Universitätsverlag Göttingen Advances in Fibre
Production Science in South American Camelids and
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Martina Gerken, Carlo Renieri, Daniel Allain, Hugh Galbraith, Juan Pablo Gutiérrez, Lisa Mckenna, Roman Niznikowski, Maria Wurzinger

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Advances in Fibre Production
Science in South American
Camelids and other Fibre Animals

Edited by
Martina Gerken
Carlo Renieri
Daniel Allain
Hugh Galbraith
Juan Pablo Gutiérrez
Lisa McKenna
Roman Niznikowski
Maria Wurzinger

<figure><img src="image_2.png" /></figure>

Universitätsverlag Göttingen
<figure><img src="image_3.png" /></figure>
Martina Gerken, Carlo Renieri, Daniel Allain, Hugh Galbraith, Juan Pablo Gutiérrez, Lisa McKenna, Roman Niznikowski, Maria Wurzinger (eds.)

Advances in Fibre Production Science
in South American Camelids and other Fibre Animals

This work is licensed under a
Creative Commons Attribution-ShareAlike 4.0 International License.

<figure><img src="image_4.png" /></figure>
Martina Gerken, Carlo Renieri,
Daniel Allain, Hugh Galbraith,
Juan Pablo Gutiérrez, Lisa
McKenna, Roman Niznikowski,
Maria Wurzinger (eds.)

Advances in Fibre
Production Science
in South American Camelids
and other Fibre Animals

<figure><img src="image_5.png" /></figure>

Universitätsverlag Göttingen
2019
Bibliographic information published by the Deutsche Nationalbibliothek

The Deutsche Nationalbibliothek lists this publication in the Deutsche Nationalbibliografie; detailed bibliographic data are available on the Internet at http://dnb.dnb.de.

Editors-in-chief:

Martina Gerken, Carlo Renieri, Daniel Allain, Hugh Galbraith, Juan Pablo Gutiérrez, Lisa McKenna, Roman Niznikowski, Maria Wurzinger

E-Mails: mgerken@gwdg.de; Carlo.Renieri@Unicam.it; d.allain@free.fr; h.galbraith@abdn.ac.uk; gutgar@vet.ucm.es; lisa.mckenna@agr.uni-goettingen.de; roman_niznikowski@sggw.pl; Maria.Wurzinger@Boku.Ac.At

Suggested citation:

M. Gerken, C. Renieri, D. Allain, H. Galbraith, J. P. Gutiérrez, L. McKenna, R. Niznikowski, M. Wurzinger (eds.) (2019): Advances in Fibre Production Science in South American Camelids and other Fibre Animals. Universitätsverlag Göttingen, Göttingen

This work is protected by German Intellectual Property Right Law.

It is also available as an Open Access version through the publisher's homepage and the Göttingen University Catalogue (GUK) at the Göttingen State and University Library (https://www.sub.uni-goettingen.de).

The license terms of the online version apply.

Revision of English language: Lisa McKenna

Revision of Spanish language and abstracts: Juan Pablo Gutiérrez

Typesetting and layout: Lisa McKenna, Marvin Heuduck, Martina Gerken

Fotos (cover, session titles): Alexander Riek and Martina Gerken

Cover design: Margo Bargheer and Jutta Pabst

© 2019 Universitätsverlag Göttingen

https://univerlag.uni-goettingen.de

ISBN: 978-3-86395-408-6

DOI: https://doi.org/10.17875/gup2019-1158
Preface

Animal fibres from South American camelids and other fibre or wool bearing spe-
cies provide important products for use by the human population. The contempo-
rary context includes the competition with petrocarbon-based artificial fibres and
concern about excessive persistence of these in the natural environment. Animal
fibres present highly valuable characteristics for sustainable production and pro-
cessing as they are both natural and renewable. On the other hand, their use is
recognised to depend on availability of appropriate quality and quantity, the pro-
duction of which is underpinned by a range of sciences and processes which sup-
port development to meet market requirements.

Such support includes the efforts of the Animal Fibre Working Group (AFWG) of the European Federation of Animal Sciences (EAAP) which was instituted in 2007 and tasked with creating a network for investigation and dissemination of information in Europe and internationally. One task has been the organisation of scientific meetings, and continuing the tradition of previous European Symposia on South American camelids. These include the recent 5th Meeting in Sevilla (Spain: 2010) and 6th Meeting at EAAP, Nantes (France: 2013). References to these and other meetings, workshops and publications may be found on the AFWG website: http://www.eaap.org/presentation/scientific-structure/commis sions-working-groups/animal-fiber-working-group/.

The present publication derives from the 7th European Symposium on South American Camelids and 3rd European Meeting on Fibre Animals (http://www.sympcam.org/). This meeting was held in the conference facility of the Domus Pacis Hotel, Assisi, Italy, on 12 to 14 June 2017. It was organised by Prof Dr Carlo Renieri and his colleagues Dr Attilio De Cosmo, Dr Francesco Fantuz, Dr Antonietta La Terza, Prof Alessandro Valbonesi (University of Camerino), Dr Marco Antonini (ENEA), and Maurizio Gubbiotti (University Marconi, Roma) with support from the scientific board comprising AFWG colleagues. We wish to thank Dario Pediconi, Cristina Nocelli, Irene Pazzaglia, Stefano Pallotti (University of Camerino) who helped us during the symposium. We also thank all participants who readily agreed to chair sessions or to participate in the Round Table.

We are very grateful to Loro Piana (http://www.sympcam.org/loropiana/) for
generous funding support which enabled the attendance of international speakers
and provided scholarships for three young scientists from Latin American countries.
Individual papers and abstracts, where full papers were not available, were printed from the manuscripts supplied by the authors. The assistance of Marvin Heuduck (Göttingen University) and the editors of Göttingen University Press in the editorial process is acknowledged.

Martina Gerken (Göttingen University), Carlo Renieri (University of Camerino, Italy), Daniel Allain (INRA, France), Hugh Galbraith (University of Aberdeen, UK), Juan Pablo Gutiérrez (Complutense University of Madrid, Spain), Lisa McKenna (Göttingen University), Roman Niznikowski (University of Life Sciences, Warsaw, Poland), and Maria Wurzinger (University of Natural Resources and Life Sciences (BOKU), Vienna, Austria)

The editors
Selection and Evaluation of Fiber Characteristics of an Extreme Fine Alpaca Strain at Victory Farm in Missouri .................................................................................. 121
T. Wuliji

Merino Breeding Program Improves Wool Quality in US Wool Sheep Flocks .... 135
T. Wuliji, L. Wuri, H. Glimp and T. Filbin

Selection Strategies for Fiber Quality in Alashan Cashmere Goat......................... 149
M. Antonini, P.R. Tang, F. Panella, G. Attard, E. Lasagna, S. Ceccobelli
and F.M. Sarti

Interaction between ASIP and MC1R in Black and Brown Alpaca......................... 163
C. Bathrachalam, C. Nocelli, I. Pazzaglia, S. Pallotti, D. Pediconi, A. La Terza
and C. Renieri

Alpaca FGF5: Hypothetical Post-Transcriptional Readthrough Regulation in Skin Biopsies.................................................................................................................. 171
Pallotti S., Pediconi D., Morelli M.B., Dharaneedharan Subramanian, Molina M.G., Antonini M., Renieri C. and La Terza A.

Alpines Steinschaf (Alpine Stonesheep) ................................................................................ 185
Christian Mendel, Isabelle A. Ketterle

## Reproduction and Pathology

The Alpaca Cria, Clinical and Immunological Aspects .................................................. 195
P. Walter Bravo

Addition of Seminal Plasma to Frozen-Thawed Llama Spermatozoa does not Preserve Sperm Motility.................................................................................................................. 201
Fumuso, F.G., Carretero, M.I., Chaves, M.G., Neild, D.M., Miragaya, M.H. and Giuliano, S.M.

Alpaca Semen Quality throughout the Breeding Period.................................................. 213
P. Walter Bravo, W. Garcia and V. Alarcon

The Sperm Chromatin Dispersion Assay (HALO Test) Correlates with the Tunel Technique in Llama Sperm.................................................................................................................. 221
M.I. Carretero, F.G. Fumuso, S.M. Giuliano, D.M. Neild, P. Cetica and M.H. Miragaya

Teeth in Camelids: Myths, Facts and Problems .................................................................................. 229
I. Gunsser

## Nutrition

Advances in Nutrition on Chinese Cashmere Goat: A Review........................................ 239
Sun Haizhoua, Li Shenglia, Zhang Chongzhia, Jin Lua, Sang Dana and Zhang Chunhuaa
Alfalfa Hay Supplementation to Improve Llama Meat Production
for Smallholders in Pasco Region, Peru................................................................255
G. Gutierrez, A. Corredor, R. Robles, J. Mendoza, V. Hidalgo and M. Wurzinger

Water Metabolism in South American Camelids ....................................................267
M. Gerken, L. Brinkmann, R. Amin Runa and A. Riek

Meat and Fibre Production, Fibre Metrology

Carne y charqui de llama ....................................................................................................279
C. Ayala, G. Condori, C. Renieri, S. Pilco and J.L. Quispe

Wool Scouring in Europe: Urgent and Ecological Solutions....................................301
M.T. Chaupin

Proteomic Method for Determination of Animal Hair Fibres ....................................305
C. Tonetti, S. Paolella, D.O. Sanchez Ramirez, R.A. Carletto, C. Vineis, A. Varesano
and S. Sforza

The Use of Near-infrared (NIR) Reflectance Spectroscopy to Predict
Mohair Quality in Greasy Fleece Samples of Angora Goats ....................................313
D. Allain, S. Brenot, G. Auvinet, B. Pena-Arnaud and P. Martin

Variability of Fiber Quality of Chinese Alashan Left Banner White Cashmere
goat ........................................................................................................................................325
S. Pallotti, J. Wang, P. Tang, M. Antonini, Y. Lou, C. Pieramati, A. Valbonesi and
C. Renieri

Effects of Year and Sampling Site on Mean Fibre Diameter of Alashan
Cashmere Goat................................................................................................................333
Marco Antonini, Jun Wang, Yujie Lou, Peirong Tang, Carlo Renieri, Irene Pazzaglia,
Alessandro Valbonesi

Abstracts

Sustainable Cashmere, Pastoralism, and Coexistence with Predators in Europe .. 341
N. Kravis

Efecto de la precipitación pluvial en la seja de selva y la zona alto andina
de la región Puno sobre la producción ganadera de altura ....................................342
Pineda B., Zeballos J., Mamani R. and Huanca T.

Evaluation of Population and Social Composition of Vicunas (Vicugna vicugna)
in Different Environment Sites of the Laguna Blanca Biosphere Reserve
(Catamarca, Argentina) ....................................................................................343
Riva de Neyra, L. A., Hick, M.V.H. and Frank, E. N.

Animal Welfare Problems in South American Camelids Kept in Europe............344
Gauly, M.
<table>
  <tbody>
    <tr>
      <td>Breeding Objectives for Alpacas of the Highlands Central of Peru</td>
      <td>345</td>
    </tr>
    <tr>
      <td><i>Candio, J.R. and Gutiérrez, G.A.</i></td>
      <td></td>
    </tr>
    <tr>
      <td>Vicugna Pacos As1-Casein: Identification of New Polymorphisms at the Csn1s1 Gene</td>
      <td>346</td>
    </tr>
    <tr>
      <td><i>Erhardt, G., Gu, M., Wagner, H., Di Stasio, L. and Pauciullo, A.</i></td>
      <td></td>
    </tr>
    <tr>
      <td>Estimación de la heredabilidad de seis caracteres de calidad de fibra de alpacas huacaya del INIA Puno</td>
      <td>347</td>
    </tr>
    <tr>
      <td><i>Mamani-Cato, R.H., Huanca, T., Pineda, M., Naveros, M. and Gallegos, R.</i></td>
      <td></td>
    </tr>
    <tr>
      <td>Effect of the Brown Coat-Coding Gene (Tyrp-1) on Wool and Skin Color of Żelaźnieńska and Wrzosówka Sheep</td>
      <td>348</td>
    </tr>
    <tr>
      <td><i>Niżnikowski, R., Swiątek, M. and Zymańska, Z.</i></td>
      <td></td>
    </tr>
    <tr>
      <td>Relationship between Classes Assigned by Visual Appraisal and a Selection Index in Function of Live Weight, Fleece Weight and Fiber Diameter in Huacaya Alpacas from Pasco</td>
      <td>349</td>
    </tr>
    <tr>
      <td><i>Corredor F.A. and Gutiérrez G.</i></td>
      <td></td>
    </tr>
    <tr>
      <td>Preliminary Comparative Analysis and Localization of <i>Bos Taurus</i> SNPS on <i>Vicugna Pacos</i> Chromosome 10 (Vpa10)</td>
      <td>350</td>
    </tr>
    <tr>
      <td><i>Farfán K.A., Gutierrez G.A. and Ponce de León F.A.</i></td>
      <td></td>
    </tr>
    <tr>
      <td>Innovative Andrological Evaluation to Optimize the Selection of Fiber Animal</td>
      <td>351</td>
    </tr>
    <tr>
      <td><i>Stelletta, C.</i></td>
      <td></td>
    </tr>
    <tr>
      <td>Use of Seminal Plasma on Interval to Ovulation, Susceptibility of Corpus Luteum to Prostaglandin and Improving of Reproductive Performance in Alpacas (<i>Vicugna Pacos</i>) under Peruvian Highland Conditions</td>
      <td>352</td>
    </tr>
    <tr>
      <td><i>Huanca, W., Turin, J., Huanca, W.F., Mamani, C., Sanchez, S. and Cordero, A.</i></td>
      <td></td>
    </tr>
    <tr>
      <td>Induction of Superovulation in Alpacas According to the Number of Follicles Recruited to the Emergence of Follicular Wave</td>
      <td>354</td>
    </tr>
    <tr>
      <td><i>Pozo A., Vásquez A., Zevallos J., Olivera L., Cordero A. and Huanca W.</i></td>
      <td></td>
    </tr>
    <tr>
      <td>Farmers Wool and Traceability</td>
      <td>355</td>
    </tr>
    <tr>
      <td><i>Thompson, N.</i></td>
      <td></td>
    </tr>
    <tr>
      <td>Feed Intake and Animal Behaviour of Alpaca and Llamas Co-Grazing on Andean Highlands in Peru</td>
      <td>356</td>
    </tr>
    <tr>
      <td><i>Hoehn D., Castro-Montoya J., Gomez C. and Dickhoefer U.</i></td>
      <td></td>
    </tr>
    <tr>
      <td>Daily and Seasonal Changes in Body Temperature and Activity Patterns of Llamas in the High Andes of Peru</td>
      <td>358</td>
    </tr>
    <tr>
      <td><i>Riek, A., Stölzl, A., Marquina Bernedo, R. and Gerken, M.</i></td>
      <td></td>
    </tr>
  </tbody>
</table>
Blood Levels of Phosphorus in Pubescent Alpaca (Vicugna Pacos) and the Effect of Dietary Phosphorus on Growth of Female Alpacas Post Weaning in Peruvian Andes
Quispe, C.E., Ancco, E., Van Saun, R. and Gomez, C.

Digestibility of Bean Pulp Granulated in Rabbits
Arce, O., Alagón, G., Ródenas, L., Martínez-Paredes, E., Moya, V.J., Pascual, J. and Cervera, C.

Correlation between Diameter of Fiber, Medulation and Ancestrality in Alpacas
Melo, C., Zapata, C. and Bravo, W.

Apelin, a New Adipokine Acting on Hair Follicle: an Immunohistochemical Study on Ovine Skin
Mercati, F., Dall'Aglio, C., Guelfi, G., Scocco, P. and Ceccarelli, P.

ICAR – Guideline for the Animal Fibre Production in Alpaca and Cashmere and New Rules for the Organization of the Fibre and Fleece Collection Centers
Antonini, M., Pazzaglia, I., Nocelli, C., Lou, Y. and Thompson, N.

Technological Characteristics of White and Coloroured Huacaya Alpaca Fibre in Apurimac, Perú
Corredor, F.A., Bustinza, V., Machaca, V., Paucara, V., Paúcar, R. and Quispe, E.C.

The Prickling Issue in Fabrics Made of Camelid Fibres: Possible Mechanical or Genetic Solutions
Frank E.N.

Determination of the Optimal Number of Runs Using AM2 Dehairing Technology in Fibers of Patagonian Goats (Patagonian Cashmere)
Frank, E.N., Hick, M.V.H., Castillo, M.F. and Frondizi Seghetti, D.G.

Dehairing of Alpaca Fibres Top with Am2 Dehairing Technology
Frank, E.N., Frondizi Seghetti, D.G., Hick, M.V.H., Castillo, M.F., Burgos, A. and Cruz, A.

Modelación de curvas de crecimiento de llamas q'ara utilizando modelos de crecimiento no lineales
Mamani-Cato, R.H., Huanca, T., Naveros, M. and Gallegos, R.

Genetic Basis of Early Activation of Hair Follicle in Cashmere Goat: An Approach with Candidate Genes
Pazzaglia, I., Mercati, F., Antonini, M., La Terza, A., Nocelli, C., Pallotti, S., Pediconi, D. and Renieri C.
<figure><img src="image_6.png" /></figure>
Sustainable Development, Climate Change
and Biodiversity

<figure><img src="image_7.png" /></figure>
<figure><img src="image_8.png" /></figure>
# Sustainable Development of Livestock Production:
What and how can Research Contribute?

**M. Wurzinger**

Department of Sustainable Agricultural Systems, University of Natural Resources and Life Sciences, Gregor-Mendel-Strasse 33, A-1180 Vienna, Austria

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

**Abstract.** World-wide livestock constitutes a substantial component of agricultural production by contributing to the agricultural GDP of many countries. Livestock contributes to food and nutrition security through the provision of animal-sourced foods. It is a source of income and capital assets, as well as a source of draught power and manure for farming. In pastoral societies, livestock is the prime resource and the largest non-land asset they own. Smallholder farmers contribute a large share to the agricultural production throughout the developing world. Poor households especially depend on and keep livestock. At the same time, we can observe an increasing demand, particular in developing countries, for animal-source food. This so-called livestock revolution is often seen as an opportunity for livestock owners to improve household incomes, secure food for their families and escape poverty. But simultaneously livestock keepers are confronted with many challenges such as climate change, volatile prices and new market demands. There is increasing consensus that change is accelerating and becoming less predictable, as global interconnections lead to events having consequences beyond their immediate context. Research, innovation and (higher) education can play an important role in addressing many of the challenges livestock keepers are facing. But this requires a paradigm shift towards more inter- and transdisciplinary research. Beyond the engagement of the relevant disciplines, it is now increasingly acknowledged that the understanding of and solutions to societal challenges may be better achieved by involving academic and non-academic partners in all stages of the research process. Collaboration of different stakeholders can lead to new thinking and might bring new solutions – not only technical, but also institutional ones. In recent years, innovation platforms have gained some attention as it gives different actors along a certain value chain the opportunity to interact on a regular basis. These platforms provide the necessary room to discuss current and relevant topics. For researchers it will remain difficult to balance their efforts between basic and more applied research – both equally important for the sustainable development of the livestock sector. By supporting resource-poor and often marginalized livestock keepers, research can contribute to achieving the SDG-Sustainable Development Goals endorsed by the United Nations.

**Resumen.** La ganadería a nivel mundial constituye un componente importante de la producción agrícola mediante su contribución al PBI del sector agrícola en muchos países. El sector ganadero contribuye a la alimentación y seguridad alimentaria a través de la pro-visión de alimentos de origen animal. También, la ganadería es fuente de ingresos y de
bienes de capital, así como aporta con fuerza de tracción y guano para la agricultura. En sociedades pastoriles, la ganadería es el recurso principal y el bien de capital, sin considerar a la tierra, más grande. Los agricultores pequeños contribuyen en gran escala a la producción agrícola en los países en desarrollo. Las familias pobres especialmente crían y dependen del ganado. Al mismo tiempo, nosotros podemos observar un incremento de la demanda, particularmente en países en desarrollo, de alimentos de origen animal. La llama da “Revolución ganadera” es vista como una oportunidad para los propietarios de ganado para mejorar sus ingresos familiares, asegurar la alimentación de sus familias, y escapar de la pobreza. Pero simultáneamente los criadores de ganado se enfrentan a muchos retos como el cambio climático, precios volátiles y nuevas demandas del mercado. Existe un creciente consenso que el cambio se está acelerando y es menos predictable, así como las interconexiones internacionales nos están llevando a eventos con consecuencias más allá del contexto inmediato. La investigación, la innovación y la educación (superior) jugaran un rol importante en encontrar soluciones a varios de los cambios que los criadores de ganado están enfrentando. Pero, se requerirá un cambio de paradigma hacia una mayor investigación inter y trans-disciplinaria. Más allá del involucramiento de las disciplinas relevantes, cada vez se está reconociendo que el entendimiento y el planteamiento de soluciones a los retos sociales podrán ser mejor logrados con el involucramiento del sector académico y no académico en todas las fases del proceso de la investigación. En años recientes, las plataformas de innovación han recibido mucha atención debido a que los diferentes actores de una cierta cadena de valor tienen la oportunidad de interactuar de manera regular. Estas plataformas proveen el espacio necesario para discutir temas actuales y relevantes. Para los investigadores persistirá la dificultad de mantener un balance entre sus esfuerzos de realizar investigación básica versus la investigación aplicada- ambos son importantes para el desarrollo sostenible del sector ganadero. Mediante el apoyo a los ganaderos de recursos pobres y marginalizados, la investigación puede contribuir al logro de los Objetivos de Desarrollo Sostenible (ODS) endosado por las Naciones Unidas.

**Keywords:** sustainable production, multi-actor, livestock

**Introduction**

In 2015, the Sustainable Development Goals (SDG) were adopted by the United Nations member states. This agenda calls for action by all countries to improve the livelihoods of all people globally. By addressing challenges in the livestock sector, contribution to SDG1 (no poverty), SDG2 (no hunger), SDG3 (good health and wellbeing), SDG12 (responsible consumption and production) and SDG17 (partnerships for the goals) can be achieved as the livestock sector plays an important role in many economies of low to middle income countries. The livelihoods of millions of rural households depend partly or totally on income from livestock products (Banerjee et al., 2015, IFPRI, 2016). In Peru the area of alpaca production overlaps with high incidences of rural poverty. The figures from 2017 indicate that 34% of all alpaca farmers are considered as poor and 12% as extreme poor (MINAGRI, 2017).
Income growth in low- and middle-income countries increases consumption of products of animal origin, contributing to food security, wellbeing and human health (FAO, 2017). This so-called Livestock Revolution is often referred to as a possible pathway out of poverty for livestock keepers. Global growth of meat and milk consumption is projected to be 1.6 and 1.3 percent per year, respectively, in the 2007–2030 period (FAO, 2015). In India, per capita demand for milk is expected to increase by 57 percent between 2007 and 2030 (FAO, 2011) and milk consumption is likely to triple by 2050 in sub-Saharan Africa, mostly led by East Africa (Herrero et al, 2014). Although animal fiber has a relatively low share on the global fiber market, it is very well positioned in the high-quality fiber sector. Since 2001 in Peru the alpaca fiber production increased annually by 1.79 %, which is mainly a result of an increase in number of animals. In 2015 the Peruvian fiber production reached 4,478 t, of which 90 % was for export market and 10 % for the national market (mainly for handicraft). Main destinations are Italy, China, South Korea and Taiwan. In 2016 the export volume of tops was 51 million US $ (Minagri, 2017).

At the same time, livestock production is facing many challenges such as com-
petition of resources like water and land and the negative impact of climate change
and disease outbreaks. Climate change increases stress for livestock (HLPE, 2016)
and could cause a 10 to 25 percent decrease in milk production (IPCC, 2014),
which is an important source of nutrition for children (Marquis et al., 1997). An
estimated 1.96 billion people rely on livestock to supply part, or their entire daily
needs (Anderson, 2003). Currently, societies are confronted with multiple trends
that exert pressure on socio-ecological systems such as population dynamics and
urbanization (UN 2011; Długosz 2011), globalization (Guillén 2001), climate
change and natural resource depletion or degradation (IPCC 2017; Prior et al.
2012). These changes make the need for livestock professionals, able to respond to
changing needs and demands of the sector stakeholders, apparent. The necessary
transformation to a sustainable society will require guidance by responsible actors
who have a deep and contextualized understanding of current trends and challeng-
es in socio-ecological systems (Webster 2007). Socio-ecological systems have so-
cial, economic and ecological dimensions that characterize a multi-level complex
adaptive system (Smit and Smithers 1994; Tittonell 2014). Complementing their
knowledge, future decision makers in such systems will need particular capabilities
and skills to facilitate societal innovation (WBGU 2012; Brown et al., 2000). A key
skill, according to Fortuin and Bush (2010), will be the ability to cross boundaries
between disciplinary knowledge, cultures and theory and practice. Indeed, the dis-
cussion on how we can facilitate sustainability learning in higher education has
gained momentum in the past several years (Tàbara and Pahl-Wostl 2007;
Shephard 2008; UNESCO 2005; Wright and Horst 2013). As universities educate
future decision makers, they have a key responsibility for the transformation to-
wards sustainable socio-ecological systems (Shephard 2008; Gadotti 2008; Ciurana
and Filho 2006; Moore 2005; Stephens et al. 2008; Mochizuki and Fadeeva 2010).
In turn, Gadotti (2008) argues that education today is part of the causes of unsustainable lifestyles. Krizek et al. (2012) and Beringer (2007) underline that the institution university with all its components, including curricula, has to undergo a sustainability transition.

This paper presents two different case studies in which scientists aim to engage in a dialogue with different stakeholders in the sector of South American camelid production in Peru and sheep production in Argentina. Based on these two case studies some general conclusions will be drawn.

## Description of case studies

The case studies presented in this paper differ in their research scope, number and type of stakeholders involved and scale of impact.

### Case study 1 – Development of a breeding program for llamas in Peru

From 2011 – 2013 the 1st project phase “Strengthening llama production in the Central Andes of Peru” and from 2014-2017 the 2nd project phase “Design and implementation of community-based breeding strategies for llamas in the Peruvian Andes” were jointly implemented by Universidad Nacional Agraria La Molina in Lima and BOKU-University of Natural Resources and Life Sciences, Vienna. This project was designed using a transdisciplinary research approach. Participatory, action research methodology was used to ensure the interaction and communication between the research team and the livestock keepers. At the beginning of the project the main goal was to characterize the llama production systems and llama populations in three locations of the Peruvian Central Andes and to develop and to use this information for the joint implementation with the livestock keepers a breeding program for a sustainable use of the local llama populations (Wolfinger, 2012; Radolf, 2014). In the second phase different intervention strategies in the herd management accompanying the breeding program were tested together with livestock keepers. Further details of this research are reported by Apaza et al. 2015, Gómez et al. 2015, Pizarro et al. 2015, Gutierrez et al. 2017 and Wurzinger & Gutierrez 2017.

Llama farmers were in the spotlight of the research. In individual meetings and workshops they could express their needs and ideas. Based on this information, training sessions with livestock keepers were held. Mutual learning for both groups, scientists and livestock keepers, was an essential component of the project. Masters students got the opportunity to learn this new research approach, which puts the farmers in the center of all activities. Other stakeholders like representatives of the Ministry of Agriculture and persons from an NGO were also invited to participate as the project evolved. The idea was that both, the ministry and the NGO could take over responsibilities in the long run and support the local farming community. Keeping the constant contact with livestock keepers in very re-
mote areas posed a challenge for the research team. Time consuming and expen-
sive travels over a long period are difficult to maintain. A field assistant, who fre-
quently travelled could ensure a constant interaction with all people involved.
Bridging the scientific excellence (e.g., publications in well-recognized journals)
with local, tailor-made solutions for smallholder farmers was a challenge for the
researchers.

*Case study 2 – Transformation of higher education in the area of animal science in Peru and Argentina*

The EU-funded project “EDULIVE-Transforming higher education to strengthen links between universities and the livestock sector in Argentina and Peru” addresses the integration of universities within society at large using the concept of knowledge triangle. According to this concept, “the contribution of higher education to jobs and growth, and its international attractiveness, can be enhanced through close, effective links between education, research, and innovation (...)” (EU, 2017). Applying this to the livestock sector, it seems vital to strengthen the cooperation between universities for animal science and the relevant livestock sector stakeholders, promoting the offer of demand-driven higher education and with it the ability to react to current needs and expectations of the sector. Further details under: http://edulive-international.eu/

The aim of the project is to make the academic offer more demand-oriented and addresses topics of relevance for the different stakeholders. In Peru two different value chains, alpaca fibre and dairy value chain, and in Argentina the sheep (wool and meat) value chain were taken as an example. In each country, two public universities, the national research organisations, farmers´ organisations, NGOs and private companies participated in the project. A series of workshops with universities and all stakeholders were carried out to get a better understanding of the different needs, but also to identify mechanisms and possible areas of interest for future cooperation. Internships for students for a period between 2 weeks and 3 months (depending on the requisites of the universities) were implemented. The aim of these internships was to offer students the opportunity to test their knowledge in a real world setting, but also to provide the stakeholders with insights of the capacities of possible future employees and give universities an immediate feedback loop on their curriculum.

All universities revised their policies for internships and made adjustments in their curricula. The idea is that not only the students, but also the university and the receiving institution can benefit from the internship of students. Therefore, an accompanying, well-documented process is launched in the universities. Nevertheless, it remains a challenge for universities to address the large variation of demands coming from the livestock sector and reflect these in their training programs.
## Conclusions

Addressing the demands from the livestock sector in research and higher education is an important and relevant topic. Research has the responsibility, especially as it often receives public funding, to identify possible solutions for the different stakeholders. But it remains a challenge for many researchers to engage in a dialogue with farmers as it requires specific skills, open-mindedness, resources and time. Therefore, it is imperative that universities constantly get feedback from the representatives of the sector to improve and up-date their training programs on a regular basis. This can ensure that graduates are equipped with the necessary skills and knowledge to further develop their working environments and contribute to a sustainable development of the livestock production.

## References

* Anderson, S. 2003, Animal genetic resources and sustainable livelihoods. *Ecological Economics*, 45(3):331-339.

* Apaza, C.; Gutierrez, G.; Salva, B.; Elias, C. 2015, Propiedades fisicoquímicas de hot dog de carne de llama con inclusión de albúmina, goma de tara y proteína de soya. *VII Congreso Mundial de Camélidos, Puno, Peru*.

* Banerjee, A.; Duflo, E.; Goldberg, N.; Karlan, D.; Osei, R.; Pariente, W.; Shapiro, J., Thusbaert, B.; Udry, C. 2015, A multifaceted program causes lasting progress for the very poor: Evidence from six countries. *Science* 348 (6236).

* Beringer, A. 2007, "The Lüneburg Sustainable University Project in International Comparison: An Assessment against North American Peers." *International Journal of Sustainability in Higher Education* 8 (4): 446-61.

* Brown, A.L.; Bransford, J.D.; Cocking, R.R. 2000, *How People Learn: Brain, Mind, Experience, and School*. Washington DC: National Research Council.

* Ciurana, A.M.; Geli de, L.L.; Filho, W. 2006, "Education for Sustainability in University Studies: Experiences from a Project Involving European and Latin American Universities." *International Journal of Sustainability in Higher Education* 7 (1): 81-93.

* Dlugosz, Z. 2011, "Population Ageing in Europe." *Procedia – Social and Behavioral Sciences* 19: 47-55.

* EU, 2017, ec.europa.eu. In: http://ec.europa.eu/education/policy/higher-education/knowledge-innovation-triangle_en, as of 26.09.2017

* FAO. 2011, Mapping supply and demand for animal-source foods to 2030, by T.P. Robinson & F. Pozzi. *Animal Production and Health Working Paper.No. 2*. Rome.

* FAO. 2015, The Second Report on the State of the World's Animal Genetic Resources for Food and Agriculture, edited by B.D. Scherf & D. Pilling. *FAO Commission on Genetic Resources for Food and Agriculture Assessments*. Rome.

* FAO. 2017, The future of food and agriculture – Trends and challenges. Rome.

* Fortuin, I.; Karen, P.J.; Bush, S.R. 2010, "Educating Students to Cross Boundaries between Disciplines and Cultures and between Theory and Practice." *International Journal of Sustainability in Higher Education* 11 (1): 19-35.

* Gadotti, M. 2008, "What We Need to Learn to Save the Planet." *Journal of Education for Sustainable Development* 2 (1): 21-30.
Gomez, A.; Gutierrez, G.; Wurzinger, M.; Infantes, M.; Elias, C.; Salva, B. 2015, Características químicas y propiedades tecnológicas de la carne de llama (lama glama) procedente de Marcapomachoca, Junín, Perú. VII Congreso Mundial de Camélidos, Puno, Perú.

Guillén, M.F. 2001, "Is Globalization Civilizing, Destructive or Feeble? A Critique of Five Key Debates in the Social Science Literature." Annual Review of Sociology 27: 235–60.

Gutierrez, G.A.; Mendoza, J.G.; Hidalgo, V.; Wurzinger, M. 2017, Alfalfa hay supplementation to improve llama meat production for smallholders in Pasco Region, Puno. Proceedings of the 7th European Symposium on South American Camelids and 3rd European Meeting on Fibre Animals, June 12-14, 2017, Assisi, Italy.

Herrero, M.; Havlik, P.; McIntire, J.; Palazzo, A.; Valin, H. 2014, African livestock futures: realizing the potential of livestock for food security, Poverty reduction and the environment in sub-Saharan Africa. Geneva, Switzerland, Office of the Special Representative of the UN Secretary General for Food Security and Nutrition and the United Nations System Influenza Coordination (UNSIC) (available at http://preview.tinyurl.com/ocuqphk).

HLPE, 2016. Sustainable agricultural development for food security and nutrition: what roles for livestock? A report by the High Level Panel of Experts on Food Security and Nutrition of the Committee on World Food Security. Rome. http://www.fao.org/3/a-i5795e.pdf

IFPRI-International Food Policy Research Institute. 2016, 2016 Global Food Policy Report. Washington, DC: International Food Policy Research Institute.

IPCC (Intergovernmental Panel on Climate Change) 2014, Change 2014: Impacts, Adaptation, and Vulnerability. Part A: Global and Sectoral Aspects. Contribution of Working Group II to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change. C.B. Field, V.R. Barros, D.J. Dokken, K.J. Mach, M.D. Mastrandrea, T.E. Bilir, M. Chatterjee, K.L. Ebi, Y.O. Estrada, R.C. Genova, B. Girma, E.S. Kissel, A.N. Levy, S. MacCracken, P.R. Mastrandrea & L.L. White, eds. World Meteorological Organization, Geneva, Switzerland, 190 p.

IPCC. 2017, Climate Change 2017: Impacts, Adaptation, and Vulnerability.

Krizek, K.J.; Newport, D.; White, J.; Townsend, A.R. 2012, "Higher Education's Sustainability Imperative: How to Practically Respond?" International Journal of Sustainability in Higher Education 13 (1): 19–33.

Marquis, G.S.; Habitcht, J.P.; Lanata, C.F.; Black, R.E.; Rasmussen, K.M. 1997, Breats milk or animal-product foods improve linear growth of Peruvian toddlers consuming marginal diets. American Journal of Clinical Nutrition 66(5): 1102–1109.

Minagri (Ministerio de Agricultura y Riego) 2017, Diagnóstico de Crianzas Priorizadas para el Plan Ganadero 2017-2012, Lima. Peru.

Mochizuki, Y.; Fadeeva, Z. 2010, "Competences for Sustainable Development and Sustainability: Significance and Challenges for ESD." International Journal of Sustainability in Higher Education 11 (4): 391–403.

Moore, J. 2005, "Seven Recommendations for Creating Sustainability Education at the University Level: A Guide for Change Agents." International Journal of Sustainability in Higher Education 6 (4): 326–39.
Pizarro, D.M.; Gutierrez, G.A.; Ñaupari, J.A.; Wurzinger, M. 2015, Modelo de simulación dinámica para la evaluación de la sostenibilidad de la crianza de llamas en la región Pasco, Perú. VII Congreso Mundial de Camélidos, Puno, Perú.

Prior, T. D.; Giurco, G.; Mudd, L. Mason; Behrisch, J. 2012, "Resource Depletion, Peak Minerals and the Implications for Sustainable Resource Management." Global Environmental Change 22 (3): 577–87.

Radolf, M. 2014, Livelihood and production strategies of smallholder livestock keepers in the Central Peruvian Andes. Master thesis. University of Natural Resources and Life Sciences, Vienna, Austria.

Shephard, K. 2008, "Higher Education for Sustainability: Seeking Affective Learning Outcomes." International Journal of Sustainability in Higher Education 9 (1): 87–98.

Smit, B.; Smithers, J. 1994, "Sustainable Agriculture and Socio-ecological system Health." In Agroecosystem Health. Proceedings of an International Workshop., edited by N.O. Nielson, 31–38. Guelph: University of Guelph.

Stephens, J.C.; Hernandez, M.E.; Roman, M.; Graham, A.C.; Scholz, R.W. 2008, "Higher Education as a Change Agent for Sustainability in Different Cultures and Contexts." International Journal of Sustainability in Higher Education 9 (3): 317–38.

Tábara, J.D.; Pahl-Wostl, C. 2007, "Sustainability Learning in Natural Resource Use and Management." Ecology and Society 12 (2).

Tittonell, P. 2014, "Livelihood Strategies, Resilience and Transformability in African Agroecosystems." Agricultural Systems 126: 3–14.

UN 2011, World Populations Prospects: The 2010 Revision.

UNESCO 2005, UN Decade of Education for Sustainable Development 2005-2014: The DESD at a Glance. Paris.

WBGU 2012, Research and Education: Drivers of Transformation. Berlin.

Webster, K. 2007. "Hidden Sources: Understanding Natural Systems Is the Key to an Evolving and Aspirational ESD." Journal of Education for Sustainable Development 1 (1): 37–43.

Wolfinger, B. 2012, Characterisation of the production system of llamas and description of breeding strategies of smallholders in the Central Peruvian Andes. Master thesis, University of Natural Resources and Life Sciences, Vienna, Austria.

Wright, T.; Horst, N. 2013, "Exploring the Ambiguity: What Faculty Leaders Really Think of Sustainability in Higher Education." International Journal of Sustainability in Higher Education 14 (2): 209–27.

Wurzinger, M.; Gutierrez, G. 2017, Analysis of a multi-stakeholder process during the start-up phase of two community-based llama breeding programs in Peru. Livestock Research for Rural Development 29 (10).
# Animal Fibre Production in Europe: Biology, Species, Breeds and Contemporary Utilisation

H. Galbraith

School of Biological Science, University of Aberdeen, 23 ST Machar Drive
Aberdeen, AB 23 8EQ, United Kingdom

E-Mail: h.galbraith@abdn.ac.uk

**Abstract.** Animal fibre “wool” of the pelage is a keratin protein-based product formed in specialised hair follicles located in the skin of a range of animal species and breeds farmed in Europe. It is described as a natural, renewable and biodegradable product of long-standing historical value. Its use in recent decades has been affected by issues such as political changes, poor economic returns and from competition with petro-carbon-based artificial fibres. Such artificial fibres from clothes are typically poorly degraded and are now reported to wash into, and pollute, the world’s oceans and the food chain. Ecologically-sensitive methods for wool production and processing and its inherent degradability have recognised importance. Primary hair follicles typically, although with some exceptions, produce outer fibres of greater diameter, than the underlying and more valuable finer fibres from secondary follicles. Aspects of biology important for production and properties of fibres are recognised. The coarse and fine fibre products require separation after harvesting. This report concerns the animal species, goats (cashmere: Angora) and South American camelids (alpaca: Llama), which are present in limited numbers and supply relatively small, and frequently niche, markets in Europe. Angora wool from rabbits supplies an established but reducing market. These species of animals generally produce fibres of fine quality. The multi-million populations of different breeds and cross breeds of sheep, with meat and milk as primary products, are also reviewed. These produce wool of a wide range of qualities much of which is exported overseas particularly to China. The fibres from all species have properties, such as colour, medullation, tensile strength, diameter (fineness) and staple length. These properties determine end-use from small-diameter superfine yarns to the coarser fibres utilised in carpets and furniture upholstery. Poor quality wool may be used in geotextiles, or disposed of in landfill, or incinerated.

Numbers of these animals, systems of husbandry, economics, social contribution, sustainability and arrangements for collection and marketing of fibre are described in different countries in Europe. Sources include statistical reports and case studies provided by members of the European Federation of Animal Sciences (EAAP), Animal Fibres Working Group (AFWG). Reference is made to international and European initiatives for the promotion of wool such as The Campaign for Wool and The Wool Group. Note that all websites were accessed on 16 April 2018.
**Resumen.** La estructura de la fibra de lana de origen animal presenta una base proteínica de queratina que se forma en folículos capilares especializados localizados en la piel de varias especies y razas de animales criadas en Europa. Se describe como un producto natural, renovable y biodegradable de valor histórico muy antiguo. Su uso en las últimas décadas se ha visto afectado por cuestiones como cambios políticos, bajos rendimientos económicos y la competencia con fibras sintéticas derivadas del carbón y del petróleo. Este tipo de fibras presentes en la ropa presenta una muy pobre degradación natural, y recientemente se ha descrito que acaban contaminando los océanos del mundo afectando a la cadena alimenticia. Por ello se ha reconocido la importancia de los métodos ecológicamente sensibles para la producción y el procesamiento de lana. Aunque con algunas excepciones, los folículos capilares primarios producen típicamente fibras externas de mayor diámetro que las fibras más finas subyacentes más valiosas de los folículos secundarios. Existen aspectos biológicos de reconocida importancia en la producción y las propiedades de las fibras. Se precisa la separación de las fibras finas de las gruesas a partir del vellón esquilado. Este trabajo se centra en especies de animales domésticos, como cabras (cashmere y angora) y camélidos sudamericanos (alpacas y llamas), con números pequeños en Europa y que abastecen pequeños mercados, que normalmente constituyen nichos de mercado. La lana de conejos de angora abastece mercados establecidos pero en reducción. Estas especies de animales generalmente producen fibras de buena calidad. También se revisa la producción de lana de multitud de millones de ovejas de diferentes razas y cruces destinadas a la producción de carne y leche. Estas ovejas producen lana de una amplia gama de calidades, muchas de las cuales se exportan al extranjero, especialmente a China. Las fibras de todas las especies tienen propiedades, como el color, la medulación, la resistencia a la tracción, el diámetro (finura) y la longitud de la fibra. Estas propiedades determinan el uso final desde las prendas superfinas de pequeño diámetro a las fibras más gruesas utilizadas en alfombras y tapicería de muebles. La lana de baja calidad se puede usar en geotextiles o desecharse en vertederos o incinerarse. Se presentan las poblaciones de animales, los sistemas de cría, la economía, la contribución social, la sostenibilidad y la organización para la recolección y comercialización de fibra en distintos países de Europa. La información se obtuvo de informes estadísticos y estudios de casos proporcionados por miembros de la Federación Europea de Ciencias Animales (EAAP) y el Grupo de Trabajo de Animales de Fibra (AFWG). También se hace referencia a la participación del grupo en iniciativas internacionales y europeas para la promoción de la lana, como La Campaña por la Lana (The Campaign for Wool) y El Grupo de Lana (The Wool Group). Téngase en cuenta que el último acceso a las Web con información fue el 16 de abril de 2018.

**Keywords:** animal fibre production, fibre markets

**Introduction**

Natural fibres such as wool from animals have a long history of high economic value and effective utilisation but have been affected in recent decades by competition from artificial synthetics and those based on plant-derived materials (Galbraith, 2010a). Wool fibres contain polymers of keratins and other proteins
located in the cytoskeleton of cells. Plant fibres contain cellulose-based polymers while artificial fibres derived from petrocarbons are polymers of simpler organic molecules and include polyester, nylon, and polypropylene.

However, one feature of the latter is their limited degradability and which, along with certain plastics and “microfibres” shed from clothes, have been linked with pollution in land and water (Brown et al., 2011). One consequence of this may be a greater focus on better utilisation of natural fibres which, while retaining properties of durability in use and potential for recycling, have ultimately greater degradability and lesser ecological persistence.

It is also important to be aware of the impact of wool production and processing on the environment and to note developments in eco-labelling and application of life cycle assessment (IWTO, 2018). In this context, the availability of scouring resources, such as provided by the Nejdek Wool Combing (2018) facility in the Czech Republic is recognised as important in Europe along with a focus on maximising ecological outcomes (Chaupin, 2018). Ketterle (2018) has commented on the objective of deregulating the EU-Regulation 1069/2009 which classifies greasy wool as a Category 3 animal by-product.

IWTO (2018) has summarised the wool industry supply chain including production, processing, product manufacturing, recycling and biodegradation. Considering collectively all fibres, these have many uses which depend on their physical and compositional properties. For animal fibres, their properties derive from biology of production in follicles in the skin of animals and are dependent on species and breeds and mechanisms of production within the animal. Such biological mechanisms in animals has been the subject of extensive study for considerable time (eg Ryder, 1958). Interest has been maintained in Europe by the EAAP AFWG and the sharing of developments in knowledge and practices in production such as described in the current Symposium (Sympcam, 2017).

## Basic biology and husbandry

In considering basic biology (Galbraith, 2010b, 2010c), the fibre of major interest is the “wool” covering the body of the animal and referred to as the pelage or fleece coat. Wool fibre is produced in hair follicles embedded in the skin, uniquely of mammals. There are two main types of follicles, primary and secondary and a range of subtypes, the hair fibre products of which vary in property according to physical characteristics.

Primary hair follicles have accessory structures of apocrine sweat glands, sebaceous gland and arrector pilae muscle and produce fibres which may be coarse and function as protective guard hair in overlying fibres of the secondary follicle. The latter is typically of finer diameter, has a role in thermoregulation by providing
insulation (Gerken, 2010), and on harvesting, has greater economic value. Secondary follicles differ in having only a sebaceous gland. The products of glands contribute to environmental protection of the fleece, but require removal in processing of harvested fibre. Lanolin from sebaceous glands of sheep has important commercial value in treatment of human skin conditions.

The fibre products of the (coarser) primary and (finer) secondary follicles in double-coated animals require to be separated as the end-uses, generally determined by diameter, differ.

## Fibre production in follicles

Hair fibre is produced by division of keratinocyte (epidermal) cells which line the base of the follicle in contact with underlying dermal tissue (Galbraith, 2010b, 2010c). These cells divide and migrate towards, and beyond, the skin surface. The rate of proliferation of these cells also determines the rate of growth of fibre (cortex and cuticle), and fleece, and determines staple length. As the cells migrate they deposit a range of proteins in the internal skeleton and other proteins and lipids which are important in adhesion between cells and which contribute to properties of softness, flexibility, moisture absorption and tensile strength (Lyons, 2009). In addition, consistency of husbandry and nutrition of animals has implications for uniformity of chemical and physical composition along the length of fibres (Galbraith, 2000).

The number of epidermal cells across each fibre contributes to its diameter. Small diameters are typically associated with secondary fibres, finer products and greater economic value. Certain fibres have medullation which is shown by a poorly formed or hollow central shaft which contributes to insulative properties. The pattern of growth of fibres is typically cyclical in nature with an anagen growth phase followed by cessation of growth in catagen and a telogen resting phase before appearance of renewed anagen and regeneration of the follicle. The timing of the following shedding or moulting of the hair is important in harvesting of fibre, particularly if a combing, rather than shearing, method is used.

The hair growth cycle may be influenced by photoperiod, such as seasonally-produced cashmere production in goats with short anagen (months), or where largely uninfluenced by external light stimuli, and with endogenous genetic control determining long anagen (in years) and typical of wool production in Merino sheep (Allain and Renieri, 2010).

Seasonal producers of wool, if unshorn, may also shed fleeces usually in summer, whereas other species and breeds which do not, or only partially, shed and require shearing to remove the fleece. Such removal of fleece provides a harvest of wool as well as improving the physical status of the animal in hot environments.
Where wool has little or no commercial value, and does not shed, for example in sheep, shearing is still required on grounds of animal welfare. There is, in addition, interest in genetic selection for natural shedding traits, such as in the Romane sheep breed (Allain et al. 2011) in France and Exlana sheep in the UK (https://www.exlana.co.uk/exlana) to avoid the need for physical removal of wool with related financial and environmental costs. Breeding and selection indices for the most valuable product of follicles, include fineness (smallness of diameter), also frequently associated with juvenile animals, high numbers of follicles and long anagen fibre growth phase Allain and Renieri (2010). Other properties of commercial interest include colour, derived from pigmentation-containing vesicles in cells in the hair fibres, light reflection giving sheen and lustre, medullation and crimp (bending of fibres) frequently associated with fineness and spinning capacity. A smooth covering of cuticle cells on the outer surface of fibres is recognised to contribute to softness. IWTO (2018) has summarised cellular structure in wool fibre in relation to properties of value in end-use. The quality of the fleece is reduced by contaminating material and matting of the wool.

## Summary of processing and typical end-uses for animal fibres

Following harvesting by combing or shearing, fibres are cleaned and processed to form the major “worsted” or “woollen” yarns. (British Wool Marketing Board (BWMB), 2018). The worsted system involves fibres of longer staple length which are combed to lay end-to-end in parallel with much of the natural crimp of wool removed and which forms a smooth, variably tight, yarn when spun. Following weaving, these typically make lighter weight and finer fabrics used in suitings, outerwear, other garments where a smooth finish is preferred and uphosterly and carpeting.

The woollen system processes wool fibres of shorter staple length. These are carded to lie together for processing into yarn and used, for example, in knitting or weaving into fabrics. Woollen fabrics are typically softer and fluffier than worsted and are used for jackets, coats, rugs and blankets where a bulkier and more textured finish is required. The end use of animal fibres is determined by their physical properties and the preferences of human consumers either in clothing (typically smallest diameters and highest monetary value for knitwear and suiting), in domestic “home” environment (typically coarser and hard wearing for upholstery, carpets, insulation) and similarly externally (materials in transport vehicles (carpets, uphostery); horticulture (plant beddings; fertiliser) or if inadequate quality, unused and deposited in land fill or incinerated.

## Species and breeds of importance

The species of major interest in production of fibre (“wool”) in Europe are goats, South American Camelids, rabbits and sheep.
*Cashmere-bearing goats*

These animals (*Capra hircus*) have seasonal production and are double-coated with both primary and secondary follicles. The latter follicles produce the fibres of small diameter (12-18 µm) and with annual yields per European goat, typically of 50-150 g. This is in contrast with yields of cashmere up to 1kg in Chinese breed types. Harvesting may be by combing, or shearing. The raw fibre is processed which includes separating from course guard hairs and contaminating material and cleaning to produce “pure” cashmere. World production is of the order of 6,500 tonnes predominantly from China and Mongolia (Common Fund for Commodities, 2009).

With respect to Europe, there was considerable interest in the 1980's and 1990's (Ibraheem et al. 1992; Galbraith, 2006) and one analysis (Russel, 1998), concluded that “cashmere production would constitute a viable alternative to traditional forms of livestock farming throughout the EU”. However, the 2017 perspective suggests limited success in Europe due to inadequacies in numbers of animals and yields and quantities of cashmere. Utilisation largely derives from smaller scale production and manufacture such as that described for Chianti Cashmere (Kravis, 2018) in Italy: http://www.chianticashmere.com/en/) and BowmontTM (2018) in the UK.

As regards end uses, cashmere is a fibre combining light weight and softness with insulative properties. It can be dyed and spun into yarns and utilised in high value knitware such as sweaters, hats, gloves and socks, or woven into fabrics and used to produce garments such as outer coats, jackets, trousers and scarves. Larger scale fabric and garment producers in, for example, Scotland and Italy utilise cashmere fibre imported from outwith Europe (eg. https://www.loropiana.com/en/).

*Angora goats*

The other major fibre of goats is mohair produced by the Angora genotype (*Capra hircus angorensis*). These animals have predominantly secondary follicles generating fibres in the range in diameter of 22-35 µm and 2-5 kg annual yield. Contaminating primary follicle products require to be separated to ensure homogeneity of fibres. Fibres from young animals have smallest diameters and are softest in handle.

World production total up to approximately 5,000 t/year (Common Fund for Commodities, 2009). Similarly to cashmere, attempts to expand production in Europe in the 1980's and 1990's have met with limited success. Examples of contemporary production in European countries include the UK where 2.8 tonnes of unprocessed mohair were recently auctioned in South Africa by British Mohair Marketing Ltd (2016). Highest monetary values of kid and adult fleeces respectively, reflecting differences in diameter, and after deduction of post-farm costs, were (£ / kg) 10.31 and 9.08. Data for France indicate a population of 5,000 goats in 140 flocks and producing typically 5.00 kg/animal giving an annual production of 25 tonnes (http://en.france-genetique-elevage.org/Conservation-
breeds, 405.html). A selection scheme comprising 2,500 goats is described on 35 farms belonging to Capgènes-section Angora (http://www.capgenes.com/les-races-caprines/race-angora-francaise/).

Typical properties of mohair fibres include durability, flame- and crease-resistance and significant moisture- and dye-absorbing capability, the latter contributing to dense colors in processed products. Its smooth surface, arising from relatively flat cuticle scales gives a lustrous appearance. Mohair is used to manufacture many products. These include knitting yarn for hand or machine knitwear, socks and sweaters and fabric for lightweight suitings, scarves, blankets and upholstery velours. Fine mohair from younger animals is used in clothing, while thicker and coarser fibres from older animals are used in rugs and carpets and heavy fabrics for jackets and coats.

## South American Camelids (SAC)

The main interest is alpaca (*Lama pacos*) with lesser focus on llama (*Lama glama*) and with little or no production of guanaco (*Lama guanicoe*) and vicuña (*Vicugna vicugna*) fibre in Europe.

Alpaca are predominantly single-coated and produce “alpaca” fibre from secondary follicles with typical diameters of 18-30 µm and yields of raw fibre of 1.5-5.5 kg (Antonini, 2010). The main producing country is Peru with an estimated population of 3,685 m yielding 6,500 tons. The fibre of cria, (young animals) has the smallest diameter and is considered lighter, warmer and softer than cashmere (Common Fund for Commodities, 2009). Alpaca fibre is partially medullated which contributes to insulative properties and is produced in a range of natural colours. The alpaca SAC are typically sheared once per year. Two different phenotypes are described in alpaca, identified as Suri and Huacaya which differ for the type of the fleece (Presciuttini et al., 2010). The biological basis for the differences between Huacaya and Suri has been the subject of recent genetic research (Presciuttini et al., 2010). There are about 45,000 registered alpaca in the UK where individual pregnant female alpaca are described as having a price tag of £3,000 to £15,000 (Inca Alpaca, 2018).

Alpaca fibre is used primarily for knitware in addition to woven products for clothing, including those for outdoor sports, and accessories such as shawls, rugs and duvets. Blends of alpaca with wool, cotton and silk are also made and used in both woven cloth and knitware. (Common Fund for Commodities, 2009). In the UK, with an estimated processing quantity of 35-40 tons of raw fibre in 2016, many farmers process their own alpaca fibre and sell directly to the public, yarns and products ranging from insoles for boots and wellingtons with coarser fibre, to babywear and exclusive luxury fashion items with the finest fibres (British Alpaca Fleece Buyer, 2018).

CBI (2016) provides a useful overview on many aspects of utilisation of alpaca products in Europe and of assistance to exporters from developing countries.
For llama, typical diameters of the undercoat are 20-30 µm with annual yields of 1.5-2.0 kg in a range of colours. Two main breed types, “Chaku” and “Q’aras” are described by Antonini, (2010). The British Llama Society (2018) describes the limited production of llama fibre and identifies three subspecies: tapada, lanuda and ccara.

Uses of llama fibre include fine yarns made from the separated undercoat which are very soft and characteristically lighter than wool of sheep. Typical knitware products include cardigans, jumpers, scarves, gloves and socks. Coarser fibres from the outer coat are used to make rugs and tapestries. Llama fibre may be combined with other fibres to make blends with variable properties.

In terms of populations for South Americans camelids in Europe, data appear variable. For example, Bonavia (2008) provides, what appear as underestimate, at least for alpaca, of presence in certain countries which give totals of 7,000 llamas, 2,000 alpacas, and 1,000 vicuñas. Estimates were for France, 2-3,000 alpacas and llamas; England, 2-3,000 alpacas and llamas (contrasting with the values given earlier of 45,000); Germany, 2,500 alpacas and llamas and Italy 150 alpacas. Numbers of alpacas and llamas in Austria of up to 6,000 were recently described (Trah and Wittek, 2013). Following shearing, fleeces and fibre of these animals are mostly processed at home or given to small companies for processing (M. Wurzinger, personal communication).

The Llama & Alpaca Registries Europe (LAREU: http://www.lareu.org/) (Kiesling, 2018) provides an online registration service, with database comprising in excess of 19,000 SAC, for alpaca and llama breeders and owners in Europe.

## Rabbit angora

The third species of interest comprises angora-producing rabbits (*Oryctolagus cuniculus*). Angora wool comprises very fine medullated fibres with diameters in the range 14-16 µm and flat cuticle scales (Rougeot and Thebault, 1984). Current world production approximates to 3,000 tons with 90% in China which farms 50 million rabbits (Common Fund for Commodities, 2009). European production approximates to 10 tons, with 3.5 tons in France in 2015, produced by 2,500 rabbits and which has seen major decreases in production since the 1960s. Other European countries which produce angora wool include Hungary and Czech Republic.

There are two main breed types which are both double-coated with an outercoat of coarse fibres and a fine down undercoat. The fleece is harvested three or four times each year. The French type characteristically produces an annual yield of 1.5 kg. While expressing natural moulting, the fleece is typically harvested following feeding of a forage (leucaena) (Allain and Renieri, 2010) containing the chemical anti-nutrient mimosine. This produces a break in the connection of the fibre to the follicle and results in shedding of the fleece. This fleece may be
collected by plucking. The fleece from these animals produces yarn with a
“fluffing” effect.

The German type has a fleece of coarse hair and fine down, producing an annual yield of 1.5-2.0 kg. The fleece does not moult and removal is by shearing. The yarn produced is well suited for warm and thermal clothes particularly for people suffering from arthritis or allergies to sheep wool. The price (2015) of Chinese Angora wool imports was 30 €/kg and 60 €/kg for French-produced. The French and EU Angora industry is described as a niche market vertically controlled by farmer/breeder organisations with direct marketing of final products by farmers.

Angora wool is soft to the touch, has good absorption of water and absorbs dyes effectively. Its uses are in luxury products in knitted clothes, such as pullovers, scarves, socks and gloves It is frequently blended with up to 20 % of sheep wool. One negative feature of the Angora rabbit husbandry, particularly large scale, is the perception of poor welfare conditions. This has resulted in retail outlets declining to sell angora wool garments with associated reductions in production of angora fibre in Europe.

## Sheep wool

The animal fibre produced in largest quantities internationally and in Europe is the wool of sheep (*Ovis aries*). The world sheep population is given as 1,163,045 million (British Wool Marketing Board, 2018). The major producing country (with numbers of sheep (x103) and clean raw wool, including skin wool from abattoirs, (tonnes)) is China (162,062: 276,803) with the largest production in Europe in the United Kingdom (33,743: 20,158). Numbers for Europe (FAO, 2016) are 131,059,072. Different sources vary in the values reported. A selection of published data from Eurostat and FAO is shown in Table 1. The greatest numbers of sheep are seen in the order of the United Kingdom, Spain, Romania, Greece, France and Italy. The data for wool production appear only approximately consistent in respect of sheep numbers and similarly for monetary values of raw wool. The European Parliament Briefing (2017) indicated that raw wool production was valued at € 159 million in 2015 in the EU and referred to the value in the United Kingdom as more than € 40 million which is not consistent with the value recorded in Table 1 from the same source. Reference is made to the value of raw wool, as a percentage, of national value of animal products in the EU as averaging at 0.2 % with 2.2 % and 1.7 % for Romania and Slovenia respectively.

## Uses of wool and contemporary statistics

As for other fibres, the use of sheep wool is determined by a range of properties such as fineness and includes suits and jackets, shirts, knitwear, ties, plaids and blankets, Japanese futons, rugs and carpets. IWTO (2018) summarizes uses in
categories of apparel, smart textiles, sports, manufacturing, medical, architecture, protective apparel, aviation and protection in industry.

Wool is generally recognized a secondary product of meat and milk production by sheep. EU-Regulation 1069/2009 classifies it as a category 3 animal by-product. Current statistics describe dairy and non-dairy ewe populations in European geographical locations (Eurostat 2014). The “Northern Group” countries (Germany, Ireland, Hungary, Netherlands, Sweden and the United Kingdom) are described as having 35,000 dairy ewes (0.2 % of total) against a population of non-dairy animals of 15.374 million. In contrast “Southern Group” countries (Bulgaria, Spain, Greece, France, Croatia, Italy, Portugal and Romania) are recorded as having 24.525m dairy (59.7 % of total) and 16.559 million non-dairy animals.

<table><caption>**Table 1:** Estimates of sheep populations, greasy wool production and monetary value of raw wool for a range of European countries according to statistical sources.</caption><thead><tr><th rowspan="2">Countries</th><th>Sheep</th><th>Greasy Wool</th><th>Raw wool value</th></tr><tr><th>(million heads)<br>2016*</th><th>(tonnes x10<sup>3</sup>).<br>2011**</th><th>(million Euro)<br>2016***</th></tr></thead><tbody><tr><td>France</td><td>7.2</td><td>14</td><td>3.4</td></tr><tr><td>Germany</td><td>1.6</td><td>3****</td><td>2.75</td></tr><tr><td>Greece</td><td>8.8</td><td>8</td><td>2.08</td></tr><tr><td>Hungary</td><td>1.2</td><td>4</td><td>4.22</td></tr><tr><td>Ireland</td><td>3.4</td><td>14</td><td>5.01</td></tr><tr><td>Italy</td><td>7.3</td><td>9</td><td>9.41</td></tr><tr><td>Netherlands</td><td>1.0</td><td>3</td><td>4.68</td></tr><tr><td>Portugal</td><td>2.1</td><td>6</td><td>8.17</td></tr><tr><td>Romania</td><td>9.9</td><td>18</td><td>44.65</td></tr><tr><td>Spain</td><td>16.0</td><td>22</td><td>16.46</td></tr><tr><td>UK</td><td>23.8</td><td>67</td><td>30.76</td></tr><tr><td>Total</td><td>82.3</td><td>168</td><td>131.59</td></tr></tbody></table>

Another important factor is the reduction in sheep populations in recent decades, for example, 11.4 million animals between 2005 and 2014 in the 14 countries with more than 0.5 million sheep (Eurostat 2015). The major decreases are in sheep meat-producing countries such as Spain, (31 %), Portugal (30 %), Ireland (22 %) Germany (21 %), France (18 %). Reductions in the United Kingdom, with 28 % of EU-28 production, are reported as 3 %.

Niznikowski and Strzelec (2012) describe reductions in populations of sheep, from 55 to 35 million head, and annual production of wool from 60,000 to 30,000 tonnes in countries in Central and Eastern European between 1990 and 2010.

<footnote>* Eurostat, 2017a; ** FAO, 2014; *** Eurostat, 2017b; **** corrected.</footnote>
These followed the change of the political system in 1990 and removal of subsidies for wool. The largest reduction was observed in Poland (94 %) with lesser reductions in Hungary (41 %) and Romania (42 %). Sheep populations in these countries in 2010 are given as 0.261 m, 1.223 m and 9.141 m respectively (FAO, 2018). Contemporary utilisation includes a focus on agro-tourism and production of artistic handicrafts and different types of thermal insulation.

## Case studies in Europe

In addition to evaluation of published statistics, members of AFWG provided a summary of contemporary production and utilisation of animal fibre in certain European countries.

### *United Kingdom*

For example, in the UK which has the largest population of sheep and production of wool, the British Wool Marketing Board (2018) has a central role in collecting, grading, promoting and selling British Wool. More than 70 breeds of sheep are recognised by the Board. The breeds are divided into groups: Fine, Medium, Cross, Lustre, Hill, Mountain and Naturally Coloured. Clean raw wool production in the UK is estimated at 21,600 tons from a sheep population of 33 m.

Historically, as in other countries, wool has had major economic importance, such as exemplified by the wealth of the family of William Shakespeare in the 17th century. As indicated above, meat is recognised as the primary product, with wool sometimes a nuisance “by-product”. There are many specialist wool producers and breed organisations in the UK, which include the Shetland Sheep Society (2018) and BowmontTM (2018) (Superfine Merino and Cashmere). Additional interest is maintained in fine wool merino which serves a niche market for worsted processing and fine suiting. Reference is made to the weaving and finishing of high quality wool in Yorkshire and supplying Hirsch Tailoring (2018).

Sheep wool production maintains importance in the UK, and Scotland as the base of the present author. There is considerable interest in cross breeds of sheep with a variety of genetics contributing to meat production, reproductive proliфication, milk production and resilience under intensive husbandry conditions. Moreno Martinez et al. (2012), describe the considerable variation in mean values for diameter of individual wool fibres from individual cross breed animals, among other traditional pure breeds, with raw wool yield of 2-4 kg. There was variation in the presence of crimp and lustre in fleeces of sheep, indicating a heterogeneous product in collected wool in the same flock.

These higher-yielding genotypes contrast with the breeds such as the lower-producing Backface reared under more demanding extensive environmental conditions and contributing to the rural economy such as in the west coast of Scotland. As in other countries, economic returns from wool are small and do not always
cover the cost of wool removal. Interestingly, strong, coarse Blackface wool finds a market in the Italian mattress trade (http://www.scottish-blackface.co.uk/blackface-sheep-information.cfm). An example of economic returns (data from 16 August 2016 and average clip value guide), after all costs, and excluding tax, was for breeds as follows (£/kg), Fine, 0.93: Medium, 1.00: Cross, 0.92: Lustre, 4.13 (Blue Faced Leicester): Hill, 0.78-1.18: Mountain, 0.4-0.6). (http://www.britishwool.org.uk/page/wool-marketing/price-indicator.php).

### *Italy*

Selected information for sheep includes reference to fourteen breeds which are recognized in the national stud book (Carlo Renieri, personal communication). Of these, three are the wool breeds, Gentile di Puglia (6,800: 394 rams and 6,406 ewes), Sopravissana (10,001: 592 rams and 9,409 ewes) and Merinizzata Italiana (New breed 44,841: 2,073 rams and 42,138 ewes) (Assonapa, 2014).

The population of sheep is described as approximately 8 million which are mainly milk breeds. Total wool production is approximately 9,000 tons raw wool of which 250 tons are derived from the Merino breed type. Approximately 90 % is exported in the international market (similar to Germany) with approximately 10 % processed in Italy.

Official arrangements for collection of wool include the established centre at Biella The Wool Company (2018) and more recent developments in the Abruzzo Region for collection of wool in the Gran Sasso and Monti della Laga National Park. Additional centres are in the process of formalization in Puglia Region (Alta Murgia National Park), Sicily Region and Umbria Region.

Thirty three breeds of sheep are described by Biella The Wool Company which in addition to providing a service in processing of wool is a source of expertise and information in the wool supply chain (Thomson, 2018). Gubbiotti et al (2012) have described the structural characteristics of fleece from a range of Italian meat and dairy breeds of sheep. As in other European countries, regions of Italy have had a long tradition of financial wealth based on the wool trade. One important example is the Wool Guild (Arte della lana) of Florence and relationship with the Medici banking family up to the 16th century. Interestingly, Munro (2012) refers to the importance of imported English wool in the success of the Guild. Currently, Italy has been recognised as second in the world, for manufacture and third for export, of textiles internationally (Greta and Lewandowski, 2010).

### *Germany*

Details for Germany include information provided by Martina Gerken (personal communication). The sheep population is estimated at 1.579 m (Statistisches Bundesamt, 2017). The population is made up of mainly meat-producing breeds and is more limited for milk at 12,300 head. 51 breeds of sheep are registered.
These are categorized as follows: (numbers in parenthesis: Schafzuchtverbände Niedersachsen (2016), as merino (3), meat (9), landrace (22), mountain (9) and hair (5). It is notable that in common with certain Eastern European countries, (Niznikowski and Srezlec 2012), production from fine-wool Merino maintained in East Germany up to reunification in 1991, is no longer economically viable. Subsidies are available according to regional location, for certain endangered and mainly landrace breeds producing course wool fibre. There is interest in preserving rare indigenous sheep breeds (Ketterle, 2018).

Production of greasy/raw wool is currently estimated at 3,000 tons. This is valued at 0.5 to 1.0 €/kg, and at 50 % yield, provides an estimated 1,500 tons of pure wool. The revenue for wool is barely adequate to cover the cost of shearing. Approximately 10 % of the greasy wool product (300-400 tons/year) is processed in Germany, with the remaining 90 % exported to Asian countries, particularly China. The end-use of such wool, typically with diameters more than 28 µm, includes carpets, socks and “Regia Wool™ for hand knitting. Additional uses are insulation material (due to low flammability) for roofs and walls of houses, filling and padding material such as in automobiles and airplanes, and geotextiles including pads made from wool infiltrated with plant seeds for greening of land endangered by soil erosion. Wool is also frequently combined with synthetic fibres in the textile industry.

## France

Information for France includes data and statistics compiled by Daniel Allain (personal communication) and sourced from Dispositif INOSYS Réseaux d'élevage – Institut de l'Elevage/Chambres d'Agriculture, France. Totals for populations of sheep (million) and annual wool production (tonnes) for different breed types are reported as follows, for dairy (1.47: 1,537), meat (1.253: 3,901), “hardy” (2.150: 3,642) and conservation (0.089: 248). Overall population and annual wool production calculates to 4.962 m and 9,330 tons respectively. Similarly, data for mean values for different fineness of wool and approximate population (million) are reported as follows: fine merinos (21 µm: 0.78), medium merinos (25 µm; 0.351), fine medium (28.6 µm, 4.117), medium crossed (32.1 µm; 1.571), coarse (37.0 µm; 0.028), kempy (35.6 µm; 2.307) and pigmented (29.2 µm; 0.173).

France benefits from a national genetic improvement program involving 4.1 million ewes of high genetic potential (France Génétique Elevage, 2018a). For selection of meat breeds, 10 specialised breeds and a range of hardy meat breeds adapted to adverse environments are included. Among the characteristics tested is genotyping for resistance to scrapie. On-farm performance testing of rams is conducted on 1100 farms with rigorous further selection on test stations. This genetic approach has been successful for both pure breed and cross breed selection
schemes. A similar approach is taken to selection programmes for milk production
(France Génétique Elevage, 2018b).

Additional study on genetic diversity has been investigated, for example, by Leroy et al. (2015) in 51 populations of 49 breeds of sheep reared in France, where 57 breeds are officially recognised. The analysis utilised 21 microsatellite markers and concluded that the Merino genetic influences included predominantly until the end of the 19th Century to improve wool production, was weaker than that related to the UK breeds which replaced them with a view to improving meat production. In current practice, costs of shearing are given as 1.5-2.0 €/sheep which, similar to other countries, may not be covered by the value of fleece produced. As indicated above, there is interest in genotypes which shed wool and do not require active shearing (Allain et al., 2011).

Spain

Selected data for Spain include those provided by J P Gutiérrez (personal communication). Spain is well recognised to have the second largest population of sheep in Europe, composed of 80% meat breeds and the remainder dairy. In common with other countries, populations of sheep, particularly meat breeds have been reducing in recent decades. For example, the breeding population of 11.3 m ewes (Eurostat, 2017a) has been reduced from approximately 18m in 2002. These breeding animals contribute to a total population of 18.55m sheared which produced 22,000 tons of wool. This was composed (tons), for different qualities as follows: fine (4,179) extra fine (12,784), course (4,792) and black wool (174) respectively. Given the poorer economic returns for wool production, there has been recent strategic focus on using genetic selection to improve meat production from Spanish Merino sheep (Valera et al., 2009). These workers estimated heritabilities and phenotypic and genetic correlations for wool characteristics (greasy fleece weight; fibre diameter: staple length; crimp frequency) following decades of selection. They produced simulations of different selection strategies and concluded that wool quality could be improved at the same time as growth rate and meat production.

There are 43 officially recognized ovine breeds in Spain, which have developed from long term artificial selection for production traits of wool, meat and milk (Manunza et al., 2016). These workers have reported on genotype analysis on 11 of these breeds and showed differentiations in populations in certain breeds while others were shown to have a similar genetic background. The work served to highlight the complications of explaining practical outputs of artificial selection in terms of identifiable “signatures” at the level of the genome.
## Austria

Selected information was provided by Maria Wurzinger (personal communication). Information relating to Austria indicates a population of 14,955 sheep farmers and 349,087 sheep for which there are 30 recognised pure-breeds (BMLFUW 2015). The economic value of these animals is recognised as limited.

The wool crop in 2014 was slightly in excess of 1,000 tons produced mainly in Styria (284 tons), Upper Austria (212 tons) and Tyrol (187.5 tons). The wool crop is either sold to trading companies or processed by farmers and sold directly to end-consumers.

Sheep breeder associations organize collection of wool, for example, for selling to the industry, mainly as insulating material for house constructions (Styrian), processing to produce organic fertilizers for crop production (Upper Austrian), or for use in out-door textiles (Tyrolean).

## Promotion of wool

Reference is made in the above text to the long history of utilisation of wool in the textile trade, a tradition continued in the current international Campaign for Wool (2018). The patron of the Campaign is Prince Charles, The Prince of Wales. The Campaign seeks to promote the use of wool as a natural, renewable and biodegradable resource and supports the aims of environmental responsibility and sustainability in the development of a commercially viable wool industry. The Campaign is active in Europe and supports events held in different countries including those recently in France, Germany, Italy, Netherlands and the United Kingdom. Good welfare in wool sheep production is recognised in the IWTO Guidelines for ethical wool sheep production (IWTO, 2018). The European Wool Group (2018) is also recognised as acting to improve the value of European-produced wool.

## Conclusions

Animal fibre continues to be produced in Europe in variable quantities by animal species such as cashmere and Angora (mohair) goats and South American camelids, particularly alpaca. Markets for these tend to be small scale, local and niche. Production of rabbit angora wool continues although has been in decline in recent years following adverse consumer response. Fibres from these animals tend to be finer and of higher value than much of the product from sheep which are present in considerably greater numbers throughout the continent. The main product from the many breeds of sheep is either meat or milk with fibre generally a by-product. There is some production from fine wool sheep in certain countries which does contribute to uses such as higher value apparel and knitware. Sheep wool of lesser quality has value such as in carpets and uphostery and insulation products and geotextiles. Competition with fibres from plant sources and
petrochemicals is recognised, along with problems with the environmental persistence of the latter.

Some reference is made to the science underpinning fibre production by animals ranging from hair follicle biology, thermoregulation and quantitative and molecular genetics. This includes studies on the natural shedding of wool and avoiding shearing which is often uneconomic. National breeding programmes are documented along with initiatives designed to promote the utilisation of wool in Europe and internationally.

**Acknowledgments.** The author acknowledges the contribution of colleagues on the EAAP Animal Fibre Working Group and the information provided in the case studies for individual countries. These are Carlo Renieri, Daniel Allain, Martina Gerken, Maria Wurtzinger, Juan Pablo Gutiérrez and Roman Niznikowski with additional assistance from Marco Antonini. Support from the University of Camerino for attendance at Sympcam 2017 is acknowledged along with longstanding friendship from colleagues there in the subject of animal fibres and science. Margaret Merchant is also thanked for information on fine wool and its utilisation in the UK.

## References

- Allain, D.; Renieri, C. 2010, Genetics of fibre production and fleece characteristics in small ruminants, Angora rabbit and South American camelids. Animal (2010), 4, 1472–1481.

- Allain, D.; Pena, B.; Foulquieoulquié, D.; Bourdillon, Y.; François D. 2011, Genetic variability of fleece shedding in the Martinik hair, Romane sheep breeds and their crossbreds. In: Pérez-Cabal M.Á., Gutiérrez J.P., Cervantes I., Alcalde M.J. (eds) Fibre production in South American camelids and other fibre animals. Wageningen Academic Publishers, Wageningen. pp15-24.

- Antonini, M. 2010, Hair follicle characteristics and fibre production in South American camelids. Animal 4(9):1460-71.

- Assonapa, 2014, Associazione Nazionale della Pastorizia. www.assonapa.it

- Biella the Wool Company. 2018, http://www.biellathewoolcompany.com/

- Bonavia, D. 2008, The current distribution of South American Camelids. Chapter 12 in The South American Camelids. https://trove.nla.gov.au/version/46533832

- Bowmon, T.M. 2018, Superfine merino and cashmere. http://www.bowmontuk.com/

- British Alpaca Fleece Buyer. 2018, https://www.british-alpaca.com/

- British Llama Society 2018, http://www.britishllamasociety.org/Activities/Fibre/Fibre.html

- British Mohair Marketing Ltd 2016 https://www.angoragoats-mohair.org.uk/2016/05/2015-mohair-fibre-collection-sale-results/

- Brown, M.A., Crump, P., Niven, S.J, Teuten, E., Tonkin, A., Galloway, T., Thompson, R. 2011, Accumulation of Microplastic on Shorelines Woldwide: Sources and Sinks

- Environtal Science Technology.45 (21), 9175-9179.

- BMLFUW (Austrian Federal Ministry of Agriculture, Forestry, Environment and Water Management), 2015, Green report 2015.

- British Wool Marketing Board 2018 http://www.britishwool.org.uk

- Campaign for Wool 2018 http://www.campaignforwool.org/
CBI. 2016. CBI. Ministry of Foreign Affairs. https://www.cbi.eu/market-information/apparel/alpaca/europe/

Chaupin, M. T. 2018, Wool scouring in Europe: urgent and ecological solutions. Sympcam 2017.

http://www.sympcam.org/wpcontent/uploads/2016/10/Book_of_Abstacts.pdf.
P 22.

Common Fund for Commodities 2009, International Year of Natural Fibres.

http://www.naturalfibres2009.org

European Parliament Briefing 2017, The sheep and goat sector in the EU.

http://www.europarl.europa.eu/RegData/etudes/BRIE/2017/608663/EPRS_BRI(2017)608663_EN.pdf

The European Wool Group 2018 http://atelierlainesdeurope.eu/eurowoolgr.htm

Eurostat, 2017a. Sheep population – annual data.

http://appsso.eurostat.ec.europa.eu/nui/show.do?dataset=apro_mt_lssheep&lang=en

Eurostat 2017b. Economic accounts for agriculture – values at current prices

http://appsso.eurostat.ec.europa.eu/nui/submitViewTableAction.do

Eurostat, 2014. http://ec.europa.eu/eurostat/statistics-explained/index.php?title=File:National_ewe_flocks_and_dairy_share,_by_location_o_f_main_breeding_countries,_2014.png

Eurostat, 2015 Statistics explained. http://ec.europa.eu/eurostat/statistics-explained/index.php?title=Meat_production_statistics

Exlana Sheep 2018. Exlana sheep in the UK (https://www.exlana.co.uk/exlana)

FAO 2014. Table 14. Production of eggs and wool. http://www.fao.org/3/a-i3621e.pdf

FAO 2016. Live animals.Sheep. http://www.fao.org/faostat/en/#data/QA

FAO, 2018. Live animals.Sheep. 2010. http://www.fao.org/faostat/en/#data/QA.

France Génétique Elevage, 2018a http://en.france-genetique-elevage.org/Meat-sheep-breeds-selection.html

France Génétique Elevage, 2018b http://en.france-genetique-elevage.org/Dairy-sheep-breeds-selection.html).

Galbraith, H. 2000, Protein and sulphur amino acid nutrition of hair fibre-producing Angora and Cashmere goats. Livestock Production Science, 64, 1, 81–93.

Galbraith, H. 2006, A current perspective on the biology of fibre production in animals. in South American Camelids Research Volume 1, pp. 195-206.

Galbraith, H. 2010a, Animal fibre: Connecting science and production. Animal,4, 1447-1540. Galbraith, H 2010b. In vitro methodology, hormonal and nutritional effects and fibre production in isolated ovine and caprine anagen hair follicles. Animal 4, 1482-1489.

Galbraith, H 2010c, Fundamental hair follicle biology and fine fibre production in animals. Animal 4, 1490-1509.

Gerken, M 2010, Relationships between integumental characteristics and thermoregulation in South American camelids. Animal 4, 1451-1459.

Greta, M.; Lewandowski, K. 2010, The textile and apparel industry in Italy: current state and challenges to further growth. Fibres & Textiles in Eastern Europe.18, No. 6 (83) pp. 20-25.

Gubbiotti, M.; Pazzaglia, I.; Lebboroni, G.; Antonini, M.; Renieri, C. 2012, Fleece structure of Italian dairy and meat sheep. Book of Abstracts of the 63rd Annual Meeting of the European Federation of Animal Science. Wageningen Academic Publishers. P 374.
Hirsch Tailoring 2018 http://www.hirschtailoring.com/

Ibraheem, M.; Galbraith, H.; Scaife, J.R.; Ewen, S. 1994, Growth of secondary hair follicles of the Cashmere goat in vitro and their response to prolactin and melatonin. Journal of Anatomy. 185, 135-142.

Lyons, B. 2009, Australian Merino wool. In Proceedings of the Symposium on Natural Fibres. Common Fund for Commodities, Technical Paper no. 56. FAO, Rome, pp. 83-109.

Inca Alpaca. 2018. http://www.incaalpaca.co.uk/alpaca-info/about-alpacas/faq/

IWTO 2018. http://www.iwto.org/work

Ketterle, N. 2017, Collection of diversity- preserving rare indigenous sheep breeds in Germany. Sympcam 2017. http://www.sympcam.org/wp-content/uploads/2016/10/Book_of_Absttracts.pdf. P 14.

Kiesling, N. 2018, The camelid registry LAREU: what are we breeding in Europe. Sympcam 2017. http://www.sympcam.org/wpcontent/uploads/2016/10/Book_of_Absttracts.pdf. P.31.

Kravis, N. 2018, Sustainable cashmere, pastoralism, and coexistence with predators in Europe. Sympcam 2017. http://www.sympcam.org/wp-content/uploads/2016/10/Book_of_Absttracts.pdf. P 15

Leroy, G.; Danchin-Burge, C.; Palhière, I.; SanCristobal, M.; Nédélec, Y.; Verrier, E.; Rognon, X. 2015, How do introgression events shape the partitioning of diversity among breeds: a case study in sheep. Genetics Selection Evolution. 47, 48

Manunza, A.; Cardoso, T. F.; Noce, A.; Martínez, A.; Pons, A.; Bermejo, L.A.; Landi, V.; Sànchez, A.; Jordana, J.; Delgado, J. V.; Adán, S.; Capote, J.; Vidal, O.; Ugarte, E.; Arranz, J.J.; Calvo, J.H.; Casellas, J.; Amills, M. 2016, Population structure of eleven Spanish ovine breeds and detection of selective sweeps with BayeScan and hapFLK. Scientific Reports volume 6, 1-10. doi:10.1038/srep27296

Moreno Martinez, L.; Mcdonald, A.J.S.; Galbraith, H. 2012, Variation in wool fibre characteristics in sheep differing in genotype. Book of Abstracts of the 63rd Annual Meeting of the European Federation of Animal Science. Wageningen Academic Publishers. P 373.

Munro, J H. 2012, The Rise, Expansion, and Decline of the Italian Wool-Based Cloth Industries, 1100-1730: A Study in International Competition, Transaction Costs and Comparative Advantage. Studies in Medieval and Renaissance History, 3rd Series, Vol. 9 (2012). AMS Press, Inc.

Nejdek Wool Combing, 2018. http://www.gmodiano.com/#our-processor

Niznikowski, R.; Strzelec, E. 2012, Sheep and wool production in Central and Eastern Europe. Book of Abstracts of the 63rd Annual Meeting of the European Federation of Animal Science. Wageningen Academic Publishers. P 372.

Presciuttini, S.; Valbonesi, A.; Apaza, A.; Antonini, M.; Huanca, T.; Renieri, C. 2010, Fleece variation in alpaca (Vicugna pacos): a two-locus model for the Suri/Huacaya phenotype. BMC Genetics. 11, 70. doi: 10.1186/1471-2156-11-70

Rougeot, J.; Thébault, R.G. 1984, Le lapin Angora, sa toison, son élevage. Le Point Vétérinaire, Maisons, Alfort, France.

Russel, A. 1998, The establishment of cashmere production in the European Union. http://www.macaulay.ac.uk/livestocksystems/feasibility/cashmere-feasibility.pdf
Ryder, M.L. 1958, Investigations into the distribution of thiol groups in the skin follicles of mice and sheep and the entry of labelled sulphur compounds. Proceedings of the Royal Society of Edinburgh. 67, 65-82.

Schafzuchtverbände Niedersachsen. 2016. http://www.schafzucht-niedersachsen.de/Schafzucht-Verbaende-Niedersachsen/index.php?lang=de

The Shetland sheep Society 2018. http://www.shetlandsheep.org.uk/page.php?Plv=2&P1=5&P2=9&P3=

Statistisches Bundesamt, 2017. https://www.destatis.de/DE/Publikationen/StatistischesJahrbuch/StatistischesJahrbuch2017.pdf?__blob=publicationFile

Sympcam, 2017. 7th European Symposium on South American Camelids and 3rd European Meeting on Fibre Animals. June 12 -14, 2017. Assisi, Italy. http://www.sympcam.org/

Thomson, N. 2018, Farmers wool and traceability. Sympcam 2017.

http://www.sympcam.org/wp-content/uploads/2016/10/Book_of_Abtracts.pdf. P.23

Trah, M.; Wittek, T. 2013., Alpakas und Lamas, Fütterung und fütterungsbedingte Erkrankungen. 1. Auflage, LAMAS Verlag, Kaufbeuren, 9-12.

Valera, M.; Arrebola, F.; Juárez, M.; Molina, F.A. 2009, Genetic improvement of wool production in Spanish Merino sheep: Genetic parameters and simulation of selection strategies. Animal Production Science, 2009, 49, 43-47.
<figure><img src="image_9.png" /></figure>
# Effect of Technological Alternatives in the Mitigation of Climate Change in the Aging of Alpacas above 4,000 msnm Puno-Peru

T. Huanca, R.H. Mamani-Cato, M. Naveros and M. Gonzales

National Institute of Agrarian Innovation (INIA) – Peru

**Abstract.** The objective of the study was to evaluate the effect of technological alternatives in the mitigation of climate change in the breeding of alpacas over 4,000 meters above sea level. The validation study was conducted in 06 farming communities of two agroecological zones of the Puno region. The variables under study were weight at birth, weight at weaning, weight of fleece, pregnancy and mortality percentage, during 2014 in which no technological alternatives were applied, while in 2015 and 2016 this set was applied of alternatives. The data were analyzed in a completely randomized design using the SAS statistical software version 9.4. The results show that the birth weights were of 6.01, 6.254 and 6.83 kg for the years 2014, 2015 and 2016 respectively (p <0.05). The weaning weights were of 29.02, 29.73 and 30.99 kg for the years 2014, 2015 and 2016 respectively (p <0.05). The fleece weights were 2.28, 2.35 and 2.75 kg for the years 2014, 2015 and 2016 respectively (p <0.05). The pregnancy percentages were of 68.33, 72.84 and 83.77 % in the years 2014, 2015 and 2016 respectively. The mortality percentages were 10.39, 5.11 and 2.44 % for the years 2014, 2015 and 2016 respectively (p<0.05). The set of technological alternatives applied during the years 2015 and 2016 had a significant effect on the birth weight, weaning weight, fleece weight, pregnancy percentage and mortality of the alpacas.

**Resumen.** El objetivo del estudio fue evaluar el efecto de las alternativas tecnológicas en la mitigación del cambio climático en la crianza de alpacas sobre los 4,000 msnm. El estudio de validación se realizó en 06 comunidades campesinas de dos zonas agroecológicas de la región Puno. Las variables en estudio fueron el peso al nacimiento, peso al destete, peso de vellón, porcentaje de preñez y mortalidad, durante el año 2014 en el que no se aplicaron alternativas tecnológicas, en tanto que en los años 2015 y 2016 se aplicaron este conjunto de alternativas. Los datos se analizaron en un diseño completamente al azar mediante el programa estadístico SAS versión 9.4. Los resultados muestran que los pesos al nacimiento fueron de 6.01, 6.254 y 6.83 kg para los años 2014, 2015 y 2016 respectivamente (p<0.05). Los pesos al destete fueron de 29.02, 29.73 y 30.99 kg para los años 2014, 2015 y 2016 respectivamente (p<0.05). Los pesos vellones fueron de 2.28, 2.35 y 2.75 kg para los años 2014, 2015 y 2016 respectivamente (p<0.05). Los porcentajes de preñez fueron de 68.33, 72.84 y 83.77 % en los años 2014, 2015 y 2016 respectivamente. Los porcentajes de mortalidad fueron 10.39, 5.11 y 2.44 % para los años 2014, 2015 y 2016 respectivamente (p<0.05). El conjunto de las alternativas tecnológicas aplicadas durante los años 2015 y
2016 tuvieron un efecto significativo sobre el peso al nacimiento, peso al destete, peso
vellón, porcentaje de preñez y mortalidad de las alpacas.

**Keywords:** alpaca, technological alternatives, climate change

**Introduction**

The CSA have played a fundamental role in the development of Andean societies
from the old hunter communities to the current peasant communities (Mengoni,
2008). Before colonization domestic camelids were widely distributed from the
altitudes of the Andes to sea level. During the colonization they suffered the
uncontrolled sacrifice and were displaced by the domestic animals introduced by
the Europeans. This fact remains a clear example of ecological imperialism
(Crosby, 1986). As a consequence, both domestic and wild CSAs suffered a severe
reduction in number and their geographic distribution was drastically affected,
being reduced to the altitudes of the Andean highlands (Wheeler et al., 1995). The
CSA have the advantage of resisting adverse environments such as the one existing
in the Andean highlands. It is estimated that there are about seven million CSAs in
the Andean countries: Argentina, Bolivia, Chile, Colombia, Ecuador, Paraguay and
Peru (Fernández Baca, 2005, Raggi, 2005). Of these CSA, 51 % are in Peru and
34 % in Bolivia. Only in Peru are the four species of CSA, being this country
which houses the largest population of alpacas and vicuñas. The largest population
of llamas is found in Bolivia and guanacos in Argentina. Interest in llamas and
alpacas has increased in recent years in other countries including the United States,
Canada, Australia, New Zealand and some European countries such as the United
Kingdom, Germany, Italy and France (Brown, 2000; Sharpe et al., 2009). The
objective of the study was to evaluate the effect of technological alternatives in the
mitigation of climate change in the breeding of alpacas over 4,000 meters above
sea level.

**Materials and Methods**

The study was conducted in eight rural communities of the department of Puno
located above 4,000 meters above sea level in the agro-ecological zones of dry
puna and humid puna. The data for the analysis comes from the records of calving,
weaning, shearing, controlled enumeration and health, these were analyzed in a
completely randomized design using the statistical software SAS version 9.4. The
Duncan test was used for multiple comparisons with a level of significance of
α = 0.05.
## Results and Discussion

Table 1 shows the productive indices of three consecutive years, taking the year 2014 as a reference, as the year in which the technologies were not applied. The results show that the birth weights were of 6.01, 6.254 and 6.83 kg for the years 2014, 2015 and 2016 respectively (p<0.05). The weaning weights were of 29.02, 29.73 and 30.99 kg for the years 2014, 2015 and 2016 respectively (p<0.05). The fleece weights were 2.28, 2.35 and 2.75 kg for the years 2014, 2015 and 2016 respectively (p<0.05). The pregnancy percentages were of 68.33, 72.84 and 83.77 % in the years 2014, 2015 and 2016 respectively. The mortality percentages were 10.39, 5.11 and 2.44 % for the years 2014, 2015 and 2016 respectively (p<0.05). It is possible to observe that the effect of the application of technologies allowed to increase the weight at birth, weight at weaning, weight of fleece, also allowed to increase the pregnancy rate and decrease the percentage of mortality of the alpacas.

<table><caption>**Table 1:** Alpacas' productive indices according to production year</caption><thead><tr><th>Year of Production</th><th>Weight at birth kg</th><th>Weight at weaning kg</th><th>Weight of fleece kg</th><th>Pregnancy %</th><th>Mortality %</th></tr></thead><tbody><tr><td>2014</td><td>6.01 ± 1.36<sup>c</sup></td><td>29.02 ± 6.67<sup>b</sup></td><td>2.28 ± 0.62<sup>b</sup></td><td>68.33</td><td>10.39</td></tr><tr><td>2015</td><td>6.25 ± 1.27<sup>b</sup></td><td>29.73 ± 5.98<sup>a</sup></td><td>2.35 ± 0.65<sup>b</sup></td><td>72.84</td><td>5.11</td></tr><tr><td>2016</td><td>6.83 ± 1.21<sup>a</sup></td><td>30.99 ± 6.12<sup>a</sup></td><td>2.75 ± 0.59<sup>a</sup></td><td>83.77</td><td>2.44</td></tr></tbody></table>

## Conclusion

The application of technologies contributes to mitigate the effects of climate change at the level of peasant communities above 4,000 msnm, we observe the increase in weight at birth, weight at weaning, weight of fleece, pregnancy percentage and decrease in percentage of mortality of the alpacas.

## References

* Brown, B.W. 2000, A review on reproduction in South American camelids. Anim Reprod Sci. 58: 169-195.

* Crosby, A.W. 1986, Ecological Imperialism: The Biological Expansion of Europe, 900-1900. Cambridge University Press, Cambridge.

* Fernández Baca, S. 2005, Situación actual de los camélidos sudamericanos en Perú. Proyecto de Cooperación Técnica en apoyo de la crianza y aprovechamiento de los Camélidos Sudamericanos en la Región Andina TCP/RLA/2914. Organización de las Naciones Unidas para la Agricultura y la Alimentación (FAO).

* Mengoni, G.L. 2008, Camelids in ancient Andean societies: a review of the zooarchaeological evidence. Quaternary International. 185: 59-68.

<footnote><sup>abc</sup> Significant differences between means within column, p<0.05</footnote>
Raggi, L.A. 2005, Situación actual de los camélidos sudamericanos en Chile. Proyecto de Cooperación Técnica en apoyo a la crianza y aprovechamiento de los Camélidos Sudamericanos en la Región Andina. TCP/RLA/2914. Organización de las Naciones Unidas para la Agricultura y la Alimentación (FAO).

Sharpe, M.S.; Lord, L.K.; Wittum, T.E.; Anderson, D.E. 2009, Pre-weaning morbidity and mortality of llamas and alpacas. Aust Vet J. 87: 56-60.

Wheeler, J.C.; Russela, A.J.F.; Redden H. 1995, Llamas and Alpacas: Pre-conquest breeds and post-conquest hybrids. J Archaeol Sci. 22: 833-840.
# Collection of Diversity – Preserving Rare Indigenous Sheep Breeds in Germany

N. Ketterle

Kollektion der Vielfalt, Bosslerstraße 1, 73119 – Zell, Germany

E-Mail: info@kollektion-der-vielfalt.de

**Abstract.** The aim of the project is to preserve the following rare indigenous sheep breeds: Alpine Stonesheep, Black Mountain Sheep, Brown Mountain Sheep, Carniolan Stone Sheep, Coburger Foxsheep, German Karakul Sheep, Forest Sheep, Pomeranian Coursewool Sheep, Spectacle Sheep, Vallachian Sheep and White Mountain Sheep by using their unique wool. Wool quality becomes a very important breeding aspect in order to offer high quality products from the region for the region directly to the customer. One major challenge is to preserve the “wool infrastructure” in Europe, e.g. scouring plants and spinning mills and to modify the EU- regulation 1069/2009 which classifies greasy wool as an animal-by-product of category 3.

**Resumen.** El objetivo del proyecto es la conservación de las siguientes razas autóctonas locales: la raza ovina Alpine Stonesheep, la raza ovina Black Mountain, la raza ovina Brown Mountain, la raza ovina Carniolan Stone, la raza ovina Coburger Foxsheep, la raza ovina Karakul alemana, la raza ovina Forest, la raza ovina Pomeranian Coursewool, la raza ovina Spectacle, la raza ovina Vallachian y la raza ovina White Mountain mediante el aprovechamiento de su lana exclusiva. La calidad de la lana se convierte en un aspecto muy importante de mejora para ofrecer productos de alta calidad de la región directamente al cliente. Un desafío importante es preservar la "infraestructura de lana" en Europa, por ejemplo, limpiando plantas e hilanderías y modificando el reglamento de la UE 1069/2009, que actualmente clasifica la lana grasa como un producto derivado del animal de categoría 3.

**Keywords:** conservation, sheep, rare breeds

Many people know that wild animal and plant species are in danger of extinction but only a few are aware of the fact that a similar situation exists with regard to domesticated plant and animal species. A small number of high performance plant and animal strains produce the foodstuffs for the world's population – yet at the same time, a breed of farm animal disappears every two weeks. This means, a breed that successfully adapted to its climate and habitat, which has a genetic heritage and has become a cultural asset, is lost. Many people know about the “Red List” from the realm of wild plants and wild animals. “Red Lists” are registers of
extinct, missing and endangered species of animals, plants and fungi, plant communities as well as biotope types and biotope complexes. They are scientific, expert assessments outlining the current threat for a particular reference area, and evaluate the threat on the basis of population size and population development. Also in the agricultural context, a significant narrowing of genetic variety has already occurred and is still continuing.

In Germany, the Merinolandschaf and a few meat breeds numerically outnumber the other sheep breeds. On the other hand, there are more than 22 different sheep breeds on the Red List in Germany, mostly colored landrace breeds. Innovative ideas are required to utilize the products of these endangered breeds to ensure their long-term conservation.

In former times, farm animals were incorporated in farming in a variety of ways. In the case of cattle, not only their milk and meat supply was appreciated but cattle were also required to pull wagons and ploughs. Sheep were kept for the production of meat, milk and wool. Industrialization and mechanization in agriculture resulted in specialization, which in turn resulted in a departure from multiple-purpose breeds to one-purpose or dual-purpose breeds. In addition to this, the wool price in Europe declined more and more, resulting in wool playing a minor role in sheep husbandry if at all.

Our old and domestic sheep breeds are the result of centuries of breeding. They should therefore be considered a cultural asset worthy of conservation similar to heritage buildings, work of art or an old tree. The “other” performance capability of the old long-established sheep breeds is often underestimated and is even disregarded. They still have valuable characteristics such as a strong physique, frugality, longevity, high fertility and strong mothering abilities. These breeds are resistant to diseases and have adapted to their environment. Modern animal husbandry has led repeatedly to a dead end as a result of its performance breeding goals.

Our project is called “Collection of Diversity – Preserving Rare Indigenous Sheep Breeds” and was initiated in order to work especially with the marketing of particular endangered indigenous sheep breeds and their wool. We always aim to make the sheep breed and their typical wool attractive for both the breeder and the consumer. Furthermore we strive for environmentally friendly and local processing in the manufacturing process. All our woolen products are manufactured in rather small establishments, mostly family run businesses, in Germany, Austria and Northern Italy. Our current product range includes woolen products of ten different colored sheep breeds on the endangered list.

These breeds are:

* Alpines Steinschaf (“Alpine Stone Sheep”)

* Braunes Bergschaf (“Brown Mountain Sheep”)

* Brillenschaf (“Spectacles Sheep”)

* Deutsches Karakulschaf (German Karakul Sheep)

* Coburger Fuchsschaf (“Coburg Fox Sheep”)
* Krainer Steinschaf (“Carniola Stone Sheep”)

* Rauhwolliges Pommersches Landschaf (“Pomeranian Coarsewool Sheep”)

* Schwarzes Bergschaf (“Black Mountain Sheep”)

* Walachenschaf (“Valachian Sheep”)

* Waldschaf (“Forest Sheep”)

Every single one of these endangered colored sheep breeds has a very special wool, typical of the particular breed.

<figure><img src="image_10.png" /><figcaption>**Figure 1:** A group of Coburger Fuchsschaf at Arkfarm Ketterle. Photo: Arkfarm Ketterle</figcaption></figure>

The Coburger Fuchsschaf (“Coburg Fox Sheep”), for instance, plays a major role in the conservation of the countryside (Fig. 1). It is widespread throughout Germany and, aboveall, ideally adjusted to habitats in low mountain ranges. Its beautiful and unique wool is a distinctive mark of this old but endangered country breed. The wool of the mature animals has a golden hue; this is also referred to as “the Golden Fleece”. However, the fleece of the newborn lambs is reddish brown and brightens as they grow older. The variable tinges and the very good quality, in particular, contribute towards the popularity of its wool. The wool of this breed has a micron count of 33 and 36 (C-D assortment) and is well suited for spinning, knitting, weaving and felting. Furthermore, the wool also absorbs dye well during the dyeing process. The annual wool quantity of the ewes proves to be between 3,5 to 4,5 kg and, in the case of rams, around 5 kg. Due to the fact that the wool is colored it is not in demand with the wool industry and is therefore not used for industrial production in great quantities. As a result, the breeders were not in a position to sell the exceptional wool of their animals in the market place. This ultimate-
ly forced the breeders to dispose of their wool (burning, ploughing under and discarding) instead of selling it at a profit. Due to that there was no necessity to grow the typical “Golden Fleece Wool” of this breed. So it is important that breeders must be able to have the opportunity to sell their wool for a profit.

<figure><img src="image_11.png" /><figcaption>**Figure 2:** A mother with their Alpines Steinschaf lambs in different colours at Arkfarm Ketterle. Photo: Arkfarm Ketterle</figcaption></figure>

Another breed in our project is the Alpine Steinschaf (Fig. 2). The Alpines Steinschaf, a breed that is mostly found in the Eastern Alps of Bavaria/Germany and Salzburg/Austria and is listed on the Red List in Germany as “extremely endangered”. In 2004 there was a total stock of 123 sheep in flock books. Now, in 2016 there are 680 sheep in flock books. In 2004 the cross-border wool project of the Alpines Steinschaf began. Greasy wool is collected once a year in April at the annual breeders meeting. A big problem within this wool project is the small amounts of greasy wool. In 2004 we started the wool project with only 111kg greasy wool and two breeders. In 2017 913 kg greasy wool was collected from 19 different sheep breeders (Fig.3).
<figure><img src="image_12.png" /><figcaption>**Figure 3:** Evolution of breeders and greasy wool (kg) from the start of the project 2004.</figcaption></figure>

Since 2012 there was a decrease of greasy wool due to the EU-regulation 1069/2009 concerning the animal by-products came into effect on 1st March 2011. In this regulation greasy wool is classified as an animal by-product of category 3. In conclusion we are no longer authorized to collect greasy wool in Germany from Austrian breeders nor are German breeders authorized to collect greasy wool in Austria from Austrian breeders. So far there exists no registration form that will allow us collecting and transporting greasy wool from more than one EU-country. This is just permitted for professional wool merchants and wool trading organizations but not for small breeder organizations or NGO's.

The wool of the Alpine Steinschaf is coarse wool and it plays an important role in their survival in the high mountains. It's beautiful and unique wool is a special feature of this old breed. The Alpines Steinschaf have a double dual coated fleece with pithy, long coarse hair and fine wavy and short under coat. The woolly undercoat has a micron count of 38 to 48, an indication of the great diversity in this breed. Every wool color from white to black to brown as well as brindled is found. A ewe annually produces between 3 to 3.5 kg of wool and rams about 4 kg. Breeders have been unable to sell the wool of this breed for a profit because it was colored and too coarse. So far there was no use for the wool. Only some felters or hand spinners used the wool for handcrafts.

Another breed in our project is the Karakul. The Karakul is one of the oldest breeds of domestic animals in the world (Fig. 4). In 1900 the Karakul first came to Germany at the suggestion of a fur trade house in Leipzig. Since 1928 the Karakul is pure bred in Germany without substantial addition of homologous transfusion. The Karakul is a fat-tailed breed.
<figure><img src="image_13.png" /><figcaption>**Figure 4:** A herd of German Karakul (Deutsches Karakul) at Arkfarm Gerstner. Photo: Arkfarm Ketterle</figcaption></figure>

Their wool is coarse and carpet-like; their fleece is also strong, long-stapled and lightweight. It is also high yielding and double coated, with excellent felting qualities. The Karakuls are the only fur bearing sheep known. The lambs are born with a lustrous coat of fur instead of wool covering their bodies known as “Persian Lamb”. The lack of profitability of the breed as a result of the decline of fur prices meant that the Karakul almost extinct. In 1985 there were 1216 sheep in flock books and 125 breeders. In 2016 only 250 sheep in flock books and 6 breeders are left. For this reason and to save this rare sheep breed from extinction we started a wool project with 200 kg greasy wool and 2 breeders in 2016. The most difficulties we had in finding enterprises that are prepared and able to take in small amounts of wool and be able to produce high quality products of this really special wool.

The following example illustrates how innovative marketing concepts can pre-
serve a sheep breed that is close to extinction: The low wool price – selling the raw
wool directly after shearing does not even cover the cost of the shearing. This,
together with the low demand for smaller quantities of wool, two breeders of the
Alpines Steinschaf started to have their wool processed together in November
2004. In 2005, at the annual meeting of the breeders of Alpines Steinschaf, they
showcased the products made of their wool to attract more breeders for a joint
marketing of these unique woolen products. Here, the idea to launch a wool pro-
ject for the Alpines Steinschaf was born. With this founding, the cornerstone for a
successful marketing of exclusive, first-class woolen products of a particular region
was laid. At the annual meeting of the Alpines Steinschaf breeders in May, the
wool is jointly collected and sorted. The pre-sorted greasy wool currently fetches
2 € per kilogram in the form of a wool voucher. The aim is to further increase the
price so that it will be worthwhile for the breeders to produce wool again, which
means to breed Alpine Steinschafe with their typical wool. It was decided that
quality criteria for the wool should be incorporated in the breeding objectives. Collecting the wool at an annual meeting of breeders is also aimed at reducing transport costs as far as possible. The wool is, ideally suited for the production of felting, and knitting materials. That is why typical regional products are being manufactured from the different types of wool according to their structure and history of former production. At the same time our own logo was designed for the different woolen products. This is aimed at securing the brand recognition of these unique products and to further raise the awareness level. This is also the sign of quality, guaranteeing that all products with this special logo have been made exclusively from the wool of this breed. In this manner, the customer can ensure the survival of this rare indigenous sheep breed by buying products displaying its logo (Fig. 5).

<figure><img src="image_14.png" /><figcaption>**Figure 5:** Logo on wool products from rare indigenous sheep breeds.</figcaption></figure>

The current product range of the Alpines Steinschaf comprises, for example, carded wool, knitting wool, woolen socks, gloves and mittens, woolen beanies, jerseys, knitted sweaters, cardigans, jackets, coats, blankets, various felted bags, insoles, slippers, cushions and woven carpets. Every year more products are added.

The goals of the wool project called “Collection of Diversity – Preserving Rare Indigenous Sheep Breeds” are:

* The acquisition of pre-sorted raw wool at an appropriate price;

* Ecologically friendly processing and production of high quality woolen products;

* Transparent processing in small and mid-sized plants and businesses;

* Manufacturing in the region, thus supporting the regional economic potential;

* Manufacturing under socially acceptable working conditions;

* Supporting sustainable agriculture and conservation of the countryside by means of sheep husbandry;

* Promoting an old sheep breed on the brink of extinction thus supporting genetic variety and preserving a cultural asset.
References

AG Alpine Steinschafe (www.Alpines-Steinschaf.de)

AG Coburger Fuchsschafe (www.agfuchsschaf.de)

Bundesministerium für Ernährung, Landwirtschaft und Verbraucherschutz
(http://www.bmelv.de)

Bundesamt für Naturschutz (http://www.bfn.de)

Bundesamt für Naturschutz (http://www.bfn.de)

Gesellschaft zur Erhaltung alter und gefährdeter Haustierrassen e.V. (www.g-e-h.de)
Breeding and Genetics

<figure><img src="image_15.png" /></figure>
<figure><img src="image_16.png" /></figure>
# Advances in Llama (Llama glama) Coat Color Genetics

M.S. Daverio, M. Anello, L. Vidal-Rioja and F. Di Rocco

Laboratorio de Genética Molecular, Instituto Multidisciplinario de Biología Celular (IMBICE), CCT-CONICET-La Plata, CICPBA-UNLP, La Plata 1900, Buenos Aires, Argentina

E-Mail: fdirocco@imbice.gov.ar

**Abstract.** Domestic species present a wide phenotypic diversity. As their genomic sequences become available, finding the connection between the genotype and the observed phenotype is a goal possible to achieve. In several species, the genes and mutations responsible for the different coat colors and patterns have already been identified. Color is a Mendelian trait, with little or no environmental influence. Moreover, the biochemical pathway of pigment synthesis is well-known and highly conserved among mammals. Thus, in species such as llama whose complete genome has not yet been sequenced, the study of candidate genes results a valid alternative to identify the genetic variation responsible for different phenotypes. The objective of this work is to present the recent advances in the knowledge of the genes that control the coat color in llamas and compare our results with those published for alpaca.

**Resumen.** Las especies domésticas presentan una amplia diversidad fenotípica. A medida que sus secuencias genómicas están disponibles, encontrar la conexión entre el geno-tipo y el fenotipo observado es una meta posible de lograr. En varias especies, ya se han identificado los genes y mutaciones responsables de los diferentes colores de capa. El color es un carácter mendeliano, con poca o ninguna influencia ambiental. Además, la ruta bioquímica de la síntesis de la melanina es bien conocida y altamente conservada entre los mamíferos. Por lo tanto, en especies como la llama, cuyo genoma completo aún no ha sido secuenciado, el estudio de genes candidatos resulta una alternativa válida para identificar la variación genética responsable de diferentes fenotipos. El objetivo de este trabajo es presentar los recientes avances en el conocimiento de los genes que controlan el color de capa en llamas y comparar nuestros resultados con los publicados para la alpaca.

**Keywords:** llamas, coat color, MC1R, ASIP, polymorphisms

Melanogenesis is the process of forming melanin, a pigment that gives color to the skin, hair and eyes. The melanin synthesis is produced in the melanosome, a specialized organelle inside the melanocyte. The two types of pigments that are
responsible for color are eumelanin (dark brown/black) and pheomelanin (red/yellow). The ratio between both pigments is controlled by the interaction between the melanocortin receptor (MC1R) and its antagonist, the agouti signaling protein (ASIP). The binding of the α-melanocyte-stimulating hormone (α-MSH), which is the natural ligand of the MC1R, causes the exclusive production of eumelanin, whereas Mc1r inhibition by ASIP leads to the synthesis of pheomelanin (Lu et al., 1994). Polymorphisms in the genes encoding these two proteins have been associated with coat color variation in mammals and birds (Kijas et al., 1998; Andersson et al. 2003, Schmutz et al., 2003; 2003; Kerns et al., 2004).

In many species, dominant MC1R alleles (E) produce a uniform black color, while recessive alleles (e) produce red-yellow pigment. Conversely, dominant alleles of the Agouti locus produce a yellow coat whereas the recessive allele in homozygosis is associated with a black coat. In addition to mutations in the genes that control the type of pigment produced, mutations in other genes related to melanogenesis also affect the coat color. For example, genes that affect correct migration and differentiation of the melanocytes, such as KIT or MITF are responsible for unpigmented or spotted phenotypes in most species (Brooks and Bailey, 2005; Pielberg et al., 2002; Cooper et al., 2005). Llamas and alpacas are characterized by a wide variety of coat colors and patterns, although molecular basis of coat color determination is not yet fully understood in camelids. However, in recent years some advances have been made in the knowledge of the genes that control the pigmentation in these species. Molecular genetics studies in alpacas have identified mutations in the MC1R and ASIP genes that are associated to coat color (Powell et al., 2008; Feeley and Munyard 2009; Chandramohan et al., 2011; Feeley et al., 2011; Chandramohan et al., 2013). Much more recently, the first work characterizing these genes and their relation to the color in llamas has been published (Daverio et al., 2016).

The goal of this work is to present the progress made so far in llama coat color genetics, and discuss it with regard to the alpaca findings. To understand how color variation occurs, it is necessary to first understand how the eumelanic and phaeomelanic coats are produced. For these, the coding region of MC1R and ASIP were sequenced in animals with different phenotypes: 1) Eumelanic (animals that have black coat with their variants or dark brown coat) 2) pheomelanic (animals with reddish brown coat) 3) mixed (pheomelanic coat with black face and trims) 4) white (non-albino), representing absence of pigmentation in the coat. Moreover, samples of the llama wild ancestor, the guanaco (*Lama guanicoe*), were also sequenced. Screening for genetic variation in 84 llamas showed 13 Single nucleotide polymorphism (SNPs) in the MC1R gene, 10 of which were non-synonymous. Two of them, c.205C > A and c.638G>A, are novel SNPs that had not been previously described in alpacas. The combination of three polymorphisms, c.259G>A, c.376G>A and c.383T > C, defined three main haplotypes (or alleles) for MC1R: MC1R*1 (c.259A/c.376A/c.383T), MC1R*2(c.259G/c.376G/c.383C) and MC1R*3 (c.259G/c.376G/c.383T). The guanaco sequences showed no variation,
and were identical to MC1R3. Allele frequencies were similar in reddish and black animals. However, MC1R*1 showed a significant association with pigmented coat (P <0.0001). This allele was not found in any of the 29 white llamas analyzed. In this last phenotypic group, MC1R*2 was found at frequency significantly higher than in the other groups. Moreover, the wild genotype (MC1R*3/MC1R*3) was observed in ten individuals, four “black face” and six white animals, suggesting the influence of other genes in the production of those phenotypes.

MC1R mutations responsible for coat color in llamas appear to be different from those reported in alpacas. Feeley and Munyard (2009) found that alpacas with haplotype combination c.82A/c.126T/c.901C (E/E or E/e) in MC1R are able to produce eumelanin whereas animals which have the combination c.82G/c.126C/c.901T (e/e), only express pheomelanin. We found that SNP c.901C > T, c.82A > G and c.126T > C appeared in very low frequency, being almost fixed for the ‘eumelanic’ combination proposed by those authors in the alpaca. Different results were found by Chandramoan et al. (2011) for alpaca: animals homozygous for A82/A259/A376/C901 expressed black phenotypes whereas white animals were homozygous for the combination of 82/G259/G376/T901. In llamas, the c.259A/c.376A/c.383T haplotype is associated to a pigmented coat, but not exclusively eumelanic and c.259G/c.376G/c.383C (MC1R*2) is found in high frequency in white animals. Moreover, almost half of the white llamas were homozygous MC1R*2/ MC1R*2, a genotype not observed in any of the llamas with pigmented coat. In principle, we did not expect any particular genotype at the MC1R in white animals since white is caused in other species by mutations located in other genes, which are dominant and epistatic to MC1R. The c.383T>C substitution in the MC1R*2 produces a p.M128T change in the protein, in a site highly conserved in mammals. Functional studies in humans confirmed that this variant, found in malignant melanoma, shows marked loss of function and reduced agonist binding affinity. Nevertheless, for the moment we have no direct experimental evidence of the impact of this substitution on the llama protein. As MC1R alleles were not specifically associated with pheomelanic/eumelanic phenotypes in llamas, the complete coding region of ASIP was also analyzed in the same animals. Two polymorphisms were found within exon 4, a 57 bp deletion (c.325_381del) and a nonsynonymous SNP (c.292C > T). Both polymorphisms are predicted to have a deleterious effect on the protein (Feeley et al., 2011).

All llamas with a pheomelanic coat carried at least one copy of the non-deleted (wild type) ASIP allele. Instead, 17 from 19 black llamas were homozygous for the deletion, homozygous for the T variant of c.292C > T or heterozygous for the combination of both. This is consistent with a recessive inheritance mode for black color proposed for llamas (Frank et al, 2006). The same two variants have been previously associated to black color in alpacas (Feeley et al., 2011; Chandramohan et al., 2013). Moreover, a third substitution c.353G>A associated with this phenotype has been described but has not been yet identified in llamas. Although we have now molecular evidence of how eumelanic and phaeomelanic phenotypes can
be produced, some questions remain to be answered. Segregation analysis has shown that the white phenotype is dominant to pigmented phenotypes in llamas and alpacas (Frank et al, 2006). So, why are MC1R*2/MC1R*2 llamas white? Why they are not pheomelanic? Why llamas with wild genotype MC1R *3/MC1R*3 can also have a white coat? Non albino-white phenotypes are caused in most species by mutations in MITF and KIT genes. On these bases, these genes were afterward sequenced. To this objective, we obtained total RNA from skin biopsies and the complete cDNA sequence was determined in 23 llama samples. Neither polymorphisms in the coding region nor splicing variants associated with coat color were found. Therefore, to further understand how white phenotype in llama is produced studies on the expression of KIT, MITF and some other genes involved in melanogenesis are currently underway within our group.

## References

* Andersson L. 2003, Melanocortin receptor variants with phenotypic effects in horse, pig, and chicken. Annals of the New York Academy of Sciences, 994: 313-8.

* Brooks, S.A. and Bailey, E. 2005, Exon skipping in the KIT gene causes a sabino spotting pattern in horses. Mammalian Genome, 16:893-902.

* Chandramohan, B., Renieri, C., La Manna, V., La Terza, A. 2013, The alpaca agouti gene: genomic locus, transcripts and causative mutations of eumelanic and pheomelanic coat color. Gene, 521 (2): 303-310.

* Chandramohan, B., Manna, V.L., Renieri, C., Terza, A.L. 2011, Asip and MC1R cDNA polymorphism in alpaca. Fibre production in South American camelids and other fibre animals. pp. 93-96.

* Cooper, M.P., Fretwell, N., Bailey, S.J., Lyons, L.A. 2005, White spotting in the domestic cat (Felis catus) maps near KIT on feline chromosome B1. Animal Genetics, 37: 163-165.

* Daverio, M.S., Rigalt, F., Romero, S.R., Vidal Rioja, L., Di Rocco, F. 2016, Polymorphisms in MC1R and ASIP genes and its association with coat color phenotypes in llamas (Lama glama). Small Ruminant Research, 144: 83-89.

* Feeley, N.L., Munyard, K.A. 2009, Characterisation of the Melanocortin 1 Receptorgene in Alpaca and identification of possible markers associated with phenotypic variations. Animal Production Science Journal, 49 (8): 675-681.

* Feeley, N.L., Bottomley, S., Munyard, K.A. 2011, Three novel mutations in ASIP associated with black fiber in alpacas (Vicugna pacos). Journal of Agricultural Science, 149 (4):529-538.

* Frank, E.N., Hick, M.V.H., Gauna, C.D., Lamas, H.E., Renieri, C., Antonini, M. 2006, Phenotypic and genetic description of fibre traits in South American domestic camelids (Llamas and alpacas). Small Ruminant Research, 61: 113-129.

* Kerns, J.A., Newton, J., Berryere, T.G., Rubin, E.M., Cheng, J.F., Schmutz, S.M., Barsh, G.S. 2004, Characterization of the dog Agouti gene and a non agouti mutation in German Shepherd Dogs. Mammalian Genome, 15 (10): 798-808.

* Kijas, J.M., Wales, R., Tornsten, A., Chardon, P., Moller, M., Andersson, L. 1998, Melanocortin receptor 1 (MC1R) mutations and coat color in pigs. Genetics, 150(3): 1177-1185.
Lu, D., Willard, D., Patel, I.R., Kadwell, S., Overton, L., Kost, T., Luther, M., Chen, W., Woychik, R.P., Wilkison, W.O., et al. 1994, Agouti protein is an antagonist of the melanocyte-stimulating hormone receptor. Nature, 371: 799-802.

Pielberg, G., Olsson, C., Syvänen, A.C., Andersson, L. 2002, Unexpectedly high allelic diversity at the KIT locus causing dominant white color in the domestic pig. Genetics, 160: 305-311.

Powell, A.J., Moss, M.J., Tree, L.T., Roeder, B.L., Carleton, C.L., Campbell, E., Kooyman, D.L. 2008, Characterization of the effect of Melanocortin 1 Receptor, a member of the hair color genetic locus, in alpaca (Lama pacos) fleece colordifferentiation. Small Ruminant Research, 79: 183-187.

Schmutz, S.M., Berryere, T.G., Ellinwood, N.M., Kerns, J.A., Barsh, G.S. 2003, MC1R studies in dogs with melanistic mask or brindle patterns. Journal of Heredity, 94: 69-73.
<figure><img src="image_17.png" /></figure>
# Characterization and Expression Analysis of SLC7A11 in Llamas

M. Anello<fnref n="1" />, E. Fernandez<fnref n="1" />, M. Silbestro<fnref n="1" />, F. Veiga<fnref n="2" />, L. Vidal Rioja<fnref n="1" /> and F. Di Rocco<fnref n="1" />

E-Mail: fdirocco@imbice.gov.ar

**Abstract.** The llama (*Lama glama*) is a South American camelid which is gaining worldwide recognition for its fiber. Coat color is one of its most cherished and commercially important characteristics; however, there is very little information about the molecular mechanisms that control pigmentation in llamas. SLC7A11 (solute carrier family 7 number 11) encodes the light chain of cystine/glutamate exchanger, xCT, which has a major role in pheomelanin synthesis. Reports on these issues indicate for instance, that in mouse a mutation in SLC7A11 gene produce a diluted coat color due to a reduction in pheomelanin production. Chintala et al. (2005). Furthermore, differences in the skin expression levels of SLC7A11 have been observed in brown and white alpacas (Tian et al., 2015). The aim of this study is to describe SLC7A11 coding region and to analyze its variation and expression in llamas of different coat color. For this purpose, skin biopsies from 13 animals were collected, the RNA extracted and total cDNA obtained. cDNA was used for PCR amplification and sequencing of the full coding region. In addition, expression levels were analyzed by Real Time PCR. Coding region of SLC7A11 consisted of 1,512 bp that encoded a 503-amino acid protein. Protein sequence analysis showed 12 transmembrane helix regions with cytoplasmic N-terminal and C-terminal residues. Analysis with BLASTP showed 91-99 % identity to other mammals as well as a highly conserved amino acid permease domain. Seven SNPs were observed in the llama DNA sequences, 6 synonymous and 1 non synonymous. Finally, preliminary results indicated that expression levels of SLC7A11 in undiluted phenotypes differ significantly from those in diluted and white, but no differentiation was found between diluted and white phenotypes.

**Resumen.** La llama (*Lama glama*) es un camélido sudamericano que está empezando a ganar reconocimiento mundial por su fibra. El color de la fibra es una de las características de mayor valor comercial, sin embargo, hay muy pocos estudios sobre los mecanismos moleculares que controlan la pigmentación en llamas. El gen SLC7A11 (transportador soluble de la familia 7, número 11) codifica la cadena liviana del intercambiador de cistina/glutamato, xCT, y cumple un rol importante en la síntesis de feomelanina. Chintala et

<footnote n="1">¹Laboratorio de Genética Molecular, Instituto Multidisciplinario de Biología Celular (IMBICE), CCT-CONICET-La Plata, CICPBA-UNLP, La Plata 1900, Buenos Aires, Argentina</footnote>

<footnote n="2">²Facultad de ciencias veterinarias, Universidad de Buenos Aires, Buenos Aires, Argentina</footnote>
al. (2005) encontraron que una mutación este gen es responsable del fenotipo subtle gray (sut) en ratón. Los mutantes sut muestran un color de capa diluido que se debe a una reducción de la síntesis de feomelanina. Además, alpacas marrones y blancas presentan diferencias en los niveles de expresión de SLC7A11 (Tian et al., 2015). El propósito de este trabajo es describir la región codificante de SLC7A11 y analizar su variación y expresión en llamas con distinto fenotipo de color Para ello se tomaron biopsias de piel, se extrajo el ARN y se obtuvo el cADN total. Este último fue empleado para reacciones de PCR y posterior secuenciación de la región codificante completa. Los niveles de expresión se analizaron mediante PCR en tiempo real. La región codificante de SLC7A11 está compuesta por 1512 pb y codifica una proteína de 503 aminoácidos. El análisis de la secuencia de la proteína muestra 12 regiones de tipo hélice transmembrana con los residuos C-terminal y N-terminal de orientación citoplasmática. Mediante el análisis con BLASTP se observó entre 91-99 % de identidad con otros mamíferos y la presencia de un dominio aminoácido permeasa altamente conservado. Se identificaron siete SNPs en las secuencias de ADN de llamas, 6 sinónimos y 1 no sinónimo. Finalmente, de acuerdo a resultados preliminares de expresión, los niveles de SLC7A11 en fenotipos no diluidos fueron significativamente diferentes de los hallados en fenotipos diluidos y blancos.

**Keywords:** lamas, coat color, SLC7A11

## Introduction

The fiber of llama (*Lama glama*) is increasingly appreciated by the textile industry. Its cost depends primarily on the diameter and the color of the fiber. However, there is little knowledge about the molecular basis of coat color determination, which makes it very difficult for breeders to obtain the expected color phenotypes or reduce those of less commercial value.

In mammals, the basic coat colors are defined by the relationship between two pigments: eumelanin (from black to brown) and pheomelanin (from red to yellow). Eumelanin/pheomelanin ratio is regulated mainly by the ligand-receptor system of the agouti signaling protein (ASIP) and the melanocortin 1-receptor (MC1R). The binding of alpha-melanocyte stimulating hormone (α-MSH) to MC1R leads to eumelanin synthesis while binding of ASIP inhibits signal transduction, causing the melanocytes to produce pheomelanin (Lu et al., 1994). However, the final color phenotype will also depend on the expression and interaction of many other genes involved in processes such as the development and differentiation of melanocytes, melanosome formation and pigment synthesis, pigment transport and transference to tissues and survival of melanocyte stem cells. Mutations affecting genes involved in any of these processes can disrupt the normal pigmentation pathway producing diluted and white phenotypes.

The different coat color phenotypes in llamas were described by Frank (2001) and Frank et al. (2006). Based on these descriptions and performing classical crossbreeding analysis, those authors studied the segregation of color phenotypes and postulated that pigmented phenotypes are segregated by the Agouti locus.
Nevertheless, the molecular basis of coat color determination in these species has not yet been established. Recently, we have sequenced the coding region of MC1R and ASIP in llamas and studied the association between polymorphisms in these genes and coat color variation (Daverio et al., 2016). An interesting finding in that work was the detection of association between MC1R*2 haplotype and white coat. However, white llamas were also homozygous for haplotype MC1R*3, which was found in colored phenotypes as well. This suggests that other genes might be involved in the production of white phenotypes. In most species, mutations in MITF and KIT genes are responsible of white phenotype (Haase et al., 2007; Pielberg et al., 2002), but we have studied the complete coding region of these genes in llamas and found no differences between white and colored phenotypes (Anello et al., 2015, 2016).

Extreme dilution of pheomelanin has been proposed as a possible mechanism for white or cream phenotypes in other species. Sponenberg and Rothschild, (2001) described in dogs an Intense (I) locus that is thought to dilute only pheomelanin. However, I locus has not been characterized yet. Several genes have been involved in color dilution in different mammal species; one of them is SLC7A11 (solute carrier family 7 number 11). This gene encodes the light chain of cystine/glutamate exchanger, xCT, which has a major role in pheomelanin synthesis. Cystine is a precursor for pheomelanin synthesis but it is not necessary for the synthesis of eumelanin. Therefore, mutations in SLC7A11 are supposed to produce only pheomelanin dilution, without affecting eumelanin. There are only a few reports about this gene and even less that analyses its relationship with pigmentation. One of them is the one carried out by Chintala et al. (2005). They found that a mutation in SLC7A11 gene was responsible for the subtle gray (sut) phenotype in mouse; sut mutants show a diluted coat, due to a reduction in pheomelanin production. In another work, Tian et al. (2015) studied brown and white alpacas and found that there were differences in the expression levels of SLC7A11 between these two phenotypes.

Based on the above information, the aim of the present study is to describe SLC7A11 coding region and to analyze its variation and expression in llamas of different coat color phenotype.

## Materials and Methods

Skin biopsies from llamas with white (n=6) and pheomelanic (n=7) phenotypes were collected by disposable biopsy punch (3 mm diameter), following the Argentinean Ethical Guidelines for Biomedical investigation in Animals from Laboratory, Farm or obtained from Nature (Resolution D N° 1047/05 from CONICET, Argentina). Biopsies were conserved in RNAlater (SIGMA, Germany) until their further extraction. Total RNA was extracted by homogenization in TRIzol® and addition of chloroform to separate phases. The aqueous phase was used for the alcoholic precipitation of RNA. Then, it was washed and resuspended
in RNase free water. Reverse transcription to obtain cDNA was performed in a
20 µl reaction volume, using 1 µg/µl RNA, RevertAid Reverse Transcriptase
(Thermo Fisher Scientific) and random primers (Biodynamics), following the
manufacturer's instructions.

PCR primers were designed over conserved regions from others mammal sequences available in GenBank, to fully cover SLC7a11 coding region and flanking 5' and 3' UTRs. Amplification reactions were carried out in 25 µl PCR mix containing 1X PCR Buffer (Invitrogen, Carlsbad, CA, USA), 2 mM MgCl2, 0.2 mM dNTPs, 60 µg BSA, 1 U Taq DNA polymerase (Invitrogen, Carlsbad, CA, USA), 0.5 µM of each primer and 50 ng of cDNA. The cycling profile consisted of an initial denaturation step at 94 °C for 3 min, 30 cycles of 40 sec at 94 °C, 50 sec at 55-57 °C, 40 sec at 72 °C and a 5-min final extension at 72 °C. PCR products were checked on a 1% agarose gel stained with GelRedTM, purified by PEG precipitation and sequenced. Sequences obtained were aligned and analyzed using Geneious (v.6.1.8, Biomatters). Additionally, comparison with the alpaca sequence available at GenBank (KM095134) was carried out. Protein sequence analysis was performed with TMHMM server (http://www.cbs.dtu.dk/services/TMHMM) for the prediction of transmembrane regions, Pfam (http://pfam.xfam.org) for the identification of conserved domains and BLASTP (blast.ncbi.nlm.nih.gov) for protein homology.

For the expression analysis the samples were divided into three color groups: undiluted pheomelanic (n=3), diluted pheomelanic (n=3) and white (n=6). Quantitative Real Time PCR was carried out using specific primers designed over the llama SLC7A11 coding region; 18S gene was used as endogenous control. Amplification reaction consisted of 20 µl, including 4 µl of HOT FIREPol® EvaGreen® qPCR Mix Plus (ROX) (Solís Biodyne), 0.5 mM of each primer and 1 ng cDNA. The cycling parameters were: 15 min at 95 °C, 40 cycles of 15 sec at 95 °C, 20 sec at 60 °C, 20 secs at 72 °C, and a final gradient from 95 °C to 72 °C. Every sample was analyzed in triplicate on a RotorGene Q (Qiagen) equipment. Quantification of SLC7A11 transcript abundance was performed using the comparative threshold cycle (CT) method established by Livak and Schmittgen (2001). Finally, ANOVA analysis of variance was used to assess if differences in expression were significant.

Results

Complete coding region of the llama SLC7A11 gene consists of 1,512 bp divided
into 12 exons. The protein encoded has 503 amino acids and, when aligned with
other mammal species, it presented high identity with its homologues: from 99 %
with Vicugna pacos and Camelus ferus to 91 % with Mus musculus. The protein
displays one highly conserved functional domain Amino Acid Permease_2 located
at amino acids 72-461, and 12 transmembrane helix regions with citoplasmatic N-
terminal and C-terminal residues.
We detected 3 positions in the llama SLC7A11 gene that differ from the ones present in the alpaca sequence: c.15 T>C, c.18 G >T and c.445 C> T. The first two are located in exon 1 and represent synonymous substitutions, while the third one is located in exon 3 and it produces a change from His to Tyr.

Seven SNPs were found along the llama sequences, of which 6 are synonymous changes and one is a non-synonymous substitution. Table 1 shows the extension of each exon in the coding sequence (CDS) and the location of the polymorphisms.

<table><caption>**Table 1:** SLC7A11 exons and polymorphisms in llama</caption><thead><tr><th>Exon</th><th>Extension in CDS</th><th colspan="2">Polymorphisms</th><th colspan="2">Change in protein</th></tr></thead><tbody><tr><td>Exon 1</td><td>1 - 277</td><td colspan="2">-</td><td colspan="2">-</td></tr><tr><td>Exon 2</td><td>278 - 403</td><td>c.298 T>C</td><td>c.381 C>T</td><td>no</td><td>no</td></tr><tr><td>Exon 3</td><td>404 - 520</td><td colspan="2">c.418 G>T</td><td colspan="2">p.140 A>S</td></tr><tr><td>Exon 4</td><td>521 - 646</td><td colspan="2">c.522 T>C</td><td colspan="2">no</td></tr><tr><td>Exon 5</td><td>647 - 746</td><td colspan="2">-</td><td colspan="2">-</td></tr><tr><td>Exon 6</td><td>747 - 791</td><td colspan="2">-</td><td colspan="2">-</td></tr><tr><td>Exon 7</td><td>792 - 915</td><td colspan="2">c.837 T>C</td><td colspan="2">no</td></tr><tr><td>Exon 8</td><td>916 - 1019</td><td colspan="2">c.984 T>C</td><td colspan="2">no</td></tr><tr><td>Exon 9</td><td>1020 - 1116</td><td colspan="2">-</td><td colspan="2">-</td></tr><tr><td>Exon 10</td><td>1117 - 1266</td><td colspan="2">-</td><td colspan="2">-</td></tr><tr><td>Exon 11</td><td>1267 - 1444</td><td colspan="2">c.1380 G>T</td><td colspan="2">no</td></tr><tr><td>Exon 12</td><td>1445 - 1512</td><td colspan="2">-</td><td colspan="2">-</td></tr></tbody></table>

We also studied the relative expression of *SLC7A11* in the skin of undiluted, diluted and white phenotypes (Figure 1). SLC7A11 expression levels in the non-diluted group differ significantly from those in diluted and white (p<0.05) animals. Although expression level was consistently lower in white llamas than in the diluted ones, the difference was not significant.

## Discussion

There are very few studies addressing SLC7A11 gene variation and its role in mammal's pigmentation. Here, we described the coding region of this gene in llamas and found results that support previous information (Tian et al., 2015). SLC7A11 codes for a highly conserved protein with an AA_permease_2 domain, which presents 12 transmembrane helices and it has the function of transport of amino acids. Variation observed in the llama SLC7A11 gene was mainly due to synonymous substitutions and the unique non-synonymous SNP found does not seem to be associated to color dilution.
<figure><img src="image_18.png" /><figcaption>**Figure 1:** Relative expression of SLC7A11 in llamas.</figcaption></figure>

SLC7A11 expression levels in undiluted llamas differ significantly from those in diluted and white. This is consistent with the protein function: it allows cystine to enter the cell and cysteine is necessary for the pheomelanin synthesis. Therefore, less expression of SLC7a11 should be translated into a reduction in pheomelanin synthesis. However, though it is evident that level of the diluted group is in between the others two, mRNA expression levels between diluted and white were not significantly different. This could be due to a small difference that would need a larger sample to be detectable. Moreover, as pheomelanic coats can vary from dark red to very light brown, the results obtained may depend on the manner in which the animals from the diluted group were chosen. In other words, at the lower end of the dilution range, expression values close to those of the group of white animals would be expected, while at the other end values should be more similar to those of animals with undiluted coats. This discussion raised the question if white coat could be an extreme dilution of pheomelanin, There is not enough information to answer this question yet, but considering the results from this work and previous findings from other genes (Daverio et al., 2016, Anello et al. 2015, Anello et al., 2016) it is likely that more than one genetic mechanism is responsible for white color in llamas.

Further research in needed to understand how SLC7a11 expression is con-
trolled. We cannot rule out regulatory mutations affecting SLC7A11, but it is also
possible that another gene, located upstream in the pigmentation pathway is down-
regulating the expression of SLC7A11in white and diluted llamas.

**Acknowledgments.** Authors would like to thank for their collaboration with the samples Dr. Marcelo Miragaya and the staff from the Faculty of Veterinary Sciences, UBA, the breeder Mr. Carlos Rusconi and his staff. F. Di Rocco is researcher from the Scientific
Research Commission (CIC), province of Buenos Aires. This work was supported by institutional grants 2015 and 2016 from CIC and funds from STAN 486/12 IMBICE-CONICET, Argentina.

## References

Anello, M., Silbestro, M., Vidal Rioja, L., Di Rocco, F. 2015, Caracterización de KIT: un gen candidato para el color de capa en llamas (Lama glama). Libro virtual del VII Congreso Mundial en Camélidos Sudamericanos DOI:10.13140/RG.2.1.1863.0487

Anello, M., Silbestro, M., Vidal Rioja, L., Di Rocco, F. 2016, Characterization of MITF coding region in llamas. Abstract book: 35rd International Society for Animal Genetics Conference. Salt Lake City, Utah. p.:107

Chintala, S., Li, W., Lamoreux, M.L., Ito, S., Wakamatsu, K., Sviderskaya, E.V., Bennertt, D.C., Young-Mee, P., Gahl, W.A., Huizing, M., Spritz, R.A., Ben, S., Novak, E.K., Tan, J. and Swank, R.T. 2005, Slc7a11 gene controls production of pheomelanin pigment and proliferation of cultured cells. Proc Natl Acad Sci U S A. 2005; 102:10964-10969

Daverio, M.S., Rigalt, F., Romero, S., Vidal Rioja, L., Di Rocco, F. 2016, Polymorphisms in MC1R and ASIP genes and their association with coat color phenotypes in llamas (Lama glama). Small Ruminant Res. 144, 83-89

Frank, E.N., Hick, M.V.H., Gauna, C.D., Lamas, H.E., Renieri, C., Antonini, M. 2006, Phenotypic and genetic description of fiber traits in South American domestic camelids (Llamas and alpacas). Small Rumin. Res. 61, 113-129.

Frank, E.N. 2001, Descripción Y Análisis De Segregación De Fenotipos De Color Y Tipos De Vellón En Llamas Argentinas PhD Thesis. UBA (pp.204).

Haase, B., Brooks, S.A., Schlumbaum, A., Azor, P.J., Bailey, E., Alaeddine, F., Mevissen, M., Burger, D., Poncet, P.A., Rieder, S., Leeb, T. 2007, Allelic heterogeneity at the equine KIT Locus in dominant white (W) horses. PLoS Genet. 3 (11), e195.

Livak, K.J., Schmittgen, T.D. 2001, Analysis of relative gene expression data using real time quantitative PCR and the 2-ΔΔCT method. Methods 25, 402-408.

Lu, D., Willard, D., Patel, I.R., Kadwell, S., Overton, L., Kost, T., Luther, M., Chen, W., Woychik, R.P., Wilkison, W.O., et al. 1994, Agouti protein is an antagonist of the melanocyte-stimulating hormone receptor. Nature 371, 799-802.

Pielberg, G., Olsson, C., Syvänen, A.C., Andersson, L. 2002, Unexpectedly high allelic diversity at the KIT locus causing dominant white color in the domestic pig. Genetics 160, 305-311.

Sponenberg D.P., Rothschild M.F. 2001, Genetics of coat colour and hair texture. In: Ruvinsky A, editor. Genetics of the dog. New York: CABI Pres

Tian, X., Meng, X., Wang, L., Song, Y., Zhang, D., Ji, Y., Li, X., and Dong, C. 2015, Molecular cloning, mRNA expression and tissue distribution analysis of Slc7a11 gene in alpaca (Lama pacos) skins associated with different coat colors. Gene. 555:88-94.
<figure><img src="image_19.png" /></figure>
# PCR-RFLP Method for Testing ASIP EXON 4 Mutations in Llamas

M.S. Daverio, V. Alcoela-Ersinger, M. Anello, L. Vidal-Rioja and F. Di Rocco

Laboratorio de Genética Molecular, Instituto Multidisciplinario de Biología Celular (IMBICE), CCT-CONICET-La Plata, CICPBA-UNLP, La Plata 1900, Buenos Aires, Argentina

E-Mail: fdirocco@imbice.gov.ar

**Abstract.** The basic coat colors of mammals are determined by the relative proportion of two types of pigments, eumelanin and pheomelanin. The agouti signaling protein (ASIP) plays a crucial role in melanogenesis, by increasing the production of pheomelanin and decreasing the eumelanin synthesis by blocking the signaling pathway of the melanocortin 1 receptor (MC1R). In several species, loss of function mutations of the ASIP gene is responsible for the black coat color. The polymorphisms of ASIP exon 4, c.325_381del and c.292C> T, have been previously associated with eumelanic phenotypes in llamas (Daverio et al., 2016) and also in alpacas (Chandramohan et al., 2013). The objective of this work was to develop an alternative method to DNA sequencing for genotyping both polymorphisms. Fifty one llama DNA samples, including controls of known sequence were analyzed. PCR products spanning ASIP exon 4 were digested with the enzyme Bvel, that recognizes the c.292C>T mutation, followed by electrophoresis in 8% polyacrylamide gels to simultaneously detect the SNP and the deletion. Analysis of the band patterns presented complete concordance between DNA sequencing and PCR-RFLP genotypes. Genotyping results from the samples showed that all dark llamas (n=13) were homozygous for the deletion, homozygous for the c.292C> T polymorphism or heterozygous for both, while none of these combinations were observed in the pheomelanic animals here analyzed (n=20). The results of this work support the findings of previous studies and also show the usefulness of the PCR-RFLP technique as a relatively fast, simple and cost-effective method to determine the ASIP exon 4 variants in llamas.

**Resumen.** Los colores de capa básicos de los mamíferos están determinados por la proporción relativa de dos tipos de pigmentos, eumelanina y feomelanina. La proteína de señalización agouti (ASIP) juega un rol crucial en la melanogénesis, incrementando la producción de feomelanina y disminuyendo la síntesis de eumelanina por bloqueo de la vía de señalización del receptor 1 de melanocortina (MC1R). En varias especies, mutaciones con pérdidas de función del gen ASIP son responsables del color de capa negro. Los polymorfismos del exón 4 de ASIP, c.325_381del y c.292C> T, han sido asociados previamente con fenotipos eumelánicos en llamas (Daverio et al., 2016) y también en alpacas (Chandramohan et al., 2013). El objetivo de este trabajo fue desarrollar un método alternativo a la
secuenciación de ADN para genotipar ambos polimorfismos. Se analizaron 51 muestras de ADN de llamas, incluyendo controles de secuencia conocida. Los productos de PCR que abarcan el exón 4 de ASIP fueron digeridos con la enzima Bvel, que reconoce la mutación c.292C> T, seguida por electroforesis en geles de poliacrilamida al 8 % para detectar simultáneamente el SNP y la delección. El análisis de los patrones de banda presentó concordancia completa entre los genotipos de secuenciación y los de PCR-RFLP. Los resultados de genotipado de las muestras mostraron que todas las llamas oscuras (n=13) eran homocigotas para la delección, homocigotas para el polimorfismo c.292C> T o heterocigotas para ambos, mientras que ninguna de esas combinaciones se observó en los animales feomelánicos aquí analizados (n=20). Los resultados de este trabajo apoyan los hallazgos de estudios previos y también muestran la utilidad de la técnica de PCR-RFLP como un método relativamente rápido, sencillo y de bajo costo para determinar las variantes del exón 4 de ASIP en llamas.

**Keywords:** llamas, PCR-RFLP, ASIP-genotyping, eumelanic

**Introduction**

Coat color in animals has important functions in camouflage, protection against UV radiation and communication, among others. Compared to the wild species, domestic animals present a fascinating diversity of colors product of artificial selection (Reissmann & Ludwig, 2013). During the last decade, molecular techniques have allowed the identification of genes and alleles underlying this variation in a large number of species. The color of the coat basically depends on the proportions of the pigments eumelanin (black or dark brown) and pheomelanin (reddish-yellow). The production of these pigments is mainly controlled by the melanocortin-1 receptor (MC1R) and agouti signaling peptide (ASIP) (Bultman et al., 1992). MC1R interaction with its antagonist ASIP produces pheomelanic pigments by blocking the interaction with its natural ligand alpha-MSH (Lu et al., 1994). Dominant black is usually produced by variations in the coding sequence of the MC1R gene (Klungland et al., 1995; Marklund et al., 1996; Kijas et al., 1998). Gain of function mutations that produce constitutive activation of MC1R lead to the synthesis of eumelanin and a dark coat. On the other hand, in species where the black color is recessive, non-functional alleles of the ASIP gene are responsible for the dark coloration. Loss-of-function ASIP mutations have been described in species such as dog (Kerns et al., 2004), horse (Rieder et al., 2001), sheep (Royo et al., 2008). In alpaca, three mutations in ASIP exon 4 c.325_381del, c.292C>T and c.353G>A have been also associated to black coat (Feeley et al., 2011; Chandramohan et al., 2013). In llama, the mutations c.325_381del and c.292C>T have been also showed association with black color but the c.353G>A has not been identified so far (Daverio et al., 2016).

In Argentina, the llama is the most abundant domestic camelid and exhibits a wide variety of coat colors that meet different commercial interests. Although the
value of camelids fiber greatly depends on its diameter, the color is also important for the textile industry (Frank et al., 2006, Mueller et al., 2010). In this context, black, reddish brown and golden colors are usually some of the most required (Frank et al., 2006). However, the color of offspring based only on the parents' phenotype is hard to predict in camelids. Thus, the use of molecular tests to determine the genotype of the breeding animals could be useful in selection programs. Nevertheless, if the number of animals is high, sequencing may be expensive and additionally it requires specialized equipment, not always available in small laboratories. The main objective of this work was to develop a simple technique based on Polymerase Chain Reaction and Restriction Fragment Length Polymorphism (PCR-RFLP) to detect c.325_381del and c.292C>T polymorphisms, as an alternative to DNA sequencing.

## Materials and Methods

Thirty three blood llama samples (20 pheomelanic and 13 eumelanic) from diverse herds from Argentina were collected by jugular puncture. Total genomic DNA was isolated following the LiCl protocol described in Daverio MS (2015).

Using the PCR technique, exon 4 of the ASIP gene was amplified in the 33 samples and 18 controls of known sequence. Amplification reactions were carried out as described in Daverio et al., (2016), using as template 60-70 ng of genomic DNA. The PCR products were digested using the restriction enzyme Bvel (BspMI). The enzyme cuts the amplicon into two fragments, one of 83 bp and the other one of 374 bp or 317 bp depending on whether the allele carries or not the c.325_381del. If the c.292C>T polymorphism is present, it creates a new cutting site for Bvel generating a different banding profile.

Digestion mix consisted of 6 µl of PCR product, 6.5 U of Bvel (Thermo Fisher Scientific Inc.), 0.3 µl of Oligos 50X, 1 µl of Buffer O 10X and 6 µl of bidistilled water that was incubated at 37 °C overnight. Restriction fragments were analyzed by electrophoresis on an 8 % polyacrylamide gel for 90 min at 200 V stained with GelRedTM and visualized using an UV trans illuminator. Fragment length determination was done by comparison with a 50 bp commercial ladder (Embiotec SRL).

## Results and Discussion

We adjusted amount of PCR product, digestion time and enzyme quantity to standardize PCR-RFLP conditions. Digestion of control samples yielded band patterns in all cases concordant with genotypes previously obtained by ASIP exon 4 sequencing. Homozygous genotypes for c.292T that did not have the c.325_381del mutation (T/T, -/-) showed two fragments of 192 and 182 bp, in addition to the 83 bp band present in all digestions. Homozygous deletion genotypes (C/C, del/del), generated a 317 bp fragment while controls without the
deletion (C/C, -/-) showed a 374 bp band. Heterozygous individuals for one or both polymorphisms also had the expected combined patterns: 374 bp and 317 bp (C/C,-/del); 317 bp, 192 and 182 (C/T, -/del); 374 bp, 192 and 182 bp (C/T, -/-) (Fig. 1.)

<figure><img src="image_20.png" /><figcaption>**Figure 1:** Electrophoresis pattern of ASIP exon 4. Lines 1 to 5 and 7, 9, 11, 13, 15: digested PCR products. Lines 8, 10, 12, 14 and 16: PCR products before digestion. Line 6: molecular size marker of 50 bp. Line 5: incomplete digestion.</figcaption></figure>

After confirming that digestion patterns were as expected, unknown samples were genotyped. PCR-RFLP analysis revealed 6 different genotypes (Table 1). No shared genotypes were found between the two color groups; three were exclusive of eumelanic samples and the remaining three of pheomelanic animals.

The two mutations here studied have been associated to the recessive black coat in alpacas (Feeley et al., 2011; Chandramohan et al., 2013) and llamas (Daverio et al., 2016). Substitution c.292C> T, causes a p.R98C change that introduces an additional cysteine into the region of ASIP which makes contact with MC1R (Kiefer et al., 1997). The other mutation, c.325_381del, leads to the loss of 19 amino acids (p.C109_R127del) within the same domain. Both mutations are predicted to be deleterious and to alter normal protein function. Therefore, the mutated ASIP protein is unable to bind MC1R to induce pheomelanin production and only eumelanin is synthesized, resulting in a black coat. In agreement with that, we found that black llamas were all homozygous for c.292T (T/T), homozygous c.325_381del (del/del), or in a lower frequency (3/13), compound heterozygotes for both mutations, whereas these genotypes were not found in pheomelanic individuals. Thus, our results provide additional support to the previously reported
association between ASIP variants and coat color in domestic camelids (Feeley et al., 2011; Chandramohan et al., 2013, Daverio et al., 2016)

<table><caption>Table 1: ASIP exon 4 genotypes in eumelanic and pheomelanic llamas</caption><thead><tr><th></th><th>Genotype</th><th colspan="2">Phenotype</th></tr></thead><tbody><tr><td>c.292C>T</td><td>c.325_381del</td><td>Pheomelanic</td><td>Eumelanic</td></tr><tr><td>C/C</td><td>del/del</td><td>0</td><td>6</td></tr><tr><td>T/T</td><td>-/-</td><td>0</td><td>4</td></tr><tr><td>C/T</td><td>-/del</td><td>0</td><td>3</td></tr><tr><td>C/C</td><td>-/-</td><td>9</td><td>0</td></tr><tr><td>C/T</td><td>-/-</td><td>2</td><td>0</td></tr><tr><td>C/C</td><td>-/del</td><td>9</td><td>0</td></tr></tbody></table>

del: deletion

-: non deleted variant

## Conclusion

The PCR-RFLP technique proved to be useful for genotyping c.325_381del and c.292C>T polymorphisms. It offers the advantage of being based on a single PCR reaction and is suitable for reading on polyacrylamide gels.

The confirmation of the association between these variants and the black coat still requires the analysis in a larger number of animals and further family based segregation studies. The methodology presented here would be adequate for this purpose, since it allows to genotype the two variants simultaneously, without needing of DNA sequencing.

**Acknowledgments.** We would like to thank Miriam B. Silbestro for her technical assistance. This work was supported by grants from The National Scientific and Technical Research Council (CONICET) and from the Commission for Scientific Research of the Province of Buenos Aires (CIC) from Argentina.

## References

* Bultman, S.J., Michaud E.J., Woychik R.P. 1992, Molecular characterization of the mouse agouti locus. Cell, 71:1195-1204

* Chandramohan, B., Renieri, C., La Manna, V., La Terza, A. 2013, The alpaca agouti gene: genomic locus, transcripts and causative mutations of eumelanic and pheomelanic coat color. Gene, 521(2):303-10

* Daverio, M.S. 2015, Caracterización de genes vinculados al crecimiento y al color de capa en la Llama (Lama glama) (Characterization of genes related to growth and coat color in Llama (Lama glama). PhD Thesis, FCE-UNLP, 109pp. http://hdl.handle.net/10915/40703

* Daverio, M.S., Rigalt, F., Romero, S., Vidal-Rioja, L., Di Rocco, F. 2016, Polymorphisms in MC1R and ASIP genes and their association with coat color phenotypes in llamas (Lama glama). Small Ruminant Research, 144:83-89
Feeley, N.L., Bottomley, S., Munyard, K.A. 2011, Three novel mutations in ASIP associated with black fibre in alpacas (Vicugna pacos). Journal of Agricultural Science, 149 (4):529-538

Frank, E.N., Hick, M.V.H., Gauna, C.D., LamaS, H.E., Renieri, C., Antonini, M. 2006, Phenotypic and genetic description of fibre traits in South American domestic camelids (Llamas and alpacas). Small Ruminant Research, 61:113-129

Kerns, J.A., Newton, J., Berryere, T.G., Rubin, E.M., Cheng, J.F., Schmutz, S.M., Barsh, G.S. 2004, Characterization of the dog Agouti gene and a non agouti mutation in German Shepherd Dogs. Mammalian Genome, 15(10):798-808

Kiefer, L.L., Ittoop, O.R.R., Bunce, K., Truesdal, A.T., Willard, D.H., Nichols, J.S., Blanchard, S.G., Mountjoy, K., Chen, W.J., Wilkison, W.O. 1997, Mutations in the carboxyl terminus of the agouti protein decrease agouti inhibition of ligand binding to the melanocortin receptors. Biochemistry, 36:2084-2090

Kijas, J.M., Wales, R., Tornsten, A., Chardon, P., Moller, M., Anedersson, L. 1998, Melanocortin Receptor 1 (MC1R) mutations and coat color in pigs. Genetics 150 1177-1185

Klungland, H., VagE Di, Gomez-Raya, L., Adalsteinsson, S., Lien, S. 1995, The role of melanocyte-stimulating hormone (MSH) receptor in bovine coat color determination. Mammalian Genome 6, 636-639

Lu, D., Willard, D., Patel, I.R., Kadwell, S., Overton, L., Kost, T., Luther, M., Chen, W., Woychik, R.P., Wilkison, W.O., and Col. 1994, Agouti protein is an antagonist of the melanocyte-stimulating hormone receptor. Nature, 371:799-802

Marklund, L., Moller, M.J, Sandberg, K., Andersson, L. 1996, A missense mutation in the gene for melanocyte-stimulating hormone receptor (MC1R) is associated with the chestnut coat color in horses. Mammalian Genome 7, 895-899

Mueller, J.P., Rigalt, F., Cancino, A.K., Lamas, H. 2010, Calidad de las fibras de camélidos sudamericanos en Argentina (Quality of the fibers of South American camelids in Argentina). Comunicación Técnica INTA EEA Bariloche Nro PA 573, 20 pág.

Reissmann, M., Ludwig, A. 2013, Pleiotropic effects of coat colour-associated mutations in humans, mice and other mammals. Seminars in Cell and Developmental Biology. 24(6-7):576-86

Rieder, S., Taourit, S., Mariat, D., Langlois, B., Guerin, G. 2001, Mutations in the agouti (ASIP), the extension (MC1R), and the brown (TYRP1) loci and their association to coat colour phenotypes in horses (Equus caballus). Mammalian Genome 12, 450-455

Royo, L.J., Alvarez, I., Arranz, J.J., Fernández, I., Rodriguez, A., Pérez-PardaL, L., Goyache, F. 2008, Differences in the expression of the ASIP gene are involved in the recessive black coat colour pattern in sheep: evidence from the rare Xalda sheep breed. Animal Genetics 39, 290-293
# Heredabilidad estimada de fibras meduladas en alpaca huacaya

R. Pinares<fnref n="1" />, A. Cruz<fnref n="2" />, R. Morante<fnref n="2" />, I. Cervantes<fnref n="4" />, A. Burgos<fnref n="2" />, G. Gutiérrez<fnref n="3" />, J.P. Gutiérrez<fnref n="4" />

**Abstract.** The aim was to estimate the heritability of medullated fibers in alpaca Huacaya. Samples from 36 fleeces of white males (0.4-10.4 years old), from Pacomarca farm (Puno, Perú) were took. The diameter was recorded in each fiber (FD) following the IWTO-8-2011 regulation, and simultaneously the class of medullation (CM) was recorded as stablished in the literature (Villarroel, 1963) using the projection microscope (PM). For estimating genetic parameters of CM and FD was used an animal model with repeated measurement, the age was considered as a linear and quadratic covariate, the additive genetic effect and the permanent environmental as random effects. Continuous and threshold models were evaluated; considering 4 univariate models to estimate the heritability of CM and 4 bivariate models to estimate the heritability as well as the genetic correlation between CM and DF which were analyzed using TM software. The estimate heritability of CM was 0.36 ± 0.13 under a bivariate continuous model where CM was non-medullated versus medullated. Using the same model, the estimate genetic correlation between CM and FD was 0.93 ± 0.12. In sum, if the amount of medullated fibers is reduced, the individual fiber diameter in alpaca fleece will decrease.

**Resumen.** El objetivo fue estimar la heredabilidad de fibras meduladas en alpaca Huacaya. Se tomaron muestras de 36 vellones blancos de alpacas machos (0.4-10.4 años) del fundo Pacomarca (Puno, Perú). Se registró el diámetro de cada fibra (DF) siguiendo la norma IWTO-8-2011, y simultáneamente se registró la clase de medulación (CM) según se describe en la bibliografía (Villarroel, 1963), mediante el microscopio de proyección (MP). Para la estimación de los parámetros genéticos de CM y DF, se usó un modelo animal con medidas repetidas. La edad se consideró como covariable lineal y cuadrática, el efecto gen-

<footnote n="1">¹Facultad de Zootecnia. Universidad Nacional Agraria de la Molina – UNALM.
Av. La Molina S/N. La Molina, Lima, Perú.</footnote>

<footnote>E-Mail: zoovet_p@hotmail.com</footnote>

<footnote n="2">²Fundo Pacomarca – Inca Tops S.A, Avda. Miguel Forga 348, Arequipa, Perú.</footnote>

<footnote n="3">³Programa de Mejoramiento Animal (PMA), Universidad Nacional Agraria La Molina UNALM. Av. La Molina S/N. La Molina, Lima 12, Perú.</footnote>

<footnote>E-Mail: gustavogr@lamolina.edu.pe</footnote>

<footnote n="4">⁴Departamento de Producción Animal, Universidad Complutense de Madrid, Avda. Puerta de Hierro s/n E-28040-Madrid, España.</footnote>

<footnote>E-Mail: gutgar@vet.ucm.es</footnote>
ético aditivo del animal y su ambiente permanente como efectos aleatorios. Se evaluaron modelos continuos y umbrales, considerando 4 modelos univariados para estimar la heredabilidad de CM y 4 modelos bivariados para estimar las heredabilidades de CM y DF así como su correlación genética, mediante el software TM. La heredabilidad estimada de CM fue de 0.36 ± 0.13 bajo un modelo continuo bivariado donde CM se consider en dos categorías: no medulada versus medulada. Utilizando el mismo modelo se estimó una correlación genética de 0.93 ± 0.12 entre CM y FD. En conclusión, si se reduce la cantidad de fibras meduladas por selección, también se disminuirá el diámetro individual de la fibra.

**Keywords:** alpaca, medullated fiber, heritability, genetic correlations

## Introducción

La calidad de la fibra de alpaca es considerada mundialmente como una de las mejores para la industria textil. Sin embargo, su precio aún está lejos de otras fibras finas como la cachemira. Quizás el principal factor que reduce su valor es la presencia de fibras meduladas por lo que se ha asociado con el factor de picazón en la prendas (Frank et al., 2014). En el vellón de alpacas, incluso las fibras más finas pueden tener medulación fragmentada o discontinua.

En el fundo Pacomarca se ha realizado grandes esfuerzos para eliminar esta característica en particular. Realizando el descerdado manual (eliminación de fibras meduladas) se podría reducir inmediatamente la picazón por ende mejoraría el confort de las prendas (Laime et al., 2016). Sin embargo, el descerdado manual no sería rentable económicamente (Quispe et al., 2015). Por lo que se busca animales con bajo contenido de medulación en el vellón y el propósito de este trabajo fue estimar la heredabilidad y la correlación genética entre la medulación y el diámetro de cada fibra.

## Materiales y métodos

Se registraron un total de 21600 fibras individuales de 36 muestras de vellones blancos de alpacas Huacaya machos entre 0.4 y 10.4 años del fundo experimental Pacomarca del Grupo Inca (Puno, Perú). El diámetro individual de fibra (FD) se registró aleatoriamente (600 fibras por animal) siguiendo la norma IWTO-8-2011 (IWTO, 2011); por microscopio de proyección (PM) se clasificaron en cinco categorías de medulación (Villarroel, 1963): no medulada, fragmentada, discontinua, continua y fuertemente medulada.

El pedigrí de los individuos registrados fue rastreado a los ancestros hasta los fundadores para completar un pedigrí de 121 individuos, con el 100 % de los padres conocidos, así como el 71 % de los abuelos, el 20 % de los bisabuelos y menos del 1 % de los bisabuelos identificados. La matriz de relaciones numéricas fue calculada utilizando el software Endog v4.8 (Gutiérrez y Goyache, 2005) donde se mostró una fuerte conexión entre los animales.
El grupo de machos fue tomado al azar, donde se observa una conexión entre ellos, justificando su representatividad en toda la población. Por otra parte, aun cuando el número de animales es escaso, el alto número de registros ayudó a obtener estimaciones confiables y la fiabilidad final de fue conocido mediante el enfoque Bayesiano que ayuda a cuantificar adecuadamente los niveles de incertidumbre, una vez que la convergencia fue probada.

La medulación aquí se ha codificado como una característica discreto. Por lo tanto, los modelos umbrales son los indicados para la estimación de los parámetros genéticos para este rasgo. Sin embargo, se ha demostrado que los modelos lineales tienen mejores resultados que los modelos umbrales cuando las bases de datos son pequeñas (Ibáñez et al., 2014).

Para la medulación las categorías evaluadas fueron: C2A, dos categorías: 1: agrupada no medulada, fragmentada y la dicontinua, 2: agrupada la continua y la fuertemente medulada. C2B, dos categorías: 1: no medulada 2: agrupada todas las meduladas.

Las dos formas de clasificación se analizaron utilizando dos metodologías diferentes. El modelo lineal continuo (L) y el modelo umbral (T); la ecuación del modelo umbral fue idéntico al modelo lineal lineal, pero en este caso el vector y explica la categoría visible conocido como umbrales (Gianola, 1982). Finalmente, todas las combinaciones también fueron probadas usando 4 modelos univariados y 4 modelos bivariados que combina el diámetro de la fibra y la medulación. Así, se llevaron a cabo 8 estimaciones diferentes de componentes de varianza para la medulación.

La ecuación del modelo animal con medidas repetidas fue:

$$y = Xb + Zu +Wp + e$$

(1) donde y vector de observaciones (categoría de medulación o diámetro de fibra individual); b vector de efectos fijos: edad, edad al cuadrado; u vector de efectos genéticos aditivos; p vector de ambiente permanente; e vector de efectos residuales; X, Z y W son matrices de incidencia de efectos fijos, efecto genético del animal y efectos del medio ambiente permanente, respectivamente.

La ecuación del modelo umbral fue idéntico, pero en este caso el vector y representa una variable subyacente llamada problema que explica la categoría visible correspondiente definida por valores específicos llamados umbrales (Gianola, 1982). El análisis se llevó a cabo con el software TM (Legarra et al., 2011). Un millón de iteraciones se realizaron para cada modelo con un período de ciclo de 100.000 y con un intervalo fino de cada 100 iteraciones.

## Resultados y discusión

Las estimaciones de los componentes de la varianza para diferentes agrupaciones de categorías de medulación, se muestran en la Tabla 1, junto con las respectivas desviaciones estándar de sus distribuciones marginales posteriores. Utilizando
diferentes modelos que combinan modelos univariados o bivariados con el diámetro de fibra, bajo modelos lineales o umbrales.

Las estimaciones de heredabilidad para el rasgo de la médula variaron de 0.30 a 0.36 en los modelos lineales y de 0.11 a 0.15 en los modelos de umbrales. Todas las estimaciones de heredabilidad para categorías de medulación pueden considerarse significativas de acuerdo con la magnitud de la desviación estándar de sus distribuciones marginales posteriores, mostrando que la cantidad de información utilizada fue suficiente para proporcionar estimaciones significativas.

Las estimaciones de la heredabilidad del diámetro de cada fibra fueron entre 0.28 ± 0.14 y 0.35 ± 0.15, que son mucho menos precisos, de acuerdo con sus respectivas desviaciones estándar, que los de la medulación.

Por otro lado, la correlación genética estimada entre el diámetro individual y la medulación fue muy alta utilizando modelos continuos de 0.88 a 0.93, aunque el uso de modelos de umbral fue moderado e impreciso de 0.26 a 0.68. En ovejas Corriedale, Sánchez et al. (2016) reportaron una correlación genética de magnitud intermedia (0.50) entre el diámetro medio de fibra y la presencia versus ausencia de fibra medulada.

En ovejas Corriedale, utilizando un modelo animal multivariado, Sánchez et al. (2016) reportaron una heredabilidad de 0.37 ± 0.10 para la presencia versus ausencia de medulación; similar a la heredabilidad estimada en el presente estudio (0.36 ± 0.13). Sin embargo, resulta mayor a 0.23 ± 0.02 del contenido de medullation OFDA en cabras Angora, aunque similar a 0.32 ± 0.02 del contenido de fibras kemp (Allain y Roguet, 2006). Se han reportado heredabilidades similares para el porcentaje de medulación total (0.29 ± 0.04) y de 0.33 ± 0.07 para el porcentaje de fibras fuertemente meduladas (Frank et al., 2011).

Por último, una limitación de nuestra investigación fue un número limitado de animales (36) debido a que el tiempo requerido para procesar una muestra tomó alrededor de 1 día por PM y además requiere personal capacitado. También Sánchez et al. (2016) indicaron 1,5 días por animal. Sin embargo, por OFDA 100 analiza en menos de 8 minutos una muestra (Lupton y Pfeiffer, 1998), lo cual permite registrar más animales, basado en la opacidad (>80 %) agrupando fibras meduladas continuas y fuertemente meduladas de forma similar a C2A que se registró por MP.

## Conclusión

Los modelos lineales continuos estimaron mejores heredabilidades de CM y altas correlaciones genéticas entre CM y DF. Así se reduciría la cantidad de fibras meduladas y el diámetro de cada fibra en el vellón de alpaca Huacaya.
<table><caption>**Tabla 1:** Parámetros genéticos estimados ± desviación estándar de sus distribuciones marginales posteriores, para categorías de medulación y el diámetro individual de fibra bajo modelos lineales (L) y umbrales (T)</caption><thead><tr><th rowspan="2">Modelo</th><th colspan="6">Categoría de Medulación</th><th colspan="3">Diámetro de Fibra</th></tr><tr><th>σ<sup>2</sup><sub>u</sub></th><th>σ<sup>2</sup><sub>ep</sub></th><th>h<sup>2</sup></th><th>c<sup>2</sup></th><th>R</th><th>r<sub>g</sub></th><th>σ<sup>2</sup><sub>u</sub></th><th>σ<sup>2</sup><sub>ep</sub></th><th>h<sup>2</sup></th><th>c<sup>2</sup></th><th>R</th></tr></thead><tbody><tr><td>Uni; L;</td><td>0.09±</td><td>0.05±</td><td>0.30±</td><td>0.15±</td><td>0.45±</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>C2A</td><td>0.04</td><td>0.04</td><td>0.12</td><td>0.11</td><td>0.06</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Uni; L;</td><td>0.14±</td><td>0.07±</td><td>0.35±</td><td>0.17±</td><td>0.52±</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>C2B</td><td>0.07</td><td>0.05</td><td>0.13</td><td>0.12</td><td>0.06</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Bi; L; C2A</td><td>0.10±</td><td>0.07±</td><td>0.30±</td><td>0.20±</td><td>0.50±</td><td>0.91±</td><td>25.5±</td><td>17.0±</td><td>0.34±</td><td>0.23±</td><td>0.58±</td></tr><tr><td></td><td>0.05</td><td>0.04</td><td>0.12</td><td>0.11</td><td>0.07</td><td>0.13</td><td>14.14</td><td>11.63</td><td>0.15</td><td>0.15</td><td>0.07</td></tr><tr><td>Bi; L; C2B</td><td>0.16±</td><td>0.10±</td><td>0.36±</td><td>0.21±</td><td>0.57±</td><td>0.93±</td><td>26.0±</td><td>17.0±</td><td>0.35±</td><td>0.23±</td><td>0.58±</td></tr><tr><td></td><td>0.08</td><td>0.06</td><td>0.13</td><td>0.12</td><td>0.07</td><td>0.12</td><td>14.09</td><td>11.88</td><td>0.15</td><td>0.14</td><td>0.07</td></tr><tr><td>Uni; T;</td><td>0.13±</td><td>0.08±</td><td>0.11±</td><td>0.07±</td><td>0.14±</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>C2A</td><td>0.07</td><td>0.06</td><td>0.06</td><td>0.05</td><td>0.04</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Uni; T;</td><td>0.19±</td><td>0.09±</td><td>0.15±</td><td>0.07±</td><td>0.22±</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>C2B</td><td>0.09</td><td>0.07</td><td>0.06</td><td>0.06</td><td>0.05</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Bi; T; C2A</td><td>0.14±</td><td>0.11±</td><td>0.11±</td><td>0.08±</td><td>0.19±</td><td>0.26±</td><td>24.1±</td><td>14.6±</td><td>0.34±</td><td>0.21±</td><td>0.55±</td></tr><tr><td></td><td>0.08</td><td>0.06</td><td>0.05</td><td>0.05</td><td>0.05</td><td>0.51</td><td>12.76</td><td>10.60</td><td>0.15</td><td>0.14</td><td>0.07</td></tr><tr><td>Bi; T; C2B</td><td>0.17±</td><td>0.11±</td><td>0.13±</td><td>0.09±</td><td>0.21±</td><td>0.56±</td><td>24.6±</td><td>15.5±</td><td>0.34±</td><td>0.22±</td><td>0.55±</td></tr><tr><td></td><td>0.09</td><td>0.07</td><td>0.06</td><td>0.05</td><td>0.05</td><td>0.45</td><td>14.08</td><td>11.63</td><td>0.16</td><td>0.15</td><td>0.07</td></tr></tbody></table>

σ²u: varianza genética aditiva, σ²ep: varianza ambiental permanente, c²: varianza de ambiente permanente/varianza fenotípica,

h²: heredabilidad, R: repetibilidad, ru: correlación genética, Uni: uniuariado, Bi: bivariado, C2A: agrupada sin medulación, medulación fragmentada y discontinua vs medulación continua y fuertemente medulada; C2B: sin medulación vs medulación
**Agradecimientos.** Animal fibre working group of EAAP. Programa de Investigación y Proyección Social en Ovinos y Camélidos Americanos-UNALM.

Referencias

Allain, D., Roguet, J.M. 2006, Genetic and non-genetic variability of OFDA medullated fiber contents and other fleece traits in the French Angora goats. Small Ruminant Research, 65:217-222

Frank, E.N., Hick, M.V.H., Molina, M.G., Caruso, L.M. 2011, Genetic parameters for fleece weight and fiber attributes in Argentinean Llamas reared outside the Altiplano. Small Ruminant Research, 99:54-60

Frank, E.N., Hick, M.V.H., Castillo, M.F., Prieto, A., Adot, O.G. 2014, Fiber-based components determining handle and skin comfort in fabrics made from dehaired and non dehaired llama fiber. International Journal of Applied Science and Technology, 4:51-66

Gianola, D. 1982, Theory and Analysis of Threshold Characters. Journal of Animal Science 54, 1079-1096

Gutiérrez, J.P., Goyache, F. 2005, A note on ENDOG: a computer program for analysing pedigree information. Journal of Animal Breeding and Genetics, 122: 172-176.

Ibáñez, B., Cervantes, I., Gutiérrez, JP., Goyache, F., Moreno, E. 2014, Estimates of direct and indirect effects for early juvenile survival in captive populations maintained for conservation purposes: the case of Cuvier's gazelle. Ecology and Evolution 4.

IWTO-8, 2011, Fiber Diameter Distribution Parameters and Percentage of Medullated Fibers in Wool and Other Animal Fibers by the Projection Microscope. International Wool Textile Organization

Laime, F.M., Pinares, R., Paucara, V., Machaca, V., Quispe, E.C., 2016, Características Tecnológicas de la Fibra de Llama (Lama glama) Chaku antes y después de Descerdar. Revista de Investigaciones Veterinarias del Perú, 27(2):209-217

Legarra, A., Varona, L., López De Maturana, E. 2011, TM Threshold Model. Retrieved on 12 February 2017, from http://snp.toulouse.inra.fr/~alegarra/manualtm.pdf

Lupton, C.J., Pfeiffer, F.A. 1998, Measurement of Medullation in Wool and Mohair using an Optical Fiber Diameter Analyser. Journal of Animal Science, 76:1261-1266

Quispe, E.C., Chipa, L., Pinares, R. 2015, Análisis económico y de la producción del descerdado manual de la fibra de llamas (Lama glama) Chaku. Archivos de Zootecnia, 64:191-198

Sánchez, A.L., Urioste, J.I., Peñagaricano, F., Neimaur, K., Sienra, I., Naya, H., Kremer, R. 2016, Genetic parameters of objectionable fibers and of their associations with fleece traits in Corriedale sheep. J Anim Sci 94:13-20

Villarroel, J. 1963, Un estudio de la fibra de alpaca. Anales Científicos de la Universidad Nacional Agraria La Molina – UNALM 1(3):246-274
# Performance Evaluation of Llama, Alpaca and Sheep Herds of a Community in Pasco, Peru

D.M. Pizzaro¹, G.A. Gutiérrez², J.A. Ñaupari³ and M. Wurzinger⁴

¹Animal Science Faculty. Universidad Nacional Agraria La Molina – UNALM. Av. La Molina S/N. La Molina, Lima 12, Perú.

E-Mail: dpizarro@lamolina.edu.pe

²Animal Breeding Program. Universidad Nacional Agraria La Molina – UNALM. Av. La Molina S/N. La Molina, Lima 12, Perú.

E-Mail: gustavogr@lamolina.edu.pe

³Rangeland Ecology and Utilization Laboratory. Universidad Nacional Agraria La Molina – UNALM. Lima, Perú, Av. La Molina S/N. La Molina, Lima 12.

E-Mail: jnaupariv@lamolina.edu.pe

⁴Division of Livestock Sciences. Department of Sustainable Agricultural Systems. University of Natural Resources and Life Sciences – BOKU. Gregor-Mendel-Str. 33, A-1180 Vienna, Austria.

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

**Abstract.** In the Peruvian highlands livestock production is especially developed in the South and Center of the country between altitudes ranging from 2,200 to 4,500 meters above sea level. Community-owned pasture is the main source of animal feed in this region. Llama rearing is commonly combined with other species, such as alpacas and sheep. One way to technically evaluate livestock systems is by calculating technical parameters. These indicators allow to synthesize the information contained in the production records and to standardize the evaluation criteria of the production unit. The following technical parameters were used in this study: Annual average capital (AAC), gross birth rate (%GBR), real birth rate (%RBR), mortality (%M), harvest (%H), gross increase (%GI), real increase (%RI) and livestock efficiency (%LE). In the present study, the technical evaluation of livestock was carried out using the monthly records of llamas, alpacas and sheep of the Communal Cooperative San Pedro de Racco, located in Pasco Region, Peru. The evaluation period for llamas and alpacas was from 2012 to 2015, whereas for sheep the evaluation period was from January 2014 to December 2015. It was concluded that the Community Cooperative San Pedro de Racco presented values of technical parameters corresponding to a good technological level company, possibly due to the capacity development of human resources in terms of animal and grassland management.

**Resumen.** La ganadería altoandina en el Perú se desarrolla especialmente en el centro y sur del país, entre los 2.200 a 4.500 metros sobre el nivel del mar. Los pastos comunitarios son la principal fuente de alimentación animal en esta región. La crianza de llamas es comúnmente combinada con otras especies, como alpacas y ovejas. Una forma de evaluar
técnicamente a los sistemas ganaderos es mediante el cálculo de índices técnicos. Estos indicadores nos permiten sintetizar la información contenida en los registros de producción y normalizar los criterios de evaluación de la unidad de producción. En el presente estudio se utilizaron los siguientes parámetros técnicos: Capital promedio anual (CPA), natalidad bruta (%NB), natalidad real (%NR), mortalidad (%M), saca (%S), incremento bruto (%IB), incremento real (%IR) y eficiencia ganadera (%EG). En el presente estudio, se realizó la evaluación técnica del ganado mediante registros mensuales de llamas, alpacas y ovejas de la Cooperativa Comunal San Pedro de Racco, ubicada en la región de Pasco, Perú. El período de evaluación para llamas y alpacas fue del 2012 a 2015, mientras que para ovinos el período de evaluación fue de enero del 2014 a diciembre del 2015. Se concluyó que la Cooperativa Comunal San Pedro de Racco presentó valores de parámetros técnicos correspondientes a una empresa de buen nivel tecnológico, posiblemente debido al desarrollo de capacidades de recursos humanos en materia de manejo de animales y pastizales.

**Keywords:** llama, alpaca, sheep, performance evaluation

**Introduction**

In the Peruvian highlands livestock production is especially developed in the South and Center of the country, between altitudes ranging from 2,200 to 4,500 meters above sea level. Livestock is usually raised by peasant communities who have pastures as the main resource for feeding their animals (Flores, 1992).

In Peru llama rearing takes place in large areas of highland grasslands where the possibility of developing agriculture is almost non-existent. Llamas convert very efficiently poor pastures at high altitudes into high quality products such as meat (Quispe et al., 2008). In the Central Andes, the llama management system is extensive, based on the use of communally-owned grasslands and its main objective is the meat production. Many farmers keep mixed herds composed of llamas, alpacas, sheep, and cattle. The predominant llama type is the K'ara variety, which is mainly considered for its good qualities for meat production (Mendoza, 2013).

The majority of alpacas and llamas are raised by smallholder farmers in a low-input system. Disease control measures are non-existent in most cases, as well as the grassland management. The other 20% of the population belong to individual small and medium farmers and big enterprises. In communities, there is a tendency to possess many animals above the carrying capacity of pastures, which leads to overgrazing and consequent degradation of this resource (Coughenour, 1991). The absence of disease control and prevention measures results in high mortality rates as well as low growth and birth rates, all of which result in low levels of production and productivity. Additionally, farmers have many difficulties for the commercialization of products that depend on a chain of intermediaries. The result is a low level of income for families that has an impact on their standard of living. This is the most relegated and unprotected sector of camelid breeders but at the same time the one that has enormous potential for development due to the high number
of livestock it owns and a valuable local knowledge that farmers keep (FAO, 2005b).

Sheep rearing in Peru is mainly developed in the highlands, mostly under extensive grazing based on degraded natural pastures with low nutritive levels which is reflected in the production of wool, meat and hides (Aliaga, 2006). The sheep sector in the country supplies meat but most of it is informal, which accounts for 70 % of the national meat production, which is mostly for self-consumption (FAO, 2004). The effect of falling wool prices on the international market by adding the low-quality wool produced by small farmers and communities, as well as the presence of wool substitutes such as synthetic fibers and other natural fibers such as cotton has not allowed to increase the demand in the consumption of wool (Mesias, 2001). Sheep in Peru are certainly the savings box of the rural Andean farmer whose one of their customs is to sell them if there is an urgent need for money (Diaz, 2007).

One way to technically evaluate livestock systems, basically productivity, is by calculating technical parameters. These indicators allow us to synthesize the information contained in the production records and to standardize the evaluation criteria of the production unit (Pumayalla, 1980). The parameters show the efficiency with which the herd is being managed, as well as the success or failure of the management that is carried out (Cuadros, 1981). Gutiérrez (1993) points out that the following parameters should be used to perform a technical evaluation: Annual average capital (AAC), gross birth rate (%GBR), real birth rate (%RBR), mortality (%M), harvest (%H), gross increase (%GI), real increase (%RI) and livestock efficiency (%LE). The objective of the present study was to evaluate the performance of llama, alpaca and sheep herds of the Communal Cooperative San Pedro de Racco, located in Pasco Region, Peru.

## Materials and Methods

Performance evaluation of livestock was carried out using monthly records of llamas, alpacas and sheep of the Communal Cooperative San Pedro de Racco. This cooperative is located in the province of Cerro de Pasco in the Central Andes at an altitude of 4,000 m. The evaluation period for llamas and alpacas was from November 2012 to October 2015. In contrast, the sheep evaluation period was from January 2014 to December 2015. The technical parameters were calculated using the following formulas:

$$ \text{Annual average capital (AAC): } \left( \frac{\text{initial existence} + \sum \text{final existence}}{13} \right) $$

$$ \text{Gross birth rate (\%GBR): } \left( \frac{\text{N}^{\text{º} \text{births}}}{\text{N}^{\text{º} \text{matted females}}} \right) * 100 $$

$$ \text{Real birth rate (\%RBR): } \left( \frac{\text{N}^{\text{º} \text{births}}}{\text{AAC}} \right) * 100 $$

$$ \text{Mortality (\%M): } \left( \frac{\text{N}^{\text{º} \text{dead}}}{\text{AAC}} \right) * 100 $$
* Harvest (%S): $\left(\frac{\text{Nº sold} + \text{slaughter} + \text{donations}}{\text{AAC}}\right) * 100$

* Gross increase (%GI): $\left(\frac{\text{final existence} - \text{initial existence}}{\text{AAC}}\right) * 100$

* Real increase (%RI): $\left(\frac{\text{final existence} - \text{initial existence} - \text{sold}}{\text{AAC}}\right) * 100$

* Livestock efficiency (%LE): $\%RI + \%H$

The AAC is an indicator that expresses the monthly average number of animals that exists in the herd during the year. The % GBR is an indicator of reproductive efficiency in breeding, while the %RBR expresses the relation between the number of born and the AAC. The %M indicates the percentage ratio of the number of deaths among the AAC and the %H considers the number of animals sold, benefited and includes those destined for self-consumption and donations. The %GI expresses the increase of animals per year in the herd and the %RI expresses the increase of the population due exclusively to the technical management of the herd. Finally, %LE is an indicator of efficiency in the technical management of the herd.

## Results and Discussion

### Annual average capital

The results showed that llama and alpaca populations are reduced during the periods of 2012 to 2014, but increase in the years 2014 and 2015; this increase was also observed in the sheep population. This could be due to the grassland management strategies that the cooperative began to use at the end of the year 2013 (LEUP, 2013). It was found that the average AAC was 507.8, 3,086.2 and 4,683 animals, for llamas, alpacas and sheep, respectively.

### Gross and real birth rate

The average gross birth rates (%GBR) during the evaluated years were 46.21 %, 59.82 % and 66.21 % for llamas, alpacas and sheep, respectively (Figure 1); while the average real birth rates (%RBR) were 25.03 %, 29.43 % and 40.68 % (Figure 2). The birth rate obtained in the present study was lower than that reported by Pantoja et al. (2012) of 78.29 % for llamas in Pasco, however it is necessary to specify that in their study, they only evaluated the year 2011. A similar result was obtained by Novoa and Leyva (1996) who report a birth rate of 50 %. In the Turco region (Bolivia), Rodriguez and Quispe (2007) indicate birth rates of 61 % and 66 % for llamas and alpacas, respectively.
<figure><img src="image_21.png" /></figure>

**Figure 1:** Gross birth rates (%GBR) of llamas, alpacas and sheep from the Community Cooperative San Pedro de Racco from 2013-2015

<figure><img src="image_22.png" /></figure>

**Figure 2:** Real birth rates (%RBR) of llamas, alpacas and sheep from the Community Cooperative San Pedro de Racco from 2013-2015

On the other hand, the gross birth rate of alpacas reported in the present study was below the one calculated for alpacas Huacaya (89.34 %) and Suri (83.54 %) in the Research and Production Center “La Raya” in Cusco (Gallegos, 2013). One possible explanation for these differences might be the different weather conditions. “La Raya” is located in the wet puna, whereas the Community Cooperative San Pedro de Racco is located in the dry puna. Therefore, pasture productivity and