 °C to 18 °C (April), 20 °C to 32 °C (July), and 7 °C to 20 °C (October). As of April 2016, the herd size had increased to 200 animals, including males, females, and crias. All animals were identified by implanted microchips for pedigrees, parentage, and breeding society registrations. Animals were grazed as a mixed-age herd on pastures during warm seasons and fed on dry feeds under shelter during the winter months. Pasture plant species on the farm are a mixture of grasses and legumes, including fescue, orchard, brome, bluegrass, and white clover. Animals were grazed on pastures and rotated more frequently and efficiently at a stocking rate of 10 head per acre. Adult males and females were penned and fed separately in a drylot during winter. High-quality forages, clean water, and mineral supplements were provided ad lib. Alfalfa hay containing 12 to 16 % crude protein was fed to females, yearlings, and weanlings, whereas brome or orchard grass hay with 10 % crude protein was fed to males during drylot feeding. Animals were regularly graded for body condition scores as nutrition and health status indicators. Animal welfare and health care were contracted with the local veterinary practitioner. A routine vaccination for recommended preventable illnesses was practiced, such as an injection of Clostridium perfringens type C + D and tetanus (CD&T) at annual intervals and an injectable dewormer (Noromectin®) at six-week intervals during seasons prevalent with internal parasites. Over 10 years of operation, the animal mortality rate was kept below 2 %. Other animal welfare and health care was performed, such as timely
inspection for eye infections, scours, and bloating as well as toenail trimming and tooth grinding at shearing operations.

## Production, selection, and breeding

Animal selection criteria were set for three objective measurements and two subjective traits, namely, fiber diameter (FD, µm), staple length (SL, mm), curvature in degree (CUR, deg/mm), coat color, and fleece density. The first breeding age was normalized at three years or older for males and two years for females; however, a female can be bred at about 18 months old if her live weight reaches 45 kg. Most animals were fall bred and fall born; however, a small number of animals could be spring bred if they were a noncarrier or not a bearer from fall breeding. Breeding was practiced as a single-pair mating in pens. Studs were fed and managed in separate barns and halter-trained prior to breeding. The breeding pair match-up was decided with reference to the offspring from previous mating allocations. Regular animal performance records were made on-farm, including birth date, sex, sire and dam, pedigree, birth weight (BW), weaning weight (WW), shearing body weight (SW), and fleece weight (FW). Animal breeding, pedigree, and production performance data were submitted to Colorado State University (Fort Collins, CO) to generate breeding value estimates. Data was recorded in an Excel spreadsheet for retrieval, communication, storage, and analysis. Animal registry information included animal ID (microchip), pedigrees, health records, and fiber test records.

Animals were estimated for breeding values using the expected progeny difference (EPD), which was updated annually to serve as a reference for individual animal sire or dam selection. As alpacas are self-induced ovulators, the estrus cycle occurs after breeding. Therefore, the female was reintroduced to a male seven days later but would reject mating if she was pregnant; otherwise, she would submit to breeding. The pregnancy was confirmed by observation of her “spitting off” for three consecutive weeks, and predicted cria due dates could then be placed on the birthing calendar. The gestation length averaged 345 d on-farm but was observed as early as 317 d to as late as 388 d for normal birthing. From weaning age, animals were trained for weighing, catching, and releasing as well as halter-training so they could develop an early interaction with humans and become less stressed by handling.

## Blood pack cell volume and cell type measurements

In 2017, blood samples were taken from 24 animals (12 males and 12 females, with each sex including four crias) representing the herd to determine blood pack cell volumes and blood cell profile survey measurements. Blood samples were drawn from the jugular vein from the right side of the neck (the C6 vertebra) using venipuncture fitted with a needle and collected into heparinized tubes (purple top).
A number of blood volume and cell parameters, including white blood cell (WBC) count and types (i.e., neutrophil, lymphocyte, eosinophil, and monocyte), red blood cell (RBC) count, hematocrit (HCT%), and platelet concentration were examined using the HEMAVET® HV950 hematology instrument (Drew Scientific, Inc.).

*Shearing, fleece classing, and fiber trait measurements*

Animals were scheduled to shear annually in April. Animals were sorted into coat color groups and penned under shelter overnight to prevent overfeeding or wetting of the fleece. The shearing floor space was cleaned and covered with a tarp/canvas sheet that was used for animal restraining and for fleece collections. Shearing started with the lighter shade fleeces, such as white, cream, tan, and light brown to dark brown. The shearing procedure followed an established alpaca shearing technique, which is similar to the procedure described by Wuliji et al. (1992), namely, the animals were shorn lying on their side on a tarp/canvas sheet-covered floor, with their two hind legs tied to a wall. Electric clippers with a sheep shearing comb and cutters were used to conduct shearing, while an assistant held the alpaca by both front legs, stretching and rolling the alpaca to expose the unshorn side fleece for the shearer. The saddle (blanket) fleece part was shorn first, followed by the rest of the fleece coat. The blanket and saddle fleece parts were weighed separately, recorded, bagged individually, labeled, and stored for further fiber classing, processing, and/or marketing. A small batch of mid-side fiber sample was collected from each animal for fiber trait measurements at each shearing and tested by the Yocom-McColl Testing Laboratories Inc. (Denver, CO). Yocom-McColl uses the Optical-based Fibre Diameter Analyser (OFDA 100) to measure a set of traits, such as the fiber diameter in microns, which is the ASTM standard (D2252) for alpaca fiber. Fiber measurements included FD, FD standard deviation (FDsd), and FD coefficient of variation (FDcv), SL, F30 (percentage of fiber diameter greater than 30 micron), CF (comfort factor, calculated as 100 – F30), curvature (CUR), and percentage of medullation rate (MD).

*Specialty camelid fiber collection, measurement, and comparison*

As a reference analysis, 15 fiber samples (dehaired) of Bactrian camel from Inner Mongolia and 17 fiber specimens of vicunas from private collectors and zoos were donated for this in-house study. These samples were measured and compared for FD and staple length with the fiber from the extremely fine fiber animals (25 %) from the Victory Farm herd in 2016 and 2017. Staple length was measured in mm, with five replicates. Fiber diameter was measured according to a standard microscopic method that measures 250 individual fibers randomly per specimen prep slide, following the standard ISO 137:2015 Wool – Determination of Fibre Diameter – Projection Microscope Method (2015). Eight of those vicuna fiber specimens were also tested at Yocom-McColl and validated for the accuracy of the
test by microscope procedures, which showed a 99.9 % agreement between the two methods. The FD, FDsd, FDcv, and SL of the extremely fine groups (2016 and 2017) were selected from the Yocom-McColl lab test results. The fiber test results of the Victory Farm herd (shearing 2016) are included in a multi-ranch reference comparison (Table 6).

*Expected progeny difference and heritability estimate*

The herd-expected progeny difference (EPD) and heritability ($h^2$) values were estimated using a data pool of the breeding group's herds, which Victory Farm had organized for the collaborative breeding program. The pedigree and performance data of 760 animals with 2,915 performance records were included in this analysis. The expression $h^2$ was estimated as the proportion of the difference in performance that is explained by a transmittable genetic difference. Multiple trait evaluation procedures were applied to estimate the correlation coefficient ($r$) among these traits, which may have a positive or negative effect on each other.

*Data analysis and presentation*

Victory Farm alpaca herd data files for birth weight, weaning weight, and shearing weight were retrieved and pooled for data analysis and presentation. Data were analyzed and compared among production year, sex, and age groups for body weight as well as fleece and fiber characteristics. Fleece weight and fiber characteristics were pooled and analyzed on 1,230 records of 11 annual shearings (2007 to 2017). However, 1,021 observations were used for most traits, as some fleece weight or fiber characteristic measurements were either incomplete or missing for the production years 2007 and 2008. This resulted from the small herd size, so data from those years were excluded from the comparisons. There were 173 birth weight (2010 to 2016) and 166 weaning weight records available for this analysis. Shearing live weight (SW) records of 478 animals were available from 2014 to 2016, which were compared by age groups with their repeated measurements. The FD, FDcv, SL, F30, CUR, (deg/mm), and MD rates were provided as part of the systematic standard measurements by the Yocom-McColl test lab, except for the farm reference study specimens. Production performance data by year, age, and sex were extracted from the Victory Farm database. SAS® statistical procedures (i.e., GLM, ANOVA, CORR, and t-test) were used to process those data sets. The main effects were primarily compared for production year, sex, and age; however, the EPD calculation and $h^2$ estimates were adopted from the Victory Farm group breeding annual reports (Enns & Speidel, 2017).
## Results and Discussion

Fleece weight and fiber characteristics are summarized by production years in Table 1. There were several significant differences (p<0.05) in SL, FD, FDcv, and CUR. The FW and SL decreased gradually in the past few years, whereas the FDcv slightly increased over time in the herd. There was no significant difference in the FW, except for the 2009 shearing, which was highly skewed due to the shearing regimen and the small number of animals. The measure for the CUR and F30 remained steady, with only a small fluctuation. Except in 2010, when there was a significant surge, the F30 regularly tested low (i.e., from 0.96 to 1.62 %), which was interpreted as the average CF of the fleeces for the herd over all the years of the study, which ranged from 98.4 to 99.9 %. The FD of the herd gradually decreased, while the number of animals in the herd increased, and the FD decreased significantly (p<0.05) in production years 2016 and 2017.

<table><caption>**Table 1:** Fleece weight (FW), staple length (SL), mean fiber diameter (FD), coefficient of fiber diameter variation (FDcv), percentage of fiber coarser than 30 micron (F30), and fiber curvature (CUR) measurements by production years</caption><thead><tr><th>Year</th><th>Numbers</th><th>FW (kg)</th><th>SL (mm)</th><th>FD (µm)</th><th>FDcv (%)</th><th>F30 (%)</th><th>CUR (deg/mm)</th></tr></thead><tbody><tr><td>2009</td><td>35</td><td>1.43<sup>n*</sup></td><td>87.4<sup>c</sup></td><td>18.02<sup>bc</sup></td><td>25.2<sup>abc</sup></td><td>1.43</td><td>57.61<sup>ab</sup></td></tr><tr><td>2010</td><td>59</td><td>1.11</td><td>75.2<sup>bc</sup></td><td>18.37<sup>cd</sup></td><td>25.5<sup>ab</sup></td><td>2.16<sup>n*</sup></td><td>53.17<sup>a</sup></td></tr><tr><td>2011</td><td>67</td><td>1.08</td><td>64.2<sup>ab</sup></td><td>18.19<sup>bc</sup></td><td>23.0<sup>a</sup></td><td>1.49</td><td>53.81<sup>a</sup></td></tr><tr><td>2012</td><td>68</td><td>1.04</td><td>65.5<sup>ab</sup></td><td>16.95<sup>a</sup></td><td>24.0<sup>ab</sup></td><td>1.13</td><td>57.81<sup>ab</sup></td></tr><tr><td>2013</td><td>77</td><td>0.85</td><td>57.2<sup>a</sup></td><td>17.03<sup>ab</sup></td><td>24.1<sup>ab</sup></td><td>1.03</td><td>61.49<sup>bc</sup></td></tr><tr><td>2014</td><td>168</td><td>0.83</td><td>56.7<sup>a</sup></td><td>17.13<sup>ab</sup></td><td>24.8<sup>abc</sup></td><td>1.05</td><td>58.15<sup>bc</sup></td></tr><tr><td>2015</td><td>143</td><td>0.78</td><td>57.5<sup>a</sup></td><td>17.36<sup>bc</sup></td><td>26.6<sup>bc</sup></td><td>1.32</td><td>53.87<sup>a</sup></td></tr><tr><td>2016</td><td>164</td><td>0.76</td><td>57.8<sup>a</sup></td><td>16.58<sup>a</sup></td><td>26.4<sup>bc</sup></td><td>0.96</td><td>57.66<sup>ab</sup></td></tr><tr><td>2017</td><td>171</td><td>0.73</td><td>59.9<sup>a</sup></td><td>16.67<sup>a</sup></td><td>27.2<sup>bc</sup></td><td>1.05</td><td>58.30<sup>bc</sup></td></tr></tbody></table>

The effect of age was more pronounced on the SL, and the CUR decreased with increasing age. There was no difference in the SL by sex; however, for crias, the SL was reported to grow longer in the warm seasons than in the cold seasons in Peru (Quispe-Peña et al., 2014). Both the FD and FDcv in the herd showed minor fluctuations with increasing age, which was significantly less varied compared to other huacaya or suri alpacas (Wuliji, 2012). The Victory Farm alpaca herd's cumulative FD measurements were plotted by production year (Figure 1). The mean FD of the herd decreased gradually and consistently from 2007 to 2017, while the herd size increased. The apparent annual average reduction in the FD was 0.21 microns per year in the last eight fleece measurements, which reduced the herd average FD

<footnote>abcd: means bearing a different superscript differ within the column at the p < 0.05 level</footnote>

<footnote>n*: significantly differed (p < 0.05) from the rest of the column</footnote>
to 16.58 and 16.68 in 2016 and 2017, respectively. The FW and F30 fluctuated and slightly decreased with age, whereas the MD rose with age. There were almost no changes in the FW and F30 measurements based on age but several significant (p<0.05) surges in the MD corresponded only to the eight-year-old age group (data were not shown). The herd average fiber diameter, staple length, fiber curvature, fiber diameter, and variation analysis by age group is shown in Figure 2.

<figure><img src="image_39.png" /><figcaption>**Figure 1:** Scatter plot of individual animal FD measurements (cumulative percentage of the herd) in the Victory Farm alpaca herd by production year (2007–2017).</figcaption></figure>
<figure><img src="image_40.png" /><figcaption>**Figure 2:** Staple length, fiber diameter and diameter variation, and fiber curvature measurements in alpaca age groups.</figcaption></figure>

<table><caption>**Table 2:** Fleece weight and fiber characteristic measurements compared by sex (pooled for years and ages)</caption><thead><tr><th>Sex</th><th>Freq. No</th><th>FW (kg)</th><th>SL (mm)</th><th>FD (µm)</th><th>FDcv (%)</th><th>F30 (%)</th><th>FC (%)</th><th>CUR (deg/mm)</th><th>MD (%)</th></tr></thead><tbody><tr><td>Female</td><td>576</td><td>0.84</td><td>60.0</td><td>17.59</td><td>24.7</td><td>1.39</td><td>98.61</td><td>56.3</td><td>2.1</td></tr><tr><td>Male</td><td>443</td><td>0.85</td><td>61.3</td><td>16.69</td><td>26.2</td><td>1.03</td><td>98.97</td><td>57.8</td><td>1.3</td></tr><tr><td>SD</td><td></td><td>0.29</td><td>18.8</td><td>2.05</td><td>4.6</td><td>1.16</td><td>1.16</td><td>10.8</td><td>5.9</td></tr></tbody></table>

Fleece weight and fiber characteristics are summarized by sex in Table 2. There was no difference for the FW, SL, and CUR, but the FD, F30, and MD were higher for females than males, and the FDcv was coarser for males than females. BW and WW were presented for birth years by sex, and the means were pooled (Table 3). There was no difference based on sex, however, there was a small but significant difference in the mean BW and WW by birth years, and both the BW and WW were higher for females than males. There were a number of significant positive or negative correlations among fleece weight and fiber characteristics (Table 4). There was a significant and positive or negative correlation (p<0.01) among all traits except MD with SL; there were no significant corrections with the FDcv. There were positive and significant correlations found among FW, SL, FD, F30, and MD, whereas there were some negative and significant associations between the FD and each of the following: FDcv, F30, FC, CUR, and CUR standard deviation (CURsd).
<table><caption>**Table 3:** Birth weight (kg), weaning weight, and the mean weights of alpacas recorded for six consecutive years after birth</caption><thead><tr><th>Birth weight (BW)</th><th>2010</th><th>2011</th><th>2012</th><th>2013</th><th>2014</th><th>2015</th><th>2016</th></tr></thead><tbody><tr><th scope="row">Female BW</th><td>6.8<sup>ab</sup></td><td>6.5<sup>ab</sup></td><td>6.5<sup>ab</sup></td><td>6.5<sup>ab</sup></td><td>7.0<sup>ab</sup></td><td>6.3<sup>a</sup></td><td>7.5<sup>b</sup></td></tr><tr><th scope="row">Male BW</th><td>6.8</td><td>6.7</td><td>6.8</td><td>7.3</td><td>6.9</td><td>6.6</td><td>7.4</td></tr><tr><th scope="row">Mean BW</th><td>6.8<sup>ab</sup></td><td>6.6<sup>a</sup></td><td>6.7<sup>ab</sup></td><td>6.9<sup>ab</sup></td><td>7.0<sup>b</sup></td><td>6.4<sup>a</sup></td><td>7.4<sup>b</sup></td></tr><tr><th scope="row">SD</th><td>1.1</td><td>1.1</td><td>1.0</td><td>0.9</td><td>0.9</td><td>0.9</td><td>1.0</td></tr></tbody><tbody><tr><th scope="row">Weaning weight (WW)</th><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><th scope="row">Female WW</th><td>23.5<sup>ab</sup></td><td>26.6<sup>b</sup></td><td>25.2<sup>bc</sup></td><td>26.8<sup>b</sup></td><td>29.8<sup>c</sup></td><td>22.2<sup>a</sup></td><td>27.9<sup>bc</sup></td></tr><tr><th scope="row">Male WW</th><td>21.8<sup>a</sup></td><td>25.5<sup>ab</sup></td><td>23.9<sup>ab</sup></td><td>25.1<sup>ab</sup></td><td>27.3<sup>bc</sup></td><td>22.8<sup>a</sup></td><td>26.3<sup>bc</sup></td></tr><tr><th scope="row">Mean WW</th><td>22.9<sup>ab</sup></td><td>25.9<sup>b</sup></td><td>24.7<sup>ab</sup></td><td>25.9<sup>b</sup></td><td>28.3<sup>bc</sup></td><td>22.5<sup>a</sup></td><td>26.7<sup>bc</sup></td></tr><tr><th scope="row">SD</th><td>2.5</td><td>3.2</td><td>1.9</td><td>3.3</td><td>3.7</td><td>3.8</td><td>3.5</td></tr></tbody></table>

abcd: means bearing a different superscript differ within rows at the p<0.05 level

<table><caption>**Table 4:** Correlation coefficient estimates among fleece weight and fiber characteristics</caption><thead><tr><th>Traits</th><th>SL</th><th>FD</th><th>FDcv</th><th>F30</th><th>FC</th><th>CUR</th><th>CURsd</th><th>MD</th></tr></thead><tbody><tr><th scope="row">FW</th><td>0.42**</td><td>0.22**</td><td>- 0.14**</td><td>0.20**</td><td>- 0.24**</td><td>- 0.21**</td><td>- 0.12**</td><td>0.16*</td></tr><tr><th scope="row">SL</th><td></td><td>- 0.12**</td><td>0.18**</td><td>0.25**</td><td>- 0.18**</td><td>- 0.28**</td><td>- 0.17**</td><td>- 0.14</td></tr><tr><th scope="row">FD</th><td></td><td></td><td>- 0.34**</td><td>0.59**</td><td>- 0.55**</td><td>- 0.63**</td><td>- 0.53**</td><td>0.68**</td></tr><tr><th scope="row">FDcv</th><td></td><td></td><td></td><td>0.28**</td><td>- 0.12**</td><td>0.16**</td><td>0.30**</td><td>0.06</td></tr><tr><th scope="row">F30</th><td></td><td></td><td></td><td></td><td>- 0.79**</td><td>- 0.53**</td><td>- 0.36**</td><td>0.55**</td></tr><tr><th scope="row">FC</th><td></td><td></td><td></td><td></td><td></td><td>0.45**</td><td>0.32**</td><td>- 0.46**</td></tr><tr><th scope="row">CUR</th><td></td><td></td><td></td><td></td><td></td><td></td><td>0.83**</td><td>- 0.35**</td></tr><tr><th scope="row">CURsd</th><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>- 0.23**</td></tr></tbody></table>

* significant association (p<0.05); ** significant association (p<0.01)

Correlations among the FW, FD, FDcv, and CUR were of a similar magnitude to those found in previous analyses of US alpacas (Lupton et al., 2006; Wuliji, 2012). The mean fiber diameter (FD), minimum and maximum variant, FD variations (FDsd and FDcv), and staple length (SL) measurements in specialty reference fibers are compared in Table 5. The mean FD was significantly lower in vicuna fiber compared to camel and alpaca fibers. The extremely fine alpaca fiber diameter measured 3.2 microns less than camel hair and only one micron coarser than vicuna fiber (p<0.01). The FD measured for the reference vicuna fiber specimens was nearly the same as the saddle fleeces of vicunas examined in Peru (Quispe et al., 2014). However, the FD measured for extremely fine fiber groups of the Victory
Farm herd in the 2016 and 2017 shearings tested within the range (i.e., 14.15 and 14.29 µm) of the fiber diameter variation among the vicunas of South America.

<table><caption>**Table 5:** Comparison of the mean fiber diameter (FD), minimum and maximum variant, FD variations (FDsd and FDcv), and staple length (SL) measurements in specialty reference fibers (camel or vicuna fiber) with extremely fine selection groups (25 %) of the Victory Farm herd for 2016 and 2017</caption><thead><tr><th>Camelid Species</th><th>Numbers</th><th>FD (µm)</th><th>Minimum (µm)</th><th>Maximum (µm)</th><th>FDcv (%)</th><th>SL (mm)</th></tr></thead><tbody><tr><td>Vicuna fiber</td><td>17</td><td>13.07<sup>a</sup></td><td>11.54</td><td>15.29</td><td>22.69<sup>a</sup></td><td>34.7<sup>a</sup></td></tr><tr><td>Extreme fine 2016</td><td>34</td><td>14.15<sup>b</sup></td><td>12.17</td><td>15.05</td><td>28.12<sup>b</sup></td><td>60.6<sup>b</sup></td></tr><tr><td>Extreme fine 2017</td><td>34</td><td>14.29<sup>b</sup></td><td>13.38</td><td>14.99</td><td>29.25<sup>b</sup></td><td>66.9<sup>b</sup></td></tr><tr><td>Camel fiber</td><td>15</td><td>17.43<sup>c</sup></td><td>15.35</td><td>21.09</td><td>31.83<sup>b</sup></td><td>80.0<sup>c</sup></td></tr></tbody></table>

abc: means bearing a different superscript differ within the column at the p<0.01 level

The Victory Farm FD test results from 2016 were compared with results from eight randomly tested alpaca farms in the US (Table 6). The Victory Farm herd's average FD in 2016 was significantly (p<0.01) finer than all of the eight alpaca farms (Wuliji, 2012). The FD of the Victory Farm herd ranked first in micron fineness, which was 4.74 µm, 6.10 µm, 7.82 µm, 8.81 µm, 8.96 µm, 10.26 µm, 12.35 µm, and 12.42 µm lower than the FD tested in the other farm herds, respectively. Blood cell counts and cell type measurements in a selected number of animals are shown in Table 7. There was no difference between the sexes for any of the blood parameters measured and mean blood parameters were within normal ranges.

<table><caption>**Table 6:** Victory Farm (1) FD test results (2016) compared with eight alpaca farms in the US</caption><thead><tr><th>Farms</th><th>1</th><th>2</th><th>3</th><th>4</th><th>5</th><th>6</th><th>7</th><th>8</th><th>9</th></tr></thead><tbody><tr><td>Animals</td><td>164</td><td>81</td><td>104</td><td>202</td><td>65</td><td>27</td><td>239</td><td>91</td><td>58</td></tr><tr><td>FD(µm)</td><td>16.58<sup>a</sup></td><td>21.32<sup>b</sup></td><td>22.68<sup>b</sup></td><td>24.40<sup>c</sup></td><td>25.39<sup>c</sup></td><td>25.54<sup>c</sup></td><td>26.84<sup>c</sup></td><td>28.93<sup>d</sup></td><td>29.00<sup>d</sup></td></tr><tr><td>SE</td><td>0.32</td><td>0.45</td><td>0.39</td><td>0.30</td><td>0.50</td><td>0.78</td><td>0.26</td><td>0.43</td><td>0.53</td></tr></tbody></table>

abc: means bearing a different superscript differ within the row at the p<0.01 level

<table><caption>**Table 7:** White blood cell (WBC) count, hematocrit (HCT), and cell types measured for the Victory Farm herd (2017)</caption><thead><tr><th>Alpacas</th><th>WBC (K/µL)</th><th>NE (%)</th><th>LY (%)</th><th>MO (%)</th><th>EO (%)</th><th>RBC (M/µL)</th><th>HCT (%)</th><th>PLT (K/µL)</th></tr></thead><tbody><tr><td>24</td><td>10.26</td><td>76.59</td><td>6.46</td><td>15.93</td><td>0.90</td><td>13.03</td><td>28.95</td><td>174</td></tr><tr><td>SD</td><td>2.64</td><td>13.63</td><td>5.91</td><td>9.30</td><td>0.48</td><td>1.62</td><td>3.37</td><td>54</td></tr></tbody></table>

NE: neutrophils; LY: lymphocytes; MO: monocytes; EO: eosinophils; RBC: red blood cell count; PLT: platelet count; K = 1 thousand; M = 1 million.
The calculated EPD for six traits, including FW, SL, FD, FDsd, CUR, and FD was in the range from 19 to 76% (Table 8). Heritability and correlations among fleece weight and fiber trait estimates for extremely fine alpaca breeding herds are listed in Table 8. The h² estimate indicated that the FD and CUR were highly (≥ 0.5) heritable traits, but the FW, FDsd, and SL exhibited only moderate levels of heritability (≤ 0.4). These results were closely comparable to published estimates for camelids (Wuliji et al., 2000; Gutierrez et al., 2009). The phenotypic correlation estimates (Table 4) for the FW and SL, and also for the FD and CUR, were in agreement with the current genetic correlation estimation (Table 9).

<table><caption>**Table 8:** Expected progeny difference (EPD) in fleece weight and fiber characteristics in the Victory Farm alpaca herd (Enns & Speidel, 2017)</caption><thead><tr><th>EPD traits</th><th>Average</th><th>SD</th><th>Minimum</th><th>Maximum</th><th>Accuracy %</th></tr></thead><tbody><tr><td>Fleece weight (kg)</td><td>-0.06</td><td>0.04</td><td>-0.07</td><td>0.10</td><td>19 - 66</td></tr><tr><td>Staple length (mm)</td><td>-0.83</td><td>4.65</td><td>-9.71</td><td>12.45</td><td>21 - 71</td></tr><tr><td>Mean curvature (deg/mm)</td><td>0.79</td><td>3.33</td><td>-10.21</td><td>7.51</td><td>24 - 74</td></tr><tr><td>Fiber diameter (μm)</td><td>-0.21</td><td>0.99</td><td>-1.84</td><td>3.94</td><td>25 - 75</td></tr><tr><td>FD standard deviation (μm)</td><td>-0.10</td><td>0.21</td><td>-0.51</td><td>0.77</td><td>22 - 71</td></tr><tr><td>FD rate of change (μm)</td><td>-0.01</td><td>0.09</td><td>-0.20</td><td>0.43</td><td>22 - 76</td></tr></tbody></table>

<table><caption>**Table 9:** Heritability (the values along the diagonal) and correlations among fleece weight and fiber traits estimated for group breeding herds (Enns & Speidel, 2017)</caption><thead><tr><th>Fiber traits</th><th>FW</th><th>FD</th><th>FDsd</th><th>SL</th><th>CUR</th></tr></thead><tbody><tr><td>FW</td><td>0.40</td><td>0.54</td><td>0.36</td><td></td><td></td></tr><tr><td>FD</td><td></td><td>0.65</td><td>0.75</td><td>0.58</td><td>-0.82</td></tr><tr><td>SL</td><td></td><td></td><td>0.39</td><td></td><td></td></tr><tr><td>FDsd</td><td></td><td></td><td></td><td>0.29</td><td>-0.82</td></tr><tr><td>CUR</td><td></td><td></td><td></td><td></td><td>0.50</td></tr></tbody></table>

The efficiency of selection and genetic gain in an extremely fine fiber diameter depends on the strength of heritability as well as correlations among the selection traits, such as the FD, FW, SL, CUR, and threshold-subjective traits. In addition to the strict selection and corrective mating in the herd, a tandem culling for some undesirable traits, such as unfitness, poor reproduction, and high medullation, should support the long-term selection goals.
## Conclusions

For the alpacas on the Victory Farm, the birth weight, weaning weight, and shearing weight were similar to the published camelid literature; however, the fiber weight and fiber characteristics of the Victory Farm herd were lower than typical huacaya and suri alpacas. Specifically, the fiber diameter, staple length, softness, and coat color traits appeared more similar to domesticated vicuna-like characteristics. The heritability, genetic, and phenotypic correlations among fleece and fiber traits were moderate to high for the major selection traits. This study indicated that strict selection and corrective breeding in the Victory Farm herd was effective for genetic gain in the fiber diameter to establish an extreme fine fiber alpaca strain.

**Acknowledgments.** The author would like to express his sincere appreciation for Don, Gloria, Seth, and Ashlee Smith for generously providing their alpaca herd data records, fiber specimens, and blood samples and for supporting this research and presentation. The author would like to thank Liga Wuri, for fiber measurements of vicuna and camel fiber using the microscopic method, and Amy Bax, for the alpaca blood cell count and cell type measurements.

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Production performance, repeatability and heritability estimates for live weight, fleece
weight and fiber characteristics of alpacas in New Zealand. *Small Ruminant Research*,
37: 189-201
# Merino Breeding Program Improves Wool Quality in US Wool Sheep Flocks

T. Wuliji¹, L. Wuri², H. Glimp³ and T. Filbin³

¹Lincoln University, Jefferson City, MO 65102, USA

E-Mail: WulijiT@LincolnU.edu

²University of Missouri, Columbia, MO 65102

³Rafter Seven Merinos, Inc., PO Box 717, Eureka, NV 89316

**Abstract.** A Merino breeding resource flock was established at Rafter 7 Ranch through cooperation with the University of Nevada and The Edwin L. Wiegand Trust. Initially, 500 Rambouillet ewes were purchased from two established breeders. These ewes were bred naturally or by artificial insemination (AI) to imported Australian Merino rams and to rams selected from within the flock. Over 16 years, 16 rams and semen from 41 Australian Merino rams were imported from Australia. Selection was based on objective wool measurements and phenotypic performance traits. The wool fiber diameter of the current flock decreased by 3 microns on average compared with the foundation flock (23 microns). The flock was expanded to 1,300 ewes and wasbred in 30 single-sire mating groups in 2006. The Merino flock is managed in two breeding lines, one as a registered Merino flock (n=650) and the other as a Rafter 7 line (n=650), which is selected for wool, lambing, and meat production traits. Sheep producers from 18 states, Canada, and Mexico have purchased breeding rams and ewes from Rafter 7 Ranch. The resource flock has made significant progress over the foundation ewe flock during the upgrading phase in major selection traits and is disseminating Merino genes in US wool sheep flocks.

**Resumen.** Se creó un rebaño de recursos para mejora de Merino en el Rancho Rafter 7 en cooperación con la Universidad de Nevada y el Edwin L. Wiegand Trust. Inicialmente, se compraron 500 ovejas Rambouillet de dos criadores reconocidos. Estas ovejas se reprodujeron por monta natural o inseminación artificial (IA) procedente de carneros merinos australianos importados y de carneros seleccionados del propio rebaño. Durante 16 años se importaron 16 carneros y semen de otros 41 carneros merinos australianos. La selección se basó en mediciones objetivas de lana y caracteres de rendimiento. El diámetro de la fibra de la lana actualmente fue 3 micras inferior en promedio al de los animals fundadores (23 micras). El rebaño se expandió hasta 1.300 ovejas y la reproducción se realize en 30 grupos con un único carnero en 2006. La reroducción se realiza independientemente en dos líneas, una registrada como rebaño Merino (n=650) y la otra como Rafter 7 (n=650), y la selección se basa en caracteres de lana, partos y producción de carne. Otros productores de 18 estados, Canadá y México han comprado carneros y ovejas del Rancho Rafter 7. El rebaño de recursos para mejora de Merino ha logrado un progreso significativo en relación a los animals fundadores en los principales caracteres de selección y difunde genes de raza Merino en rebaños de ovejas de lana de Estados Unidos.
**Keywords:** merino sheep, breeding program, wool quality

**Introduction**

In the United States, sheep and lambs are raised primarily in small farm flocks in the Midwest and East, and on large ranching operations in the West. Sheep grazing in the western rangelands can be profitable and environmentally sustainable (Glimp & Swanson, 1994). Most of the 7 million head of sheep in the country is in wool/meat dual-purpose flocks. However, over the last two or three decades, the wool clips in areas of major production have become progressively finer, while textile technology has improved for processing superfine wool types. Consequently, the raw wool premium prices set for fine and superfine wool categories increased significantly over other types of wool during those years. Two decades ago, the University of Nevada-Reno and Rafter 7 Ranch established a Merino breeding program at Rafter 7 Ranch near Yerington, Nevada. The purpose was to introduce superior fine wool Merino genetics from Australia and provide genetically improved and adapted breeding rams and ewes for the US western range regions. All animals born on the ranch are provided an individual metal ear tag at birth and an electronic ear tag number incorporated into their scrapie tag at weaning at approximately 90-120 days of age. Sheep numbers have steadily increased to meet the needs for breeding, replacements, and distribution of breeding stock to sheep producers. The flock is physically inventoried annually at breeding (approx. Nov. 20), shearing (approx. March 20), and lamb weaning/ewe culling (approx. Aug. 15). This paper describes a wool sheep selection program at Rafter 7 Ranch and the impacts of Merino genetics dissemination into western US range sheep flocks. Animal performance and wool characteristics were analyzed and presented for two selection flocks, namely, Rafter 7 Merino and Rafter 7 Line.

**Materials and Methods**

*Range and Pasture*

A flock of 1,300 breeding ewes (Rambouillet and Merino crossbred) and 35 stud rams are maintained at Rafter 7 Ranch, a University of Nevada-Reno (UNR) cooperative sheep station owned by The Edwin L. Wiegand Trust. The ranch includes 1,400 hectares of private land and grazing permits on 40,500 hectares of Bureau of Land Management Lands, and 1,800 hectares of USDA forestland. The flat pasture elevation is 1,200 to 1,500 m, with the high desert range elevation extending to 3,000 m. The annual precipitation within the area is less than 200 mm, mostly as winter snowfall, with unpredictable frosts and wind patterns. Desert shrubs include black greasewood, basin big sagebrush, black sagebrush, bud sage, white sage, and ephedra. Grass species include Indian ricegrass, bottlebrush
squirreltail, and cheatgrass. The established pastures were primarily tall fescue, overseeded with ladino clover. Improved irrigated pastures include a mix of tetraploid perennial ryegrass, improved fescue cultivars, and a variety of alfalfa and ladino clover. An additional 50 hectares of irrigated land is used for alfalfa hay production and aftermath grazing. Irrigated pastures, 35 pastures of 2-6 hectares, are set-stocked during breeding and lambing, with an intensive rotation during the rest of the grazing season.

## Animal Breeding and Ram Distribution

Natural mating and AI were used alternately during the upgrading phases. A computerized record and database program that includes individual animal pedigree, sex, birth date, birth and rearing rank, weaning, and yearling performance is maintained on the ranch. Two seasonal lambing management options were adopted since 2006, although the majority of lambs are scheduled to be born during spring lambing. Animal selection was made each year prior to the breeding season on a multi-trait performance index in conjunction with independent culling for undesirable traits, such as poor conformation and structure, wool face cover, jaws, infertility, and colored fibers. Animal selection was based on objective wool measurements as well as subjective assessment, growth rate, and reproductive performance traits. The Merino flock was fully inspected, pedigreed, and registered with the Texas Delaine-Merino Record Association. Flock management is in two breeding lines, one as a registered Rafter 7 Merino flock (n=650) and the other (Merino x Rambouillet) as the Rafter 7 Line (n=650), both of which are selected for high fleece weight and quality, twinning, and growth traits. The spring lambing flock was wintered on desert rangelands and grazed on irrigated pasture from shearing through lambing and early weaning. Lambs were subjected to preselection culling at weaning and final selection based on yearling performance, including body and fleece weight as well as wool characteristics. A final selection performance index was derived by various adjustment weightings made to birth and rearing ranks, age, body weight, weight gain, fleece weight, fiber diameter, and length. Ram distribution catalogs for selected rams and ewes with a comparative performance index were posted to potential sheep producers/clients two to four weeks before the ranch held a ram sale field day. Selected rams and ewes were presented along with their IDs, pedigree and yearling performance data sheets, and health certificates in subdivided pens on the field day. Typically, about 70-100 producers participated in the annual ram sale, and several dozen ranchers purchased their choice of breeding rams and ewes by open bidding at the field day auction. The genetic distribution and impact on range wool sheep production was monitored on a number of ranches that consistently used Rafter 7 Ranch rams. Two ranches' fleece tests and breeding animal replacement data were monitored and shared as client feedback information. Four of these associated ranches (located in Nevada at Reno, Ely, Fernley, and at Rafter 7 Ranch) were surveyed for
their superfine category (19 microns or less) wool lot weight ratios in the clips,
using the wool warehouse records and public auction information from 2004 to
2009 wool sale catalogs.

Wool Production and Clip Preparation

Individual fleece weight and wool characteristics were recorded for the lifetime of each breeding ewe and ram. Pre-shearing, mid-side wool staples were collected from each sheep, and a set of programmed wool tests for wool characteristics, including average fiber diameter (FD), fiber diameter variation coefficient (FDcv), average staple fiber length (SL), and estimated comfort factor (CF) were measured using an Optical-based Fibre Diameter Analyser OFDA 2000 (BSC Electronics, Western Australia) instrument. Shearing was scheduled at least four weeks prior to lambing. Fleece were classed according to the pre-shear test classification with some subjective alternatives, such as length, discoloration, or tensile strength. Wool clip volumes and sale values were recorded and presented for each of the five years. At the 2009 spring shearing, approximately 3,000 fleeces at Rafter 7 Ranch were pretested, shorn, and classed into five category lots, namely, ultrafine, superfine, fine, medium, and coarse lines. These were baled and transported to the wool warehouse, which provided a sale lot test certification.

Measurements and Statistics

Data was recorded for each animal in the selection flock, including post-shearing
body weight (BW), greasy fleece weight (FW), and wool characteristics. Wool
characteristics, including FD, FDcv, CF, and SL were measured using an OFDA
2000 instrument on pre-shearing, mid-side wool staple samples at the UNR Wool
Research Lab (Reno). Lamb fleeces were classed separately from the adult fleeces
into three classes of ultrafine ( < 17 µm) superfine ( < 19 µm), and fine ( < 21 µm)
wool lots, whereas mixed-age sheep fleeces were subjectively examined at wool
skirting tables and pooled into the appropriate classes according to the pre-
shearing fleece test results. The post-shearing body weight, fleece weight, and wool
characteristics of two flocks (2009, n=2,218) were analyzed. In addition, the annual
fleece FD was measured for 600 ewes from two breeding flocks, which were born
between 2001 and 2004. Their fleece FD was monitored for five years of shearing
within the period from 2002 to 2009. The changes in FD by age were available for
563 animals every year from the first (lamb wool) to the fifth shearing (Age I, II,
III, IV, and V), consecutively. These data were analyzed using a linear regression
equation and a calculated fleece FD change prediction formula (index) for lifetime
wool classing. The FD change prediction indexes were calculated by subgrouping
animals from FD classes into fiber diameter ranges corresponding to US standards
for grades of wool (USDA, 1968). The SAS® (SAS Institute Inc., Cary, NC)
procedures GLM, CORR, REG, and GLIMIX were followed for the data analysis
of the body weight, fleece weight, fiber characteristics, and fiber diameter variations relative to birth year and age. Heritability (h²) and genetic correlation coefficients were cited from the previous analysis in the flocks by a multi-trait animal model statistical procedure (Rauw et al., 2010).

## Results and Discussion

Summary data of body weight, fleece weight, and wool characteristics are shown in Table 1. There are significant (p<0.05) differences between the two selection lines for BW and FW. The pure Merinos produced more wool than the Rafter 7 Line, even though their wool was slightly finer. The Rafter 7 Line sheep were significantly heavier (p<0.05) than the Merino sheep. The mean greasy fleece weight of both the Rafter 7 Merino and Rafter 7 Line flocks were higher than in the published data (Lee et al., 2000) of typical Rambouillet ewes. The earlier analysis of Merino crossbred ewes showed that wool fiber density, clean wool yield, staple length, and greasy fleece weight increased by 41 % per unit area of skin, 15 %, 2.5 cm, and 1.14 kg per head shorn, respectively (Glimp, 2006). Over the years of upgrading, the Rafter 7 Line has closed the gap with the Merinos in average fiber diameter, while still maintaining a significant advantage in BW and other subjective traits, such as having a larger body frame and structure. However, no differences were found between the FD, FDcv, SL, and CF of the flocks.

<table><caption>**Table 1:** Means of body weight (BW), fleece weight (FW), fiber diameter (FD), fiber diameter variation (FDcv), staple length (SL), and comfort factor (CF) in flocks (2009)</caption><thead><tr><th></th><th>No Obs.</th><th>BW (kg)</th><th>FW (kg)</th><th>FD (µm)</th><th>FDcv (%)</th><th>SL (mm)</th><th>CF (%)</th></tr></thead><tbody><tr><td>R7 Merino</td><td>1291</td><td>66.5<sup>b</sup></td><td>5.32<sup>a</sup></td><td>19.4</td><td>17.2</td><td>86.0</td><td>99.0</td></tr><tr><td>R7 Line</td><td>1947</td><td>72.2<sup>a</sup></td><td>4.63<sup>b</sup></td><td>19.5</td><td>17.4</td><td>82.0</td><td>98.9</td></tr><tr><td>SE</td><td></td><td>0.9</td><td>0.9</td><td>0.1</td><td>0.05</td><td>0.5</td><td>0.05</td></tr></tbody></table>

ab: Column means with different superscript letters are different (p < 0.05).

Changes in the FD were shown to exist among ages and birth year (p<0.05) (Table 2), which reflected environmental variations and the interaction of birth year and age groups.
<table><caption>**Table 2:** Least squares means of the average fiber diameter of Rafter 7 Ranch breeding ewes by flock and age group (n = 556)</caption><thead><tr><th rowspan="2">Flock</th><th rowspan="2">Means/<br>Flock</th><th colspan="5">Means by Age Group</th><th rowspan="2">SE</th></tr><tr><th>Age I</th><th>Age II</th><th>Age III</th><th>Age IV</th><th>Age V</th></tr></thead><tbody><tr><td>R7 Meri-<br>no</td><td>20.5</td><td>18.4<sup>e</sup></td><td>20.9<sup>c</sup></td><td>20.7<sup>c</sup></td><td>21.4<sup>a</sup></td><td>21.4<sup>a</sup></td><td>0.1</td></tr><tr><td>R7 Line</td><td>20.6</td><td>18.7<sup>d</sup></td><td>21.1<sup>b</sup></td><td>20.8<sup>c</sup></td><td>21.6<sup>a</sup></td><td>21.5<sup>a</sup></td><td>0.1</td></tr><tr><td>Means</td><td>20.5</td><td>18.5</td><td>21.0</td><td>20.7</td><td>21.5</td><td>21.4</td><td>0.1</td></tr></tbody></table>

abed Means with a different superscript letter differ significantly at the p < 0.05 level within and between rows for age groups; there is no statistical difference between the pooled flock means.

Weaning weight, yearling weight, and fleece weight of the Rafter 7 Ranch sale rams have improved over the years (Table 3). The BW and FW for sale rams of birth years 2005-2007 did not differ between the two lines. The BW measures in sale rams showed a trend of the two breeding flocks converging in terms of BW and FD, although the Merinos produced more and finer wool. Weaning weight of the Rafter 7 Ranch flocks were analyzed previously (Rauw et al., 2007a), which showed that ram lambs weighed more than ewe lambs, single-reared lambs were heavier than multi-litter lambs, and lambs born from two-year-old ewes were lighter than lambs born from ewes of other age groups.

<table><caption>**Table 3:** Body weight (BW) at weaning (WW), yearling (YW), 16-months-old (SW), and 10-month-old fleece weight (FW) of the Rafter 7 Ranch sale rams (n = 120/year)</caption><thead><tr><th>Year of<br>Birth</th><th>Flock</th><th>WW<br>(kg)</th><th>YW<br>(kg)</th><th>16-mo SW<br>(kg)</th><th>10-mo FW<br>(kg)</th></tr></thead><tbody><tr><td rowspan="2">2005</td><td>Merino</td><td>30.8</td><td>60.1</td><td>68.5</td><td>2.72</td></tr><tr><td>R7 Line</td><td>31.3</td><td>60.0</td><td>71.7</td><td>2.63</td></tr><tr><td rowspan="2">2006</td><td>Merino</td><td>27.7</td><td>50.9</td><td>75.5</td><td>3.55</td></tr><tr><td>R7 Line</td><td>28.6</td><td>52.7</td><td>78.6</td><td>3.64</td></tr><tr><td rowspan="2">2007</td><td>Merino</td><td>32.7</td><td>67.3</td><td>74.0</td><td>4.09</td></tr><tr><td>R7 Line</td><td>31.4</td><td>72.7</td><td>79.1</td><td>4.09</td></tr></tbody></table>

Pearson correlation coefficients were calculated between body weight, fleece weight, and wool characteristics and are presented in Table 4. There is a significant (p<0.01) high correlation between FD and CF and also high correlations among FW, FD, and SL; a low-to-moderate correlation exists between BW and FW, whereas small or negative (p<0.05) correlations were observed between FDcv and FD, BW, FW, FD, SL, and CF.
<table><caption>**Table 4:** Pearson correlation coefficients (R-values) of body weight (BW), fleece weight (FW), fiber diameter (FD), staple length (SL), and comfort factor (CF) characteristics</caption><thead><tr><th>Traits</th><th>FW</th><th>FDcv</th><th>FD</th><th>SL</th><th>CF</th></tr></thead><tbody><tr><td>BW</td><td>0.20</td><td>- 0.06</td><td>0.21</td><td>- 0.13</td><td>- 0.10</td></tr><tr><td>FW</td><td></td><td>- 0.20</td><td>0.64</td><td>0.45</td><td>- 0.36</td></tr><tr><td>FDcv</td><td></td><td></td><td>- 0.16</td><td>- 0.09</td><td>- 0.16</td></tr><tr><td>FD</td><td></td><td></td><td></td><td>0.31</td><td>- 0.70</td></tr><tr><td>SL</td><td></td><td></td><td></td><td></td><td>- 0.13</td></tr></tbody></table>

Note: all R-values are significant at p < 0.01.

The heritability and genetic correlative values are listed in Table 5. The h² value was low to moderate for BW, SL, and FW but high for the FD and FDcv traits. The h² and genetic correlations among wool traits were generally lower but comparable to the previous findings in fine wool breeds by Wuliji et al. (2001) and Safari et al. (2005).

<table><caption>**Table 5:** Heritability and genetic correlations for body weight (BW) change, fiber diameter (FD), fiber variation (FDcv), staple length (SL), and greasy fleece weight (FW) estimated in Rafter 7 flocks (Rauw et al., 2010).</caption><thead><tr><th>Traits</th><th>BW</th><th>FD</th><th>FDcv</th><th>SL</th><th>FW</th></tr></thead><tbody><tr><td>BW</td><td>0.29</td><td>- 0.23</td><td>- 0.21</td><td>0.17</td><td>- 0.21</td></tr><tr><td>FD</td><td></td><td>0.51</td><td>0.49</td><td>0.37</td><td>0.21</td></tr><tr><td>FDcv</td><td></td><td></td><td>0.50</td><td>- 0.05</td><td>0.29</td></tr><tr><td>SL</td><td></td><td></td><td></td><td>0.39</td><td>0.34</td></tr><tr><td>FW</td><td></td><td></td><td></td><td></td><td>0.36</td></tr></tbody></table>

The sheep breeding, wool selection, and genetic resource distribution of the Rafter 7 Ranch flock impacts and the breeding stock distribution for western range sheep flocks (Wuliji et al., 2007) and the grazing efficiency of the ewes on the range were discussed previously (Rauw et al., 2007b). The genetic merit distribution and impacts on western range wool sheep breeding was presented for Borda's Ranch, Nevada, operation (Table 6). The number of two-year replacement ewes tested for fiber traits exponentially increased from 2003 to 2005 and remained high to 2008; however, the average FD of selected animals decreased significantly, from 23.9 µm to 20.9 µm in six years, whereas the CF and SL increased.
<table><caption>**Table 6:** Fleece measurements of two-year-old replacement ewes at the Borda's Ranch</caption><thead><tr><th>Year</th><th>No. of Sheep</th><th>FD (µm)</th><th>FDcv (%)</th><th>Comfort Factor (%)</th><th>SL (mm)</th><th>Mean Fiber Ends (µ)</th></tr></thead><tbody><tr><td>2003</td><td>292</td><td>23.9</td><td>16.2</td><td>92</td><td>70</td><td>23.5</td></tr><tr><td>2005</td><td>1457</td><td>22.4</td><td>16.0</td><td>96</td><td>77</td><td>22.1</td></tr><tr><td>2006</td><td>1252</td><td>21.6</td><td>16.0</td><td>97</td><td>80</td><td>21.3</td></tr><tr><td>2007</td><td>848</td><td>20.8</td><td>17.6</td><td>98</td><td>75</td><td>20.9</td></tr><tr><td>2008</td><td>626</td><td>20.9</td><td>16.7</td><td>98</td><td>87</td><td>20.7</td></tr></tbody></table>

A summary of two (Borda's Ranch and Volger's Ranch) of the client ranches' wool classing lines and clip production are presented in Table 7. Both ranches produced and classed fleeces into three sale lots, which showed a gradual reduction in FD and an increase in fine class wool lot volumes, even eliminating the coarse line lot in later years.

<table><caption>**Table 7:** Annual wool class FD lines and portions of clip changes in Rafter 7 Ranch client flocks in shearing and wool clips (2004 – 2008)</caption><thead><tr><th rowspan="2">Wool Class</th><th colspan="5">Fiber diameter of wool lots (µm)</th><th colspan="5">Portion of wool clips (%)</th></tr><tr><th>2004</th><th>2005</th><th>2006</th><th>2007</th><th>2008</th><th>2004</th><th>2005</th><th>2006</th><th>2007</th><th>2008</th></tr></thead><tbody><tr><td colspan="11">Borda's Ranch (NV) wool flock records</td></tr><tr><td>Fine</td><td>21.0</td><td>20.1</td><td>19.7</td><td>19.3</td><td>20.4</td><td>15</td><td>15</td><td>20</td><td>32</td><td>35</td></tr><tr><td>A</td><td>22.9</td><td>22.4</td><td>22.4</td><td>21.8</td><td>22.2</td><td>79</td><td>59</td><td>43</td><td>38</td><td>39</td></tr><tr><td>C</td><td>25.0</td><td>24.8</td><td>24.2</td><td>22.3</td><td>23.7</td><td>6</td><td>26</td><td>37</td><td>30</td><td>26</td></tr><tr><td colspan="11">Volger's Ranch (NV) wool flock records</td></tr><tr><td>Fine</td><td>20.3</td><td>20.6</td><td>20.6</td><td>20.5</td><td>20.3</td><td>31</td><td>29</td><td>58</td><td>35</td><td>30</td></tr><tr><td>A</td><td>22.0</td><td>22.4</td><td>22.0</td><td>21.6</td><td>22.1</td><td>39</td><td>31</td><td>35</td><td>65</td><td>70</td></tr><tr><td>C</td><td>23.0</td><td>23.1</td><td>22.4</td><td>-</td><td>-</td><td>30</td><td>19</td><td>7</td><td>0</td><td>0</td></tr></tbody></table>

Sheep flock performance and wool sale information show consistent improving trends within Rafter 7 Ranch flocks and associated clients' flocks. Wool sales from Rafter 7 Ranch increased significantly in clean wool clip weight and price values (Figure 1).
<figure><img src="image_41.png" /><figcaption>**Figure 1:** Rafter 7 Ranch clean wool clip weight (kg) and value (US dollar).</figcaption></figure>

The gradual and continual increase in superfine and fine wool ratios of the clips at Rafter 7 Ranch was consistent with an earlier within-flock analysis (Wuliji et. al., 2008). For eight-year production performance, the Rafter 7 Ranch wool clips received the highest price of any wool grown in the US. Sheep producers in 18 states, including Nevada, and in Mexico and Canada have regularly purchased breeding rams and ewes from Rafter 7 Ranch. Approximately 1,000 breeding rams and 500 replacement ewes were distributed to range flocks in the western states in the 2000s, which made a notable improvement in the fleece weight, fiber diameter, and yield in the clients' flocks (Wuliji et al., 2009). Superfine lot portions of wool clip sales at four farms in Nevada are shown in Figure 2.

<figure><img src="image_42.png" /><figcaption>**Figure 2:** The superfine ratio (%) of wool clips on four farms (in Reno, Ely, Fernley, and at Rafter 7 Ranch) by production years (2004 – 2009).</figcaption></figure>

The superfine wool ratio increases in the ranches wool sales is associated with the dissemination of the introduced Merino genetics of Rafter 7 Ranch into their sheep flocks. This has produced an improvement in wool quality by increasing the superfine ratio of the wool clip in the associated sheep producers' ranches, with an especially rapid rise in superfine production at Rafter 7 Ranch. This trend is expected
to strengthen a long-term competitive advantage for the western US states' wool sheep enterprises.

The class of FD ranges, standard wool grades, means, and the prediction formula for mixed-age ewes for lifetime fleece classing and assigned wool class color tag codes are listed in Table 8. The FD changes by age groups were small to moderate, which is similar to the pattern of increase in the fleece weight, which showed a larger increase from the first shearing to the second but small changes until four years of age. Such features of FD and FDcv, and inter-trait correlations were also observed in ultrafine Merino flocks (Wuliji et al., 1999). These indexes appeared reasonable and capable of predicting future fleece FD ($\tilde{Y} = a + bx$) using a single FD test ($x$) of individual animals at two years old.

It is recommended that the formula derived for the particular wool grade or FD range be used; however, if there is a limited flock size or operation, flock mean formulas are more suitable to cover a wide range of FDs in small flocks. The $X$ values (the second shearing FD value) were highly and significantly ($p < 0.01$) correlated to the recorded and derived dependent variates ($Y1$ and $Y2$). With 95 % confidence, the fit plot analysis ($Y1$ and $Y2$) indicated a coefficient of determination expressed by the R-squared value of 0.53 and 0.71, which are equal to the $x$ and $y$ correlative associations at 74 % and 85 % ($r = 0.73$ and 0.84), respectively. The FD prediction formula was used for Rafter 7 Ranch commercial flock wool classing operations. Several wool sheep ranches, including Borda Ranch, had adopted the one test for lifetime wool classification procedures using an OFDA analysis of the FD for their wool clip classing or breeding ewe replacements.

The selection efficiency in premium wool characteristics and rapid genetic gain were reported for various wool breeding demonstration flocks (Wuliji et al., 1999; Swan & Purvis, 2005; Brien et al., 2005). These characteristics showed moderate-to-high heritability (Atkins, 1997; Okut et al., 1999; Wuliji et al., 2001; Hansford et al., 2004). Therefore, we predict an increased likelihood of a higher rate of genetic dissemination into commercial sheep flocks followed by rapid genetic gains in wool quality traits.
<table><caption>**Table 8:** US wool grade, FD micron range (µm), wool class code (ear tag color), and the derived FD prediction formula for lifetime fleece classes (mixed-age ewes)</caption><thead><tr><th>Wool Grade</th><th>FD Range</th><th>Class Code</th><th>N</th><th>Mean X</th><th>Mean Y1</th><th>Mean Y2</th><th>y1 = a + bx</th><th>y2 = a + bx</th></tr></thead><tbody><tr><td>80s</td><td>&lt; 19.14</td><td>White</td><td>63</td><td>18.4</td><td>19.3</td><td>19.1</td><td>6.14 + 0.72x</td><td>4.61 + 0.79x</td></tr><tr><td>70s</td><td>19.15-20.59</td><td>Green</td><td>93</td><td>19.7</td><td>20.4</td><td>20.2</td><td>11.72 + 0.44x</td><td>8.79 + 0.58x</td></tr><tr><td>64s</td><td>20.60-22.04</td><td>Orange</td><td>152</td><td>20.6</td><td>20.9</td><td>20.8</td><td>7.63 + 0.65x</td><td>5.72 + 0.74x</td></tr><tr><td>62s</td><td>22.05-23.49</td><td>Yellow</td><td>112</td><td>21.6</td><td>21.6</td><td>21.5</td><td>6.46 + 0.70x</td><td>4.84 + 0.78x</td></tr><tr><td>60s</td><td>23.50-24.94</td><td>Red</td><td>94</td><td>22.6</td><td>22.6</td><td>22.5</td><td>8.45 + 0.63x</td><td>6.34 + 0.72x</td></tr><tr><td>58s</td><td>24.95-26.39</td><td>Black</td><td>49</td><td>23.9</td><td>23.0</td><td>23.2</td><td>20.11 + 0.12x</td><td>15.1 + 0.34x</td></tr><tr><td>Flock Mean Formula</td><td></td><td></td><td>563</td><td>21.0</td><td>21.3</td><td>21.2</td><td>7.16 + 0.67x</td><td>5.37 + 0.75x</td></tr></tbody></table>

Wool grade (80s): based on the spinning count of one pound clean wool that yields 80 hanks (or 44,800 yards); Mean X: the second shorn fleece FD values; Mean Y1: the average FD of three years of tests; Mean Y2: the average FD of four years of tests; y1: the predicted FD estimated by Y1; y2: the predicted FD estimated by Y2; a: intercept value; b: slope value; x: the second shearing fleece's FD.
## Conclusions

The Rafter 7 Ranch Merino flocks have made significant progress during the crossbreeding and upgrading phase as well as in the objective selection of major selection traits, including fleece weight and fiber diameter. The ranch is now disseminating elite genetics to many western range sheep flocks. The dissemination of introduced Merino genetics in the western range sheep flocks have improved wool quality and clip profits, which will strengthen a long-term competitive advantage for the US wool and sheep production sectors.

**Acknowledgments.** The Rafter 7 Ranch Sheep Breeding Program was sponsored by The Edwin L. Wiegand Trust and University of Nevada, Reno, Nevada. We would like to express our sincere appreciation to the western US region sheep breeders for their support and collaboration during establishment of the Rafter 7 Merino and Rafter 7 Line breeding flocks.

## References

Atkins, K.D., 1997, Genetic Improvement of Wool. In "The Genetics of Sheep," edited by L. Piper and A. Ruvinsky. CAB International, UK, p. 471-504

Brien, F.D., Jaensch, K.S., Grimson, R.J., Kemper, K.E., Smith, D.H., Hebart, M.L., Ramsay, A.M.M., 2005, Selection demonstration flocks - what have we learnt? Proceedings of the Association for the Advancement of Animal Breeding and Genetics, 16: 322-325

Glimp, H., 2006, The Rafter 7 Ranch sheep research and education program - Sixteen years of progress. In "The Progress Report: Edwin L. Wiegand Trust," Reno, Nevada, p.1-15

Glimp, H.A., Swanson, S.R., 1994, Sheep grazing and riparian and watershed management. Sheep Research Journal, Special Issue: 65-71

Hansford, K.J., Van Vleck, L.D., Snowder, G.D., 2004, Estimates of genetic parameters and genetic changes for reproduction, weight, and wool characteristics of Rambouillet sheep. Small Ruminant Research, 57: 175-186

Lee, J.W., Waldron, D.F., Van Vleck, L.D., 2000, Parameter estimates for number of lambs born at different ages and for 18-month body weight of Rambouillet sheep. Journal of Animal Science, 78: 2086-2090

Okut, H., Bromley, C.M., Van Vleck, L.D., Snowder, G.D., 1999, Genotypic expression at different ages: II. Wool traits of sheep. Journal of Animal Science, 77: 2366-2371

Rauw, W.M., Glimp, H.A., Jesko, W., Gomez-Raya, L., 2007a, Weaning weights in a range purebred Merino and crossbred Merino x Rambouillet flock. Sheep & Goat Research Journal, 22: 1-6.

Rauw, W., Glimp, H.A., Jesko, W., Sandstrom, M. Gomez-Raya, L., 2007b, Grazing efficiency in free range Merino sheep. Journal of Animal Science, 85 (Suppl.1): 634

Rauw, W.M., Thain, D.S., Teglas, M., Wuliji, T., Sandstrom, M.A., Gomez-Raya, L., 2010, Adaptability of pregnant Merino ewes to the cold desert climate in Nevada. Journal of Animal Science, 88: 860-870
Safari, E., Forgarty, N.M., Gilmour, A. R., 2005, A review of genetic parameter estimates for wool, growth, meat and reproduction traits in sheep. *Livestock Production Science*, 92, no. 3: 271-289

Swan, A.A., Purvis, I.W. 2005, Genetic progress in the T13 Merino breeding program. *Proceedings of the Association for the Advancement of Animal Breeding and Genetics*, 16: 165-168

USDA, 196, Official standard of the United States for grades of wool top. Consumer and Marketing Service. Last modified December 21, 1968

https://www.ams.usda.gov/sites/default/files/media/WoolStandard%5B1%5D.pdf.

Wuliji, T., Dodds, K.G., Land, J.T.J., Andrews, R.N., Turner, P.R., 1999, Response to selection for ultrafine Merino sheep in New Zealand environment I: Wool production and wool characteristics of an ultrafine fibre diameter selected and control hoggets. *Journal of Livestock Production Science*, 58: 33-44

Wuliji, T., Dodds, K.G., Land, J.T.J., Andrews, R.N., Turner, P.R., 2001, Selection for ultrafine Merino sheep in New Zealand – Heritability, phenotypic and genetic correlations of live weight, fleece weight and wool characteristics in yearlings. *Animal Science*, 72, no. 2: 1-10

Wuliji, T., Glimp, H., Perryman, B., Rauw, W., Filbin, T., 2008, Sheep production on high desert rangelands and irrigated pasture. *Proceedings of the XXI International Grassland Congress & VII International Rangeland Congress: Multifunctional Grasslands in a Changing World*, II: 83

Wuliji, T., Glimp, H., Filbin, T., 2009, Introduction of Merino genetics to improve Western range sheep flock wool quality and wool clip profits. *Proceedings of US Sheep Research Programs, American Sheep Industry Association Convention, San Diego, CA*, p. 47-49

Wuliji, T., Glimp, H., Jesko, W., Rauw, W.M., 2007, Introduction to Merino breeding resource flocks at Rafter 7 Ranch in Nevada. *Journal of Animal Science*, 85. (Suppl. 1): 572
<figure><img src="image_43.png" /></figure>
# Selection Strategies for Fiber Quality in Alashan Cashmere Goat

M. Antonini<fnref n="1" />, P.R. Tang<fnref n="2" />, F. Panella<fnref n="3" />, G. Attard<fnref n="4" />, E. Lasagna<fnref n="3" />, S. Ceccobelli<fnref n="3" /> and F.M. Sarti<fnref n="3" />

**Abstract.** Over the last ten years, many authors have attempted to study various genetic patterns and how they relate to improved cashmere fiber production. Unfortunately, the way the sector is currently structured hinders any "on the filed" attempts to apply modern quantitative and molecular techniques to improve fiber characteristics through selection. The process is complicated by the fact that cashmere fiber quality is normally assessed when the animal is approximately a year old. The study investigates the prediction of cashmere quality in adult Alashan cashmere goats from measurements on young animals. Ten fleece samples were collected from 12 kids until 14 months of age (1,441 records). Fiber quality, determined by OFDA, was measured monthly. The statistical analysis indicates that the kid random effect is highly significant ($P \le 0.0001$) throughout the test period and thus confirms a genetic effect on cashmere fleece characteristics; the age effect was also found to be consistently significant. The measured traits showed an increasing trend of mean values up to the 5th -7th month. The fiber girth stabilises by the fifth month reaching a thickness of 14.46 µm. All studied traits show a strong correlation with fiber quality beyond the fifth month. Furthermore, although adult fleece quality can be predicted beyond the fifth month of age, this usually coincides with the warm summer period and hence is an unsuitable season for the evaluation of fleece. Under such situations it may be recommended to postpone the evaluation beyond the age at eight months during the cooler months.

**Resumen.** En los últimos diez años, muchos autores han intentado estudiar diversos patrones genéticos y cómo estos se relacionan con la producción mejorada de la fibra de cashmere. Desafortunadamente, la forma en que está estructurado actualmente el sector dificulta cualquier intento de aplicar técnicas cuantitativas y moleculares modernas "en campo" para mejorar las características de las fibras mediante selección artificial. El proceso se complica porque la calidad de la fibra de cashmere se evalúa normalmente cuando el animal tiene aproximadamente un año de edad. Este estudio aborda la predicción de la

<footnote n="1">¹ENEA Unità Tecnica “Sviluppo Sostenibile ed Innovazione del sistema Agro-industriale”, Via Anguillarese 301, 00123 S.M. di Galeria, Roma, Italy</footnote>

<footnote n="2">²Livestock Improvement Station of Alashan, Inner Mongolia Autonomous District, Alashan Left Banner, 750300, China</footnote>

<footnote n="3">³Dipartimento di Scienze Agrarie, Alimentari e Ambientali, Università degli Studi di Perugia, Borgo XX Giugno 74, 06121, Perugia, Italy</footnote>

<footnote n="4">⁴Department of Rural Sciences and Food Systems, Institute of Earth Systems, University of Malta, Malta</footnote>
calidad de cashmere en cabras adultas Alashan a partir de mediciones en animales jóvenes. Se recogieron diez muestras de lana de 12 cabritos de hasta 14 meses de edad (1441 registros). La calidad de la fibra se valoró mensualmente mediante OFDA. En el análisis estadístico el efecto aleatorio cabrito resultó altamente significativo (P≤0.0001) a lo largo de todo el período, confirmando la presencia de efecto genético sobre las características del vellón de cashmere. El efecto de la edad resultó también fuertemente significativo. Los caracteres medidos mostraron una tendencia creciente de valores medios hasta el 5°-7° mes de edad. La curvatura de la fibra se estabiliza en el quinto mes alcanzando un espesor de 14.46 µm. Además los caracteres estudiados mostraron una fuerte correlación con la calidad de la fibra después del quinto mes. Aunque la calidad del vellón adulto se podía predecir después del quinto mes de edad, este período suele coincidir con el verano, temporada considerada inadecuada para la evaluación del vellón. Por ello, se recomienda aplazar la evaluación a partir de los ocho meses coincidiendo con un período más frío.

**Keywords:** animal breeding, early selection, rural production, cashmere, goat, fiber

**Introduction**

The cashmere goat is to be found over most of Central Asia; including countries such as: Tibet, Mongolia, China (mainly in Inner Mongolia, Qinchai, Xinjiang, Yinghai, Ningxia, Hebei, Shanxi, Shang Dong, Liaoning provinces); and are also common in Afghanistan, Iran, Nepal, Pakistan, India, Turkey and in some regions of the ex-Soviet Union (Uzbekistan, Turkmenistan, Tajikistan, Kazakhstan, Kirghistan). A small population is also to be found in Australia and New Zealand.

Although official cashmere production data is not readily available, the production of raw/greasy fleece quantities can be estimated as follows: People's Democracy of China (8-9,000 tons); People's Republic of Mongolia (2,700-3,000 tons); Iran and Afghanistan (1,500-2,000 tons); Pakistan, Nepal and India (800 tons) and some 80 tons originating from other countries (Turkey, Australia, New Zealand) (http://www.cashmeregoatassociation.org/). The global production of cashmere is estimated at about 15,000 tons and amounts to about 0.2% of the total produced fibers (http://www.naturalfantasy.it/).

The Alashan region is located at an altitude of 1,000-1,400 m above sea level and extends over 670,000 km² of desert area characterised with continental type of climate having an average temperature of 7 °C, with a minimum of -13 °C in January and a maximum of 24 °C in July. The Alashan region produces one of the best cashmere fiber in the People's Democracy of China (Zizhi and Degang, 2006).

Over the last 10 years many authors have studied different genetic aspects of the cashmere goat in an attempt to enhance quality and quantity of fiber production. Some studies using microsatellites to evaluate the ethnological structure of the Chinese cashmere goat populations infer that different goat groups can actually be considered as being separate breeds (Di et al., 2011; Liu et al., 2013) that are highly
to moderately polymorphic (Xu et al., 2010). Others have focused on the associa-
tion between productive traits and genetic markers. Shen et al. (2004) reports a
connection linking body weight, cashmere yield and fineness at certain blood pro-
tein loci. Jin et al. (2006a) also found favourable genotypes at six blood protein loci
for body weight and fiber yield in Liaoning cashmere goat; and obtained compara-
ble results with the use of microsatellites on the same breed (Jin et al., 2006b). The
SNPs association with cashmere production was studied by Lan et al. (2008); and
found higher cashmere yields in TC genotype of a SNP in LALBA gene and estab-
lished (Lan et al. 2009) that there exists a significant effect of 12 SNPs of POU1F1
gene on raw/greasy fleece weight and fiber length in Liaoning and Mongolian
white cashmere goats. Moreover, Lan et al. (2012) found a significant effect of 28
bp insertion on the body mass of yearlings in the Inner Mongolia white cashmere
breed. Pan et al. (2011) proposes the use of the polymorphism in Six6 gene as
suitable marker for fiber length in Inner Mongolia white cashmere goats. Wang et
al. (2012) found that the KAP9.2 gene could be a suitable marker in the selection
of cashmere goat studs.

Other researchers have investigated a variety of correlations linked with pro-
ductive traits. The quantitative approach was used to compare breeding and phe-
notypic value for body weight and fiber yield in Mongolian cashmere goats (Li et
al., 2000; Junyan et al., 2006); and it was also used to estimate the genetic parame-
ter of growth traits in Raini cashmere goat (Barazandeh et al., 2012; Mohammadi et
al., 2012). Other researchers (Zhou et al., 2002) evaluated the genetic parameters of
the production traits in Inner Mongolia Cashmere goats and found that heritability
estimates were 0.28, 0.23, 0.32 and 0.10 for cashmere weight, fiber length, fiber
diameter and live bodyweight respectively. Wang et al. (2012) studied the genetic
parameters that contribute towards fleece traits; and later Wang et al. (2015) inves-
tigated the effects of a procedure of early selection for fleece traits in a monitored
population consisting of 3,257 goats divided in 12 flocks.

Unfortunately, the current industrial structure and set up in the Alashan region
does not allow for the “in the field” application of current quantitative and mo-
lecular techniques to enhance fiber characteristics through the identification and
selection of superior sires. This poor industrial organisation is evident through the
lack of any scientific approach or application in sire / dam evaluation criteria, and
the normal practice of random matings. Furthermore, selection of potential repro-
ducers is also done randomly, sometimes at too early an age when the fiber charac-
teristics quality are still not fully expressed while others when the animals are over
1 year of age. Recently a record of performance and a genetic amelioration pro-
gram on the cashmere goat population in Alashan has been put in place (Valbonesi
et al., 2012). The objectives of this selection program are to evaluate cashmere
diameter and its coefficient of variation.

The aim of this study is to study the probability of predicting cashmere quality
in the adult Alashan cashmere goat from parameters measured on the young ani-
mal.
## Materials and Methods

Whole fleece samples including both the undercoat and out coat were collected from 12 kids on a monthly basis starting in April 2011.

The first sample was obtained at birth (MONTH0) and every month thereafter for the next 13 months (MONTH14). All animals were reared in the same flock and housed at the Animal Improvement Station of Alashan Left Banner in Alashan region.

A total of 1441 records were available for analysis (some records were missing) and the following variables were reported in the original record data file: age at sampling in months (0,...,13); LC: length of cashmere (under coat) (cm); LK: length of hair (outer coat) (cm); D: diameter of cashmere (µm); R: LK/LC ratio; CV: coefficient of variation of D; CF: comfort factor (% of fiber ≤ 30 µm).

Due to practical constraints, only the data related to the first eleven sampling events were considered for evaluation having a total of 1210 records. All the fiber parameters were evaluated with an Optical Fiber Diameter Analyser (OFDA) (Petersson and Gheradi, 1996) at the labs of the Environmental Sciences School of Camerino University (Italy). It is relevant to note that fiber quality improves as LC, LK and CF values increase and D, R and CV values decrease.

All the statistics were computed by SAS (SAS, 2000). The main statistical inference parameters were analysed, and the effects of age and kid were computed according to the following linear model:

$$y_{ijk} = \mu + \text{month}_i + K_j + e_{ijk}$$

Where:

$y_{ijk}$ = experimental observation (LC, LK, D, R, VC, CF);

$\mu$ = overall mean;

$\text{month}_i$ = fixed effect of monthly age (1, ..., 11);

$K_j$ = random/fixed effect of kid (1, ..., 12);

$e_{ijk}$ = residual

The same model was run twice, once setting the K factor as casual and again setting K as fixed effect. In the first case, the K factor is assumed to be the only indicator of the genetic variability, which due to the lack of genealogical information is actually not possible to estimate. In the second case the K factor is considered as fixed so as to estimate the GL means, the option “repeated” on PROC GLM (SAS) was used correctly since each animal had eleven observations according to age. The correlations between the observations at the more mature age (MONTH10) and the younger ages (except for the diameter measures) were computed (PROC CORR, SAS), at all the ages to observe the relationship between the baby cashmere produced at young ages and the regular cashmere produced at the more mature age. The linear, quadratic and cubic regression models were
compared through R² coefficient in order to evaluate the best one to predict fiber quality in the adult goat on the base of younger observations.

## Results and Discussion

The number of kids that were able to be sampled for fleece collection was not always constant throughout the experimental period. In fact Table 1 shows that at the several ages, fiber had a decreasing trend from MONTH0 to MONTH6, followed by an increasing one up to MONTH10. BY MONTH11 only six kids gave suitable samples of fiber. Thus in the analysis only samples from MONTH0 to MONTH10 were considered.

<table><caption>**Table 1:** Number of kids sampled at several ages</caption><thead><tr><th rowspan="2">Month</th><th colspan="6">Traits</th></tr><tr><th>LC</th><th>LK</th><th>R</th><th>D</th><th>CF</th><th>CV</th></tr></thead><tbody><tr><td>0</td><td>12</td><td>12</td><td>12</td><td>12</td><td>12</td><td>12</td></tr><tr><td>1</td><td>12</td><td>12</td><td>12</td><td>12</td><td>12</td><td>12</td></tr><tr><td>2</td><td>10</td><td>11</td><td>10</td><td>12</td><td>12</td><td>12</td></tr><tr><td>3</td><td>2</td><td>10</td><td>2</td><td>4</td><td>4</td><td>4</td></tr><tr><td>4</td><td>7</td><td>10</td><td>7</td><td>11</td><td>11</td><td>11</td></tr><tr><td>5</td><td>7</td><td>5</td><td>5</td><td>8</td><td>8</td><td>8</td></tr><tr><td>6</td><td>6</td><td>6</td><td>6</td><td>6</td><td>6</td><td>6</td></tr><tr><td>7</td><td>12</td><td>10</td><td>10</td><td>12</td><td>12</td><td>12</td></tr><tr><td>8</td><td>12</td><td>12</td><td>12</td><td>12</td><td>12</td><td>12</td></tr><tr><td>9</td><td>12</td><td>12</td><td>12</td><td>12</td><td>12</td><td>12</td></tr><tr><td>10</td><td>11</td><td>11</td><td>11</td><td>12</td><td>12</td><td>12</td></tr><tr><td>11</td><td>6</td><td>6</td><td>6</td><td>6</td><td>6</td><td>6</td></tr></tbody></table>

LC = length of cashmere; LK = length of hair; R = LK/LC ratio; D = diameter of cashmere; CF = comfort factor; CV = coefficient of variation.

The trend described above is most likely the result of the climatic conditions of the region. The kidding season starts in April (MONTH0), subsequent sampling up to MONTH6 correspond chronologically from April to September when temperatures get progressively warmer along the test period. One has to keep in mind that cashmere kids have ample fleece at birth (cashmere baby wool) that is gradually lost on the onset of photoperiod inversion which triggers off the development of new follicular activity that becomes very low in June (2 samples), therefore the ages after MONTH10 were not considered.

Secondary follicular growth was studied by Henderson and Sabine (1991) who established that the greatest development of secondary hair follicles occurred at twenty weeks of age in three groups of Australian cashmere twin goats. This result should indicate that the selection for fiber characteristics could be defined on samples collected between the fifth and tenth week of age.
In the selection of other fiber producing species, such as sheep, fiberwool quality is normally assessed at approximately one year of age during the time of first shearing (Ozcan et al., 2005; Mandal et al., 2009). This strategy requires that all the young males be reared together until this age. Under this situation it is therefore crucial to be able to identify at the most earliest stage those individuals that can accurately predict the adult fiber quality, so as to reduce the holding costs of the flock under evaluation.

In Merino sheep, 5 months could be a suitable age to evaluate fiber for an early selection; as a matter of fact, in Australian Merinos the secondary follicles are fully developed by the 17-18 week (Burns 1949, 1953, 1954a, b; Fraser, 1952, 1954; Schinckel, 1955). Despite that, in Merinos, the 1st shearing at 1 year of age is considered as the suitable age for fiber evaluation in the Merino wool improvement selection program.

In the domestic Camelids of South American (SAC) Antonini et al. (2004) defined in all three types of SAC investigated the fleece structure is established and fiber analysis could be performed as tool for an early criteria of selection on animals that were two months old.

In Table 2 the significance level of the kid (random) and age effect on the studied traits are reported. The kid factor is always highly significant ($P \le 0.0001$), hence it imparts a confirmed genetic effect on the characteristics of cashmere fleece. Similarly to what was reported by others (Wang et al., 2012); the age effect was also always significant, but at a lower level $P$ value in CF. The statistical model worked well on traits LC, LK and D; obtaining high coefficient of determination ($R^2$) values of between 0.6 and 0.7; but had a lower relevance for R, CF and CV (0.34-0.49).

<table><caption>**Table 2:** Significance and $R^2$ of the variability factors in the studied traits</caption><thead><tr><th rowspan="2"></th><th colspan="6">Traits</th></tr><tr><th>LC</th><th>LK</th><th>R</th><th>D</th><th>CF</th><th>CV</th></tr></thead><tbody><tr><th>KID*</th><td>0.0001</td><td>0.0001</td><td>0.0001</td><td>0.0001</td><td>0.0001</td><td>0.0001</td></tr><tr><th>MONTH</th><td>0.0001</td><td>0.0001</td><td>0.0001</td><td>0.0001</td><td>0.0163</td><td>0.0001</td></tr><tr><th>R<sup>2</sup></th><td>0.67</td><td>0.60</td><td>0.43</td><td>0.68</td><td>0.34</td><td>0.49</td></tr></tbody></table>

*: random

LC = length of cashmere; LK = length of hair; R = LK/LC ratio; D = diameter of cashmere;

CF = comfort factor; CV = coefficient of variation.

The mean values of the traits (Table 3) increase progressively with age and tended to stabilise by the age of 4-6 months (MONTH5-MONTH7); after which, the results indicate that there are no further significant improvements. At MONTH0 (at birth) the length of cashmere is 2.28 cm and maintains a steady rate of growth until it tapers off by the age of 5 months (MONTH6) reaching a length of 4.35 cm. A similar trend is observed in LK, starting at 8 cm at MONTH0 and achieves the largest length by the 5th month of age (16.01 cm). This value is very close to the
reported length at 1 year of age as observed by Wang et al. (2015). R achieves a value close to 3 cm by the 4-5 month of age, and could represent the beginning of new cashmere growth corresponding to the loss of baby fiber coat. Fiber diameter (D) reaches 14.46 µm by the third month and remains relatively constant thereafter; similar values were those observed by Wang et al. (2015) in the same breed. Beyond the second month, the CF registers values exceeding 99 % and CV reduces consistently implying that the fiber becomes progressively more uniform and homogeneous. Baby cashmere, which is obtained at an early age (2-4 months), has a diameter of about 13 µm; and is used in the spinning of high quality yarn.

<table><caption>**Table 3:** LSmeans (±SD) of the studied traits at the several ages</caption><thead><tr><th rowspan="2">Month</th><th colspan="6">Traits</th></tr><tr><th>LC</th><th>LK</th><th>R</th><th>D</th><th>CF</th><th>CV</th></tr></thead><tbody><tr><td rowspan="2">0</td><td>2.28 ±</td><td>8.00 ±</td><td>3.67 ±</td><td>14.27 ±</td><td>98.17 ±</td><td>35.80 ±</td></tr><tr><td>0.20</td><td>0.97</td><td>0.34</td><td>0.17</td><td>0.24</td><td>1.32</td></tr><tr><td rowspan="2">1</td><td>2.34 ±</td><td>9.47 ±</td><td>4.35 ±</td><td>13.90 ±</td><td>98.58 ±</td><td>30.89 ±</td></tr><tr><td>0.20</td><td>0.97</td><td>0.34</td><td>0.17</td><td>0.24</td><td>1.32</td></tr><tr><td rowspan="2">2</td><td>2.06 ±</td><td>8.94 ±</td><td>5.59 ±</td><td>13.61 ±</td><td>99.33 ±</td><td>26.87 ±</td></tr><tr><td>0.22</td><td>1.06</td><td>0.37</td><td>0.18</td><td>0.27</td><td>1.45</td></tr><tr><td rowspan="2">3</td><td>3.21 ±</td><td>7.66 ±</td><td>2.68 ±</td><td>13.04 ±</td><td>99.57 ±</td><td>29.15 ±</td></tr><tr><td>0.49</td><td>2.38</td><td>0.83</td><td>0.41</td><td>0.60</td><td>3.26</td></tr><tr><td rowspan="2">4</td><td>3.63 ±</td><td>12.35 ±</td><td>3.59 ±</td><td>14.46 ±</td><td>99.08 ±</td><td>30.84 ±</td></tr><tr><td>0.26</td><td>1.26</td><td>0.44</td><td>0.22</td><td>0.32</td><td>1.73</td></tr><tr><td rowspan="2">5</td><td>4.02 ±</td><td>13.93 ±</td><td>3.77 ±</td><td>14.24 ±</td><td>99.23 ±</td><td>27.90 ±</td></tr><tr><td>0.30</td><td>1.50</td><td>0.52</td><td>0.26</td><td>0.38</td><td>2.06</td></tr><tr><td rowspan="2">6</td><td>4.35 ±</td><td>16.01 ±</td><td>3.82 ±</td><td>14.52 ±</td><td>99.26 ±</td><td>27.21 ±</td></tr><tr><td>0.28</td><td>1.37</td><td>0.48</td><td>0.24</td><td>0.35</td><td>1.45</td></tr><tr><td rowspan="2">7</td><td>4.17 ±</td><td>13.65 ±</td><td>3.26 ±</td><td>14.41 ±</td><td>98.50 ±</td><td>27.81 ±</td></tr><tr><td>0.22</td><td>1.06</td><td>0.37</td><td>0.18</td><td>0.27</td><td>1.45</td></tr><tr><td rowspan="2">8</td><td>5.19 ±</td><td>13.93 ±</td><td>2.84 ±</td><td>14.34 ±</td><td>99.18 ±</td><td>28.40 ±</td></tr><tr><td>0.20</td><td>0.97</td><td>0.34</td><td>0.17</td><td>0.24</td><td>1.32</td></tr><tr><td rowspan="2">9</td><td>4.55 ±</td><td>12.69 ±</td><td>2.94 ±</td><td>14.39 ±</td><td>99.28 ±</td><td>26.97 ±</td></tr><tr><td>0.20</td><td>0.97</td><td>0.34</td><td>0.17</td><td>0.24</td><td>1.32</td></tr><tr><td rowspan="2">10</td><td>5.00 ±</td><td>12.59 ±</td><td>2.57 ±</td><td>14.73 ±</td><td>98.68 ±</td><td>31.10 ±</td></tr><tr><td>0.21</td><td>1.01</td><td>0.35</td><td>0.17</td><td>0.25</td><td>1.38</td></tr></tbody></table>

LC = length of cashmere; LK = length of hair; R = LK/LC ratio; D = diameter of cashmere; CF = comfort factor; CV = coefficient of variation.

The correlations reported in Table 4 were an attempt to identify any potential correlation between final fiber quality and parameters measured on the coat of young kids.
<table><caption>**Table 4:** Correlations between the traits at MONTH10 and at other ages</caption><thead><tr><th colspan="7">MONTH10</th></tr><tr><th>Month</th><th>LC</th><th>LK</th><th>R</th><th>D</th><th>CF</th><th>CV</th></tr></thead><tbody><tr><td>0</td><td>- 0.35</td><td>0.33</td><td>0.24</td><td>0.24</td><td>0.17</td><td>0.07</td></tr><tr><td>1</td><td>- 0.24</td><td>0.31</td><td>0.35</td><td>0.31</td><td>0.26</td><td>0.11</td></tr><tr><td>2</td><td>- 0.15</td><td>0.40</td><td>0.35</td><td>0.35</td><td>0.36</td><td>0.40</td></tr><tr><td>4</td><td>0.54</td><td>0.72</td><td>0.49</td><td>0.51</td><td>0.53</td><td>0.55</td></tr><tr><td>5</td><td>0.78</td><td>0.76</td><td>0.60</td><td>0.61</td><td>0.61</td><td>0.65</td></tr><tr><td>6</td><td>0.82</td><td>0.77</td><td>0.57</td><td>0.72</td><td>0.62</td><td>0.62</td></tr><tr><td>7</td><td>0.77</td><td>0.81</td><td>0.59</td><td>0.75</td><td>0.73</td><td>0.68</td></tr><tr><td>8</td><td>0.05</td><td>0.83</td><td>0.80</td><td>0.81</td><td>0.67</td><td>0.59</td></tr><tr><td>9</td><td>0.70</td><td>1.00</td><td>0.88</td><td>0.72</td><td>0.83</td><td>0.82</td></tr></tbody></table>

LC = length of cashmere; LK = length of hair; R = LK/LC ratio; D = diameter of cashmere; CF = comfort factor; CV = coefficient of variation.

Due to lack of sufficient data in the MONTH3 group, the correlations related to this age were not computed. This lack of data was expected as the MONTH3 coincides with the month of July when the baby cashmere coat is lost and the growth of the new fiber is about to start. All the coefficients become relevant after MONTH4; this situation leads to a practical problem that can have two possible solutions. The kids to be selected for breeding can be identified at 5 months of age, hence the rest of the group can be culled. The main constraint in this solution is that MONTH4 is achieved at the late summer, when potentially some animal could still be carrying the undercoat. Alternatively, selection can be done when the animals are 7 months old, in this option the holding time is extended by a further three months, but this will provide sufficient time for all the animals to express their fiber qualities. To solve this uncertainty, the provisional suitability of linear, quadratic and cubic regression models of MONTH4 and MONTH7 on MONTH10 were estimated for all traits and results presented in Table 5.

<table><caption>**Table 5:** R² (%) of the regression models to predict traits at MONTH10 by traits at MONTH4 and MONTH7</caption><thead><tr><th rowspan="2">MODEL</th><th colspan="12">Traits</th></tr><tr><th>LC</th><th>LK</th><th>R</th><th>D</th><th>CF</th><th>CV</th><th>M4</th><th>M7</th><th>M4</th><th>M7</th><th>M4</th><th>M7</th></tr></thead><tbody><tr><td>Linear</td><td>94</td><td>93</td><td>81</td><td>91</td><td>78</td><td>84</td><td>99</td><td>99</td><td>96</td><td>99</td><td>99</td><td>99</td></tr><tr><td>Quadratic</td><td>84</td><td>84</td><td>72</td><td>83</td><td>57</td><td>68</td><td>99</td><td>99</td><td>92</td><td>95</td><td>99</td><td>99</td></tr><tr><td>Cubic</td><td>73</td><td>71</td><td>64</td><td>76</td><td>37</td><td>52</td><td>98</td><td>99</td><td>86</td><td>88</td><td>99</td><td>99</td></tr></tbody></table>

LC = length of cashmere; LK = length of hair; R = LK/LC ratio; D = diameter of cashmere; CF = comfort factor; CV = coefficient of variation.
The linear regression proved to be the better predictor; in fact, the R² values were almost always greater than 90 %. In the case of trait R except for obvious differences, no real tangible differences are observed between MONTH4 and MONTH7 models; hence the results do not allow for the identification of the best time to measure as an indicator of mature fleece quality. Another indication on the suitability of early age selection is presented in Table 6 that shows correlations between fiber diameters at different ages.

Results indicate that this single trait may be the most important indicator in defining the fleece quality. Table 6 presents the correlations between measurements records at the baby cashmere production stage (from birth till the age of 2 months) in italic while the coefficients between the diameters in the young cashmere stage (at ages from 4-10 months) are indicated in normal fonts. The coefficients between the data at baby and young ages are represented by bold fonts. If the correlations between baby and young are significant and high, it should be possible to assume a very early selection of the best animals. From the general point of interest, it has to be noted that the estimated correlation coefficients are similar to those reported by Wang et al. (2015) in adult Inner Mongolia Cashmere goats.

In the baby phase, the correlations are often significant and range from 0.14 (not significant) to 0.40. In the young phase these coefficients are also often significant with high values (0.97: 4-6 months, 0.84: 8-10 months; 0.81: 6-8 months). The correlations between the baby and young cashmere production ages are not significant and in many cases get values close or over 0.50. These results are encouraging and need to be verified on a larger population to assess the possibility if selection for cashmere fleeces quality at early age.
<table><caption>**Table 6:** Correlation coefficients between the fiber diameter at several ages</caption><thead><tr><th>MONTH</th><th>1</th><th>2</th><th>4</th><th>5</th><th>6</th><th>7</th><th>8</th><th>9</th><th>10</th></tr></thead><tbody><tr><td>0</td><td>0.14</td><td>0.39***</td><td>0.05</td><td>0.50***</td><td>0.17</td><td>0.44***</td><td>0.53***</td><td>0.49***</td><td>0.64***</td></tr><tr><td>1</td><td>1</td><td>0.40***</td><td>0.37***</td><td>-0.36***</td><td>0.53***</td><td>0.35***</td><td>0.41***</td><td>0.40***</td><td>0.61***</td></tr><tr><td>2</td><td></td><td>1</td><td>0.30***</td><td>0.14</td><td>0.46***</td><td>0.41***</td><td>0.62***</td><td>0.21**</td><td>0.66***</td></tr><tr><td>4</td><td></td><td></td><td>1</td><td>-0.008</td><td>0.97***</td><td>0.20*</td><td>0.17</td><td>0.39***</td><td>0.31***</td></tr><tr><td>5</td><td></td><td></td><td></td><td>1</td><td>-0.38**</td><td>0.17</td><td>0.22*</td><td>0.19</td><td>-0.01</td></tr><tr><td>6</td><td></td><td></td><td></td><td></td><td>1</td><td>0.68***</td><td>0.81***</td><td>0.55***</td><td>0.72***</td></tr><tr><td>7</td><td></td><td></td><td></td><td></td><td></td><td>1</td><td>0.60***</td><td>0.59***</td><td>0.75***</td></tr><tr><td>8</td><td></td><td></td><td></td><td></td><td></td><td></td><td>1</td><td>0.53***</td><td>0.84***</td></tr><tr><td>9</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>1</td><td>0.72***</td></tr></tbody></table>

Italic: between baby cashmere ages; **Bold:** between baby and young cashmere ages; Normal: between young cashmere ages. *: P≤0.05; **: P≤0.01; ***: P≤0.001
**Conclusion**

This study represents the first scientific approach to the practice of selection for fiber quality in the Alashan cashmere goat in its local managerial environment. More data is required to generate and validate information that can be applied in practice. Some possible considerations as a result of this investigation are: 1) fiber diameter in the kid (0-2 months) is quite correlated to the fiber diameter in older animals (4-10 months), 2) adult fleece quality can be predicted after the fourth month of age, but this age is usually coincides with the warm summer period, an unsuitable season to evaluate the fleece. Therefore, it may be prudent to postpone the evaluation age to seven months, a time that usually occurs past the summer season. Probably, in the current state of reality, the breeders will to select the animals for reproduction according to their own management condition and constrains.

**Acknowledgments.** Especial thank is due to Loro Piana SpA, one of the most important Italian company in Cashmere goods, for the financial support and to allow the use of the data from its project in Alashan region.

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<figure><img src="image_44.png" /></figure>
# Interaction between ASIP and MC1R in Black and Brown Alpaca

C. Bathrachalam<fnref n="1" />, C. Nocelli<fnref n="1" />, I. Pazzaglia<fnref n="2" />, S. Pallotti<fnref n="3" />, D. Pediconi<fnref n="3" />,
A. La Terza<fnref n="2" /> and C. Renieri<fnref n="3" />

E-Mail: bcm.rna@gmail.com

E-Mail: antonietta.laterza@unicam.it

E-Mail: cristina.nocelli@unicam.it,

**Abstract.** Agouti (ASIP) and Extension (MC1R) are genes known to be involved in coat colour through pigmentation pathways by regulating type, amount and distribution of eumelanin and pheomelanin pigments in melanocytes. In alpaca genotype of ASIP and MC1R genes have already been analysed distinctly in previous studies, but so far their epistatic interaction have not been evaluated. In this study have been assessed their segregation more insights on black and brown phenotypes. In several mammals MC1R is epistatic over ASIP, id est recessive allele in Agouti (a) and dominant allele in Extension locus (E) produces black phenotype. That is confirmed in alpaca where black coat has aH/aΔ57 and aH/ahT genotype on ASIP and E/E or E/e genotype on MC1R locus. Otherwise ASIP and MC1R in brown, have a dominant profile at least in one allele as A/A, A/ahT on Agouti and E/e on Extension. Genotype and phenotype comparison clears that receptor and ligand are in concordance to produce pheomelanin and eumelanin in alpaca. Segregation analysis of 12 alpaca families genotyped by coat color, confirm the dominance of brown over black and could be helpful for coat colour classification and genotyping.

**Resumen.** Se sabe que los genes Agouti (ASIP) y Extension (MC1R) están implicados en el color de la capa al intervenir en las vías de pigmentación regulando el tipo, la cantidad y la distribución de los pigmentos de eumelanina y feomelanina de los melanocitos. Los genotipos disponibles para estos genes ASIP y MC1R ya han sido analizados en profundidad previamente, pero no se ha estudiado su interacción epistática hasta le fecha. En este estudio se estudia la segregación de estos genes en la aparición de fenotipos negro y marrón. MC1R es epistático sobre ASIP en varios mamíferos, produciendo el fenotipo negro la combinación del alelo recesivo del Agouti (a) con el alelo dominante del gen Extensión (E). Esto se confirma en la alpaca ya que la capa negra tiene un genotipo aH/aΔ57 o aH/ahT en el locus ASIP y E/E o E/e en el locus MC1R. Por el contrario, ASIP y

<footnote n="1">¹KVASU Centre for Wildlife Studies, Kerala Veterinary and Animal Sciences University, Pookode, Wayanad, Kerala 673576, India.</footnote>

<footnote n="2">²School of Bioscience and Veterinary Medicine, University of Camerino, via Gentile III da Varano, 62032 Camerino, (MC), Italy.</footnote>

<footnote n="3">³School of Pharmacy and Health Products, University of Camerino, piazza dei Costanti, 62032 Camerino, Italy.</footnote>
MC1R producirán capa de color marrón cuando el individuo porte al menos un alelo dominante siendo A/A o A/ahT para el locus Agouti y E/e en el locus Extension. La comparación entre genotipos y fenotipos permite concluir que el receptor y el ligando están en concordancia para producir feomelanina y eumelanina en alpacas. El análisis de segregación de 12 familias de alpacas genotipadas para el color de la capa, confirma el predominio del marrón sobre el negro y podría ser útil para la clasificación del color de la capa y el genotipado.

**Keywords:** Alpaca, Genetic interaction, ASIP, MC1R, Black, Brown.

**Introduction**

Fiber harvested from alpaca (Vicugna pacos) is sold as luxury yarn. Peru hosts the largest biological reserve of alpaca in the world. Alpaca are raised in a variety of coat phenotypes and, recent studies have investigated various candidate genes possible involved in coat color variation in alpaca (Feelay et al., 2011). Recently a scheme for coat colour classification was defined by the International Committee for Animal Recording (ICAR) (Antonini, 2009). As regards potential colour, it could be affected by the pigmentation process. The synthesis of both eumelanins and pheomelanins are under control of a group of genes acting at the melanosomes level. The principal genes are Extension (MC1R protein) and Agouti (ASIP protein). Melanocortin 1 receptor (MC1R) and its peptide antagonist agouti-signaling-protein (ASIP) are well known in the regulation of the eumelanin/ pheomelanin switch involved in the base color expression. As in other animals, the epistatic interaction of MC1R and ASIP, could be visible as hues palette of phenotypes, although genotype segregation patterns is not revealed. In alpaca, three significant mutations in the Agouti locus causing black and brown phenotypes were identified. In particular the SNPs (g.3836C>T and g.3881G>A) and an in-frame 57 bp deletion (g.3866_3923del57) in exon-4 are predicted to independently cause functional changes to ASIP protein. Missense mutation g. 3836 C>T involve in an amino acid substitution in R98C, would predict a change of arginine (R) to Cysteine, which seems to have minimal/ partial effect on the functional property. This is evident in one of the allele ahT associate with g.3836C>T mutations seems to have partial/minimal functionality. Missense mutation at g.3881G>A suggests that it may produces a substitution at the amino acid position R118H changes the R to histidine (H) in the cystine-rich domain, which disrupt the highly conserved Arg-Phe-Phe (R-F-F) motif in the protein. Other analysis with cloning experiment showed that one of the allele marked with g.3896G>A (aH) seems to be associated to non-functional ASIP due to the R-F-F motif disruption. Deletion of 57 bp. would result in a short 114 amino acid containing agouti protein, which lacks 19 amino acids from the cysteine (C) rich domain, which is critical in agouti function and ASIP is supposed to be not functional (Table 1). For the locus Extension (MC1R) in alpaca, four missense
mutation are relevant with the black and brown phenotypes. In particular the g.901C>T nucleotide mutation resulting in the R301C amino acid change shows that all the brown animals were heterozygous to C901T (Powell et al., 2008). The interesting fact observed with our molecular segregation analysis is that the animals homozygous to the mutations A82/ A259/ A376/ C901combination expressed black skin phenotypes with an EE genotype. The brown phenotypes were observed to have the heterozygous condition for the observed mutations (A82G/ A259G/ A376G/ C901T) with Ee genotypes (Table 2). In alpaca mutations of this two genes were analysed separately in two previous studies performed by our research group (Bathrachalam et al., 2011; Bathrachalam et al., 2013), but so far their epistatic interaction have not been evaluated. In this regard, the present study aims to fill the knowledge gap on the genetic interaction between ASIP and MC1R in alpaca and, to provide novel information useful for both applicative and basic science issues. Our outcomes provide further insights into the mechanisms of pigmentation in alpaca as well as provide "molecular tools" for the development of an efficient marker assisted breeding program for coat colors in this animal.

<table><caption>**Table 1:** The ASIP geonotypes and phenotype of Peruvian alpacas.</caption><thead><tr><th>Phenotype</th><th>g.3836C>T<br>p.R98C</th><th>g.3896G>A<br>p.R118H</th><th>g.3866-3923del57<br>p.C109_R127del</th><th>Allele</th></tr></thead><tbody><tr><td rowspan="3">Black</td><td>C</td><td>A</td><td>-</td><td><sup>H</sup><br>a</td></tr><tr><td>C</td><td>-</td><td>Yes</td><td><sup>Δ57</sup><br>a</td></tr><tr><td>C</td><td>A</td><td>-</td><td><sup>H</sup><br>a</td></tr><tr><td rowspan="3">Black</td><td>T</td><td>G</td><td>-</td><td><sup>a</sup><sup>T</sup></td></tr><tr><td>C</td><td>G</td><td>-</td><td>A</td></tr><tr><td>T</td><td>G</td><td>-</td><td><sup>a</sup><sup>T</sup></td></tr><tr><td rowspan="2">Brown</td><td>C</td><td>G</td><td>-</td><td>A</td></tr><tr><td>C</td><td>G</td><td>-</td><td>A</td></tr></tbody></table>

## Materials and Methods

### *Sampling and processing of the samples*

Alpacas were sampled from the Quimsachata Experimental Station, Peru. The Station is located on the Andean Plateau at 4,300 m under the management of the Instituto Nacional de Innovacion Agraria (INIA). The trials have been organized in a hierarchical scheme. Five Black rams have been mated to 10 black dams, and for them and their crias (baby alpaca) the colors have been assigned. Brown animals have not been crossed in this mating plan. Skin biopsies were obtained after antisepsis, and local anesthesia was injected at the border of the sampling site. Samples have been collected from parents and crias by disposable biopsy punch
(8 mm diameter) and have been stored in All Protect (Qiagen). Then, the skin fragments have been removed from preservative reagents and stored in liquid nitrogen. For molecular genetic analysis all samples have been transferred to the laboratories at the School of Environmental Sciences, University of Camerino, Italy. The genomic DNA was isolated from the skin biopsies by using DNAeasy tissue kit (Qiagen S.A., Courtaboeuf, France) according to the instruction of the industry. The DNA samples with good quantity and quality have been stored at -80 °C for further analysis. The genotyping assays have been performed for ASIP and MC1R in all the informative phenotypes according to method reported in the previous Bathrachalam works (Bathrachalam et al., 2011; Bathrachalam et al., 2013). Then the genetic interaction between ASIP and MC1R have been analysed (Table 3).

<table><caption>**Table 2:** The MC1R genotypes and phenotype of Peruvian alpacas.</caption><thead><tr><th>g.A82G<br>(p.T28A)</th><th>g.A259G<br>(p.V87M)</th><th>g.A376G<br>(p.G126S)</th><th>g.C901T<br>(p.R301C)</th><th>Phenotype</th><th>MC1R<br>Genotype</th></tr></thead><tbody><tr><td>A/A</td><td>A/A</td><td>A/A</td><td>C/C</td><td>Black</td><td><i>EE</i></td></tr><tr><td>A/G</td><td>A/G</td><td>A/G</td><td>C/T</td><td>Black</td><td><i>Ee</i></td></tr><tr><td>A/G</td><td>A/G</td><td>A/G</td><td>C/C</td><td>Black</td><td><i>Ee</i></td></tr><tr><td>A/G</td><td>A/G</td><td>A/G</td><td>C/T</td><td>Brown</td><td><i>Ee</i></td></tr></tbody></table>

*Ethics statement*

In agreement with the new European Directive on the protection of animals used for scientific purposes (Directive 2010/63/EU, Article 15, Annex VIII), all animal procedures used in the study are classified as ‘mild’ (i.e., procedures with no significant impairment of the well-being or general condition of the animals) and were preemptively approved by the Animal Ethics Committee of the University of Camerino.

<table><caption>**Table 3:** ASIP and MC1R genotype interaction and the following phenotype of Peruvian alpacas.</caption><thead><tr><th>Genotype<br>ASIP (Ligand)</th><th>MC1R (Receptor)</th><th>Phenotype</th></tr></thead><tbody><tr><td>a<sup>Δ57</sup>/a<sup>H</sup>,</td><td>E/E, E/e</td><td>Black</td></tr><tr><td>a<sup>H</sup>/a<sup>hT</sup></td><td>E/E, E/e</td><td>Black</td></tr><tr><td>A/a<sup>hT</sup></td><td>E/e</td><td>Brown</td></tr><tr><td>A/A</td><td>E/e</td><td>Brown</td></tr></tbody></table>
## Results

### *ASIP – MC1R combined Genotype*

In alpaca, the black animals observed to have two agouti genetic backgrounds. One seems to have completely recessive alleles i.e. aH/aΔ57 and another with a recessive and a partial dominant allele i.e. aH/ahT. The MC1R genotypes of black with two agouti genetic background were completely in association, thus black phenotypes with aH/aΔ57 and aH/ahT had either E/E or E/e (MC1R) genotypes (Table 3), thereby the receptor and ligand genetic background is perfectly balanced to produce eumelanin in black animals. Furthermore, black animals were heterozygous to aΔ57 allele with ahT or aH, heterozygous to both ahT and aH, homozygous to the mutation C901 (EE) and heterozygous to C901T (Ee). Brown animals were heterozygous to ahT, homozygous for aH, homozygous to g.3836C, wild allele (A). The aΔ57 and the A were not observed. All the brown animals analysed in the present study were heterozygous to C901T (Ee).

### *Segregation Analysis*

Segregation analysis was performed on 10 alpaca families constituted from 5 male (rams), 10 female (dam) and 10 crias (baby alpaca). All of them presented a black phenotype, and the genotype at the Extension and Agouti locus have been evaluated (Table 4). In all the families the allelic variants at the Agouti locus was aH/ahT or aH/aΔ57 and at the Extension locus was Ee or EE.

## Discussion

Molecular identification of the Agouti and MC1R genes provided much of the molecular groundwork for understanding the role of melanocortin signaling in pigmentation. In alpaca, the extension locus encoding for MC1R is epistatic over locus Agouti which means that a fully functioning MC1R receptor is necessary for the Agouti to be expressed. If MC1R is not functional then it cannot be activated or inactivated by either of two alternate ligands. In most of the mammals the recessive allele in agouti (a) and dominant allele in MC1R locus (E) produces black phenotypes. In black alpacas MC1R is functional at the least in one allele, and ASIP is in recessive form due to possible explained missense mutations. In these cases, ASP lost the ability to block α-melanocyte-stimulating hormone (α-MSH). Improvement of the α-MSH receptor induced cAMP production and up-regulated the transcription of MITF that activates the expression of the melanogenesis-related enzymes and regulating the gene expression of tyrosinase, TRP-1, and TRP-2. These pathways enhance the pigmentary function of melanocytes to produce more eu- and pheo-melanin and hair darkening.
<table><caption>**Table 4:** The Agouti and Extension genotype segregation in 10 black alpaca families.</caption><thead><tr><th>Family</th><th>Code</th><th>Color</th><th>ASIP</th><th>MC1R</th><th>Family</th><th>Code</th><th>Color</th><th>ASIP</th><th>MC1R</th></tr></thead><tbody><tr><td rowspan="3">1</td><td>RAM</td><td>366203<sup>4</sup></td><td>Black</td><td>a<sup>H</sup>/a<sup>bT</sup></td><td>E/e</td><td>RAM</td><td>95101<sup>2</sup></td><td>Black</td><td>a<sup>H</sup>/a<sup>Δ57</sup></td><td>E/E</td></tr><tr><td>DAM</td><td>301204</td><td>Black</td><td>a<sup>H</sup>/a<sup>Δ57</sup></td><td>E/E</td><td>2</td><td>DAM</td><td>386301</td><td>Black</td><td>a<sup>H</sup>/a<sup>Δ57</sup></td><td>E/E</td></tr><tr><td>CRIA</td><td>138108</td><td>Black</td><td>a<sup>H</sup>/a<sup>Δ57</sup></td><td>E/E</td><td>CRIA</td><td>81108</td><td>Black</td><td>a<sup>H</sup>/a<sup>Δ57</sup></td><td>E/E</td></tr><tr><td rowspan="3">3</td><td>RAM</td><td>35104<sup>1</sup></td><td>Black</td><td>a<sup>H</sup>/a<sup>bT</sup></td><td>E/E</td><td>RAM</td><td>95101<sup>2</sup></td><td>Black</td><td>a<sup>H</sup>/a<sup>Δ57</sup></td><td>E/E</td></tr><tr><td>DAM</td><td>83104</td><td>Black</td><td>a<sup>H</sup>/a<sup>bT</sup></td><td>E/e</td><td>4</td><td>DAM</td><td>275204</td><td>Black</td><td>a<sup>H</sup>/a<sup>Δ57</sup></td><td>E/e</td></tr><tr><td>CRIA</td><td>78108</td><td>Brown</td><td>a<sup>H</sup>/a<sup>bT</sup></td><td>E/E</td><td>CRIA</td><td>287208</td><td>Black</td><td>a<sup>H</sup>/a<sup>Δ57</sup></td><td>E/E</td></tr><tr><td rowspan="3">5</td><td>RAM</td><td>35104<sup>1</sup></td><td>Black</td><td>a<sup>H</sup>/a<sup>bT</sup></td><td>E/E</td><td>RAM</td><td>237204<sup>3</sup></td><td>Black</td><td>a<sup>H</sup>/a<sup>bT</sup></td><td>E/E</td></tr><tr><td>DAM</td><td>348204</td><td>Black</td><td>a<sup>H</sup>/a<sup>bT</sup></td><td>E/E</td><td>6</td><td>DAM</td><td>283205</td><td>Black</td><td>a<sup>H</sup>/a<sup>Δ57</sup></td><td>E/e</td></tr><tr><td>CRIA</td><td>264208</td><td>Black</td><td>a<sup>H</sup>/a<sup>bT</sup></td><td>E/E</td><td>CRIA</td><td>160108</td><td>Black</td><td>a<sup>H</sup>/a<sup>Δ57</sup></td><td>E/E</td></tr><tr><td rowspan="3">7</td><td>RAM</td><td>244203<sup>5</sup></td><td>Black</td><td>a<sup>H</sup>/a<sup>Δ57</sup></td><td>E/E</td><td>RAM</td><td>237204<sup>3</sup></td><td>Black</td><td>a<sup>H</sup>/a<sup>bT</sup></td><td>E/E</td></tr><tr><td>DAM</td><td>75103</td><td>Black</td><td>a<sup>H</sup>/a<sup>Δ57</sup></td><td>E/e</td><td>8</td><td>DAM</td><td>480100</td><td>Black</td><td>a<sup>H</sup>/a<sup>Δ57</sup></td><td>E/E</td></tr><tr><td>CRIA</td><td>241208</td><td>Black</td><td>a<sup>H</sup>/a<sup>Δ57</sup></td><td>E/E</td><td>CRIA</td><td>437308</td><td>Black</td><td>a<sup>H</sup>/a<sup>Δ57</sup></td><td>E/E</td></tr><tr><td rowspan="3">9</td><td>RAM</td><td>244203<sup>5</sup></td><td>Black</td><td>a<sup>H</sup>/a<sup>Δ57</sup></td><td>E/E</td><td>RAM</td><td>244203<sup>5</sup></td><td>Black</td><td>a<sup>H</sup>/a<sup>Δ57</sup></td><td>E/E</td></tr><tr><td>DAM</td><td>2160200</td><td>Black</td><td>a<sup>H</sup>/a<sup>Δ57</sup></td><td>E/e</td><td>10</td><td>DAM</td><td>130105</td><td>Black</td><td>a<sup>H</sup>/a<sup>Δ57</sup></td><td>E/e</td></tr><tr><td>CRIA</td><td>262208</td><td>Black</td><td>a<sup>H</sup>/a<sup>Δ57</sup></td><td>E/E</td><td>CRIA</td><td>89108</td><td>Black</td><td>a<sup>H</sup>/a<sup>Δ57</sup></td><td>E/E</td></tr></tbody></table>

In alpaca, the black animals observed to have two agouti backgrounds that seems to have completely recessive alleles i.e. aΔ57/ aH, and aH/ ahT. The MC1R genotypes of black with two agouti backgrounds were completely in association. All the brown animals analysed in the present study were heterozygous at the extension locus E/e and present SNP g.901C>T in one allele. Furthermore, Agouti gene has the A/ahT, A/A genotype. In this case, at the least one allele of both genes have to be functional to express this phenotype that is the typical wild-type pigmentation pattern for the alpaca. This phenotype also occurs in domestic sheep (Våge et al., 2003), in which it is understood to be a product of positive artificial selection for novel colour variants during domestication. In fact when MC1R is bound by ASIP, eumelanin synthesis is inhibited because of the down-

<footnote n="1">¹Ram1. ²Ram2. ³Ram3. ⁴Ram4. ⁵Ram5.</footnote>
regulation of several melanogenic factors, and pheomelanin production relies on the chemical status of the cell environment, i.e. the content of available tyrosine and tyrosinase activity. The depigmentation induced by ASIP results from the inhibited synthesis of both types of melanins concomitant with the dramatic decrease of tyrosinase activity. That leads preferentially to the suppression of eumelanin production during the synthesis of mixed-melanins than pheomelanin. Different shades of brown phenotypes observed in alpaca could be due to the interactive effect between agouti and MC1R followed by a mixed amount and type of melanins production. The dominance of brown phenotype over black coat was confirmed by segregation analysis. In fact occurrence of both black and brown baby alpaca (crias) in crosses of black parents could be explained by the absence of black offspring in crosses involving brown parents (Valbonesi et al., 2011). Furthermore black phenotype is under control of an allelic heterogeneity at ASIP locus. An effort of segregation about black phenotype was confirmed by genotype analysis in Agouti and Extension (MC1R) locus in 10 families. (Table 4)

**Conclusions**

Finally, clearly interaction of agouti and MC1R is in concordance with the black and brown phenotypes not the only genetic interaction through which black and brown pigments are synthetized in alpaca. As verified by the segregation analysis the black colour in alpaca is a recessive pattern. However, this research could have several implications: in fact standardize fibre production based on natural coat colour, could surely help herders farm. Breeders that perform mating program based on colour genotyping of their breeding alpacas could have a large amount of fleece with similar tone and shades. Select coat colour through biomarker assisted selection could increase breeders’ income. That is considerable in Peru, where rear alpaca is popular and poverty prevails. Furthermore natural coloured fibre does not need to be dyed and that improve the performance of eco-friendly, low economic impact and compete with synthetically dyed products, that fade with time and sometimes causes harmfulness to human body. Further studies exploring other candidate genes, especially those with regulatory functions are likely to provide great insights into our understanding of the black and brown phenotypes in alpaca.

**Acknowledgments.** The authors would like to thank the Alpaca Research Foundation (ARF) for their financial support. The authors declare no conflict of interest for this paper.

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