Piebald Camels Gabriele Volpato1, Maurizio Dioli2 and Antonello Di Nardo3*
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Volpato et al. Pastoralism: Research, Policy and Practice (2017) 7:3 Pastoralism: Research, Policy DOI 10.1186/s13570-017-0075-3 and Practice RESEARCH Open Access Piebald Camels Gabriele Volpato1, Maurizio Dioli2 and Antonello Di Nardo3* Abstract Animal breeds are the diverse outcome of the thousands-year-long process of livestock domestication. Many of these breeds are piebald, resulting from the artificial selection by pastoralists of animals bearing a genetic condition known as leucism, and selected for their productive, behavioural, or aesthetical traits. Piebald dromedary camels have not been studied or discussed before, and their same existence is often overlooked. Based on fieldwork in Western Sahara, direct observations across Northern and East Africa and the Middle East, and a literature review, we address the morphological and behavioural traits, geographical distribution, taxonomy, and material and cultural importance of piebald (painted) camels. They are a hundreds-year-old camel breed used for caravans, as mounts, and for aesthetical and cultural reasons across Sudan, Niger, Mali, Mauritania, Western Sahara, and Morocco. While they are increasingly bred out of a pastoral context for tourism and entertainment in the Canary Islands, mainland Europe, and the USA, in part of their original African range, piebald camels are under threat due to wars, droughts, and demise of pastoral livelihoods. More research is needed about these ‘beautiful and dignified’ animals. Keywords: Livestock breeds, Leucism, Paint dromedary camels, Pastoral nomads, Sahara painted, or patched of white and solid colour. Many of Have you ever seen a piebald mahri1 before?; these piebald breeds are the result of a selective breeding for a genetic condition known as leucism. Have you ever seen a camel with such grace, lightness, Animals’ colouring is the result of either presence or ab- and stature? sence of the pigment melanin in the skin, hair, and eyes. Have you ever seen anything more beautiful and Among the known conditions affecting melanin production dignified? and animal colouring are albinism, melanism, and leucism; (Al-Koni 2008) the latter resulting from defects in pigment cell differenti- ation and/or migration from the neural crest to the skin, hair, or feathers during development (Cieslak et al. 2011; Introduction Wilkins et al. 2014). In its most common form, leucism re- A breed is a group of domesticates that has specific charac- sults in irregular patches on the body surface expressing as teristics or traits artificially selected by man and transmitted white on an animal that otherwise has normal colouring through inheritance (Driscoll et al. 2009; Larson and Fuller and patterning (Rook et al. 1998; Cieslak et al. 2011); when 2014). Breeds are the diverse outcome of the thousands- this happens, the animal is ‘pied’ or ‘piebald’ (also ‘paint,’ year-long process of livestock domestication (Francis 2015). ‘spotted,or’ ‘speckled’).2 Some leucistic animals also exhibit They have been selected in accordance to productive, cul- coloration of the irises that matches the surrounding skin tural, and aesthetical traits and are often a key element of (blue eyes for pink skin, brown for dark). pastoral livelihoods and identities. Breed diversity is now- Leucism and piebald colorations occur sporadically in adays endangered by processes of livestock intensification the wild, as they usually reduce the individuals’ selective and cultural homogenization (Faye et al. 2004), and as such, fitness providing less colour protection from predators ’ its study is a fundamental step for breeds conservation. (Woolf 1991). But recessive conditions in domesticates Among the thousands of breeds from tens of domesticates have been propagated voluntarily or not by humans existing in the world, several are piebald, i.e. spotted, since the very beginning of the domestication process, and changes in size and colour are among the first ef- * Correspondence: [email protected] 3The Pirbright Institute, Pirbright, Woking, Surrey, UK fects (Epstein 1955; Trut et al. 2009; Ludwig et al. 2009). Full list of author information is available at the end of the article © The Author(s). 2017 Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. Volpato et al. Pastoralism: Research, Policy and Practice (2017) 7:3 Page 2 of 17 Voluntary propagation includes the selective breeding of camel domestication, the author states that ‘In the United specific phenotypes for their religious, ritual, social, or States, paint camels – white and brown, like pinto horses – subsistence/economic value. Involuntary propagation re- have been produced, but paint camels are virtually absent in lates to the genetic and physiological links existing be- Arabia and North Africa.’ Though it is true that there are no tween tameness (a goal of early domesticators) and painted camels in Arabia and East Africa, we contend that colour phenotypes (Wilkins et al. 2014; Cieslak et al. they indeed exist in North Africa since hundreds of years, 2011; Price 2002) and to the effects of genetic drift in and that the USA is just the latest country where they have domesticates’ isolated populations (Zeder 2012). There is been introduced after their diffusion westward from central- a general agreement that a wide variation of colour phe- eastern Sahara, where they probably originated. notypes were brought about by domestication, ‘but it is This paper departs from and contributes to the study of still largely unknown how color phenotypes were distrib- human-animal relationships and domestication from an uted in domesticated animals in earlier times, which historical and anthropological perspective (Bulliet 2005; roles they played for humans during these times and Francis 2015; Hurn 2012). Most research on camels re- when they first occurred’ (Cieslak et al. 2011). volves around their potential as milk and meat producers The piebald leucistic phenotype is known in a variety of and associated conditions (e.g. diseases) (Farah and Fischer domesticates, including Tobiano and Appaloosa horses 2004), whereas relatively less studies have been carried out (Brooks and Bailey 2005; Brooks et al. 2007), Belgian Blue on the productive and cultural links between pastoral pop- and Shorthorn cattle breeds (Seitz et al. 1999), goats, and ulations and camels. In general, camel breeds are not as sheep, as well as dogs, cats, and pigs (Cooper et al. 2005; differentiated and classified as in other livestock species Giuffra et al. 1999). Several piebald animals have evolved and their classifications are often derived from names of into valued breeds, where Holstein cows and Dalmatian ethnic groups or geographical breeding regions rather than dogs are among the most notable examples. based on phenotypic characters (Dioli 2016). Therefore, Some of the genetic mutations related to the piebald char- camel breed study could help their conservation and sup- acter bear pleiotropic effects such as deafness, developmen- port camel pastoralists (Kakar et al. 2011). Studies on the tal disorders of the eyes and night blindness, and cultural relevance of piebald livestock (e.g. cattle) have osteopetrosis (Stritzel et al. 2009; Bellone 2010; Wilkins et already proven to be useful in describing complex human- al. 2014). Because the development of the optical system is animal relationships particularly in regard to livestock highly dependent on the presence of melanin, depigmen- management (e.g. in genetic selection) and cultural identity tation has an effect on the development of the visual system (e.g. founding myths, cultural identity, and social values) (Grandin and Deesing 1998; Wilkins et al. 2014). Congenital (Coote 1994). In this paper, we address piebald camels’ stationary night blindness has been reported in a variety of physiological, genetic, and behavioural characteristics; their spotted and piebald animals (e.g. Appaloosa horses; Sand- geographical distribution; taxonomy; and material and cul- meyer et al. 2007). Animal breeders throughout the world tural roles among pastoral populations. Then, we discuss have since early times recognized that a lack of body and piebald camels’ origin and diffusion through Africa and out eye pigmentation may be accompanied by neurological de- of Africa, we address trends in piebald camels’ husbandry, fects. A relationship between depigmentation and congenital and we invite further research about and support to these deafness has been found, among other domesticates (Webb ‘beautiful and dignified’ animals. and Cullen 2010), in dogs (e.g. Dalmatian dogs with exten- sive white depigmented areas are most likely to be deaf; Study area Strain 1996) and in llamas and alpacas (Camelids), where it The area under study includes large parts of the Sahara is associated with pale blue eyes (Gauly et al. 2005). desert where camels are bred, with a focus on Western A great deal of research has been conducted on the genet- Sahara and the Sahrawi pastoralists (Figure. 1, Table 1). ics, behavioural, and productive aspects of piebald breeds The Sahara is bordered by the Atlantic Ocean on the among several species (e.g. horses, dogs). In spite of this, the western