PAPERS PRESENTED AT THE EXPERT CONSULTATION (Contd.)

C. PRACTICAL ISSUES FOR THE CONSERVATION AND IMPROVEMENT OF PRIORITY BREEDS WITH GLOBAL REVIEWS OF SPECIES (Contd.)

A GLOBAL REVIEW OF THE GENETIC RESOURCES OF CAMELIDAE

C. Novoa¹ and T. Wilson²

1 Introduction

Alpacas (L. pacos Linnaeus, 1758) and llamas (L. glama Linnaeus, 1758), which have been domesticated for centuries, together with guanacos (L. guanicoe Muller, 1776) and vicuñas (L. vicugna, Molina, 1782 or V. vicugna [Molina] Miller, 1924) that still live in the wild in Latin America, constitute the four extant species of the genus llama. These species together with the two species of the genus Camelus, namely the dromedary or one-humped camel (C. dromedarius Linnaeus, 1758) and the bactrian camel or two-humped camel (C. bactrianus Linnaeus, 1758), form the Camelidae family (Blanc and Ennesser, 1989).

Because of the common ancestry and the fact that they have a similar karyotype and diploid chromosome number (2n=74), it is not surprising that many physiological phenomena in Lamoids and Cameloids have been shown to be similar. Reproduction is a good example of this similarity. Lamoids and Cameloids are induced ovulators. The four species of Lamoids are inter fertile and their offspring, produced by all possible matches of both pure and hybrid parents, are also fertile. The two species of Cameloids also cross readily and the crosses are fertile in both sexes.

Both the Cameloids and Lamoids have evolved specialized anatomical and physiological adaptations to the arid and semi arid zones which they inhabit, but only the former can go for long periods without water. The Lamoids have also adjusted to the high elevations of the Andes through the development of haemoglobin and tissue characteristics which facilitate the concentration and utilization of oxygen under conditions of chronic hypoxia.

This paper contains a brief description of Camelidae population, distribution and status and on these grounds, priority breeds for conservation and improvement are identified.

2 Population and distribution

2.1 Lamoids

Pre-hispanic llama and alpaca herding spread well beyond the limits of the Puna ecosystem, where they were first domesticated. The maximum extension of their distribution from Southern Colombia to Central Chile, occurred under the Inca empire, but with the advent of Spanish rule and the introduction of European farm animals, camelid production and breeding control began a decline which has never been reversed. Estimates of the current population of Latin American camelids are shown in table 1.

¹ Instituto Veterinario de Investigación Tropical de Alturas, Lima, Peru.

² FAO/UNDP Regional Project RAF/88/100, Banjul, The Gambia.

Table 1. Estimated population of Latin American camelids (x 1,000)

Sources: Cajal 1983; Cajal et al 1981; Franklin 1982, 1987; Hofman et al 1983; Novoa and Wheeler 1984; Torres 1985a and b.

Llamas are found from Colombia to Chile and Argentina, with the zone of greatest concentration located between 11 degrees and 21 degrees south, whereas the distribution of alpacas extends from Cajamarca and Ancash in Northern Perú (a recent reintroduction) to lake Poopo in Bolivia with small numbers in Northern Chile and Northwestern Argentina. However, the foremost alpaca producing area is within a 50 to 100 km. radius of lake Titicaca.

Llamas and alpacas are usually kept at elevations from 2300 to 4000 and from 4000 to 4800 m., respectively. Although local llama and alpaca distribution is determined by altitudinal, topographical, and vegetational conditions and alpaca tend to occur at higher elevations, the two species can be found in the same area because of herding by man. Llama makes highly efficient use of the bunch grasses of the Puna which typically contain 5% protein, while the alpaca selects the shorter grasses (San Martín 1987). This dietary difference permits the two species to co-exist in the same ecosystem, and is reflected in incisor morphology. Llama incisors are spatulate in form and have an enamel covered crown which when worn has two cutting surfaces, while alpaca incisors have enamel only on the labial surface and, hence only one cutting surface when worn. Llamas prefer dryer forage and actually reduce food intake during the rainy season, whereas alpacas are found most frequently in wet boggy areas (San Martín 1987).

The guanaco is the widest ranging and occupies the most diverse habitat types. It is found both in shrublands and warm and cold grasslands at elevations from sea-level up to 4250 m. (Franklin 1982) or 4600 m. in the Andes. Its northernmost distribution is at present located at about 8 degrees south in the department of La Libertad, Perú. From there its range extends south along the Andean Cordillera to Navarino Island in Tierra del Fuego, east across Patagonia and as far north as the sierras of Curamalal and la Ventana in the province of Buenos Aires, Argentina. In contrast to the guanaco, the vicuña is found only at the highest elevations of the Andes between 4200–4800 m. above sea level (Franklin 1982). The northernmost distribution of the vicuña, both past and present, is 9 degrees 30 minutes south in the department of Ancash, Perú. The southern most limit is presently 29 degrees 0 minutes south in the province of Atacama, Chile.

2.2 Cameloids

In 1990 there were estimated 19 million Camels in the world, of which 93% were one-humped camels. Their distribution is shown in table 2. The one-humped camel is found in almost all the arid and semi arid regions of the old world. They are vital to the economy of northern Africa, the middle East and central Asia. In general, numbers of one-humped camel have increased over the last decade. Where numbers have declined it has been for two different reasons: in countries in which oil is the major commodity and where nomadism is no longer the major way of life and, the severe droughts of the 1970s and 1980s (Wilson 1991). The two-humped camel is cold resistant as shown by its presence in northern China, Mongolia and the southern part of the former Soviet Union (Turkmenistan, Siberia). It appears that numbers of two-humped camels have declined slowly in China and Mongolia over the last decade.

3 Description and present status of existing camelidae

3.1 Lamoids

Latin American Camelids have not been adequately evaluated and in some instances, are threatened with extinction even before they are properly described. Based on available information an attempt will be made to briefly describe both domestic and wild Camelids indicating their current status.

3.1.1 Domestic species

Llamas have been selectively bred for use as pack animals and meat producers. They are larger than their guanaco ancestor (Wheeler 1984 a and b; Wing 1977, 1986). Two different breeds are known, although it is likely that more exist. The majority (70–80%) of Andean llamas are of the Q'ara or non woolly breed, which is characterized by its nude face and limited fibre growth. Less common is the Chaku or woolly llama, which as its name implies, produces more fibre and typically has wool growing on its fore head and its ears. Colours vary from white to black and brown but is not generally uniform across the body. Intermediate phenotypes between Q'ara and Chaku, although rare, have also been recognized. Table 3, shows comparative data on llama fibre diameter.

The Andean llama has never been selected for fibre production and like the guanaco, its fleece contains up to 20% kemp (Carpio y Solari 1982) and is woven into saddle-bags and made into rope. Trade in llama wool has not developed due, in part to its coarseness and colouration irregularities.

Table 2 - Part 1. Numbers of one-humped camels by region and country (x 1000). Adapted from FAO Production Yearbooks.

Table 2 - Part 2. Numbers of two-humped camels by region and country (x 1000). Adapted from FAO Production Yearbooks.

³ Although some camels on the former USSR are one-humped, the majority are two-humped.

In contrast to llamas, alpacas have been specially bred as wool producers and two breeds are distinguished on the basis of their fibre characteristics. Approximately 90% of all alpacas exhibit the shorter, crimped fibres of the Huacaya breed, while 10% (though numbers are rapidly declining) have the long straight fibres of the Suri breed. In appearance the Huacaya is reminiscent of Corriedale, and Suri of Lincoln sheep. Animals with intermediate type wool exist, but are rare. The genetic factors which control these traits are not known; crosses Huacaya × Huacaya produce a certain percentage of Suri offspring and crosses of Suri × Suri produce some Huacaya offspring. Results in table 4 would indicate that Huacaya is recessive, however more data is needed to confirm these preliminary results.

Table 3. Fibre diameter (micron) in adult llamas by type and sex

Sources: Vidal (1967).

Alpaca colour varies from white to black and brown, including all the intermediate shades and is fairly uniform across the body. Alpaca wool has a high commercial value because it contains little kemp, has a low felting quality, is very fine (see table 5) and can be woven into lightweight, soft and lustrous fabric.

Table 4. Huacaya and Suri Crosses

Sources: Velasco (1980).

The demand for white wool greatly reduced colour variation in the Andean populations in recent years; however, local artisan production is creating more demand for other colours.

The exact mechanism of inheritance of colour pattern is unknown. Alpaca crosses between white individuals and white crossbreds, mottle or other colours produce a large proportion of offspring with white coat (Condorena 1983). On the other hand data from Velasco (1980) indicate that uniform colour (without white hairs) is dominant over mottle (with spots of white hairs); additionally brown appears to be dominant over black.

Table 5. Camelid fleece characteristics.

Sources: Sumar, 1980; Bustinza and Belon 1982; Villarroel, 1963; Franklin, 1982.

Although llamas and alpacas are often herded together in Andean communities, care is taken to prevent cross-breeding. When this does occurs, the resultant hybrid or Wari is considered to be an inferior product. Despite the fact that its wool it is not as fine as alpaca wool, and though it may be larger than an alpaca, it is not of sufficient size for use as a pack animal and it produces less meat than a llama. Wari's are, however, fully fertile and hence tend to be rapidly eliminated from the herds.

The domestic camelids are well adapted to the rigors of life in the high Andes. In addition to physiological adaptations which offset the conditions of chronic hypoxia, their stomach is adapted for greater digestive efficiency than the advanced ruminant stomach (Vallenas et al 1971). Llamas and alpacas are principally raised on marginal lands located at, and above, the upper elevational limits of agricultural production. Their ability to convert the high cellulose pasture plants into a useful source of stored protein extends productivity into areas where crops cannot be grown and provides the most reliable human food resource in the high Andes (Thomas 1973; Novoa and Wheeler 1984).

In addition to meat for immediate consumption, or storage as sun-dried charqui, the herds provide other goods and services.

The llama is utilized primarily as a pack animal. It can transport loads weighing 25– 30 kg. over distances of 15–20 km. daily (Flores Ochoa 1977), and is employed in both local and long-distance inter-Andean trade for obtaining goods and commodities which are not produced in the Puna.

The alpaca, on the other hand, is primarily a wool bearer whose fibre is utilized for the production of fine quality cloth. The skins, sinews and bones of both animals provide leather products, thongs and weaving tools. Dung is utilized as a primary source of fuel in the treeless tundra environment of the high Andes. It is also essential as a fertilizer for effective potato production in the poor soils of this zone.

At present, all llamas and 90% of all alpacas in the Andes are under the control of traditional pastoralists. These small herd holders maintain flocks of 30–500 animals on communal grazing lands. The remaining alpacas are kept in large herds which belong to rural cooperatives in Perú. Llamas and alpacas have survived within the framework of traditional, non-European socio-economic organization because they are an essential element of Andean culture. Breeding and herd management procedures are decided by traditional techniques which are not always efficient, but it is in this context that the majority of present day llama and alpaca occurs.

During the last decade llama population remained relatively stable with approximately 3.8 million in the Andean Countries. However transport by truck and rail is replacing the llama and therefore there is trend for llama numbers to decline. In relation to alpacas, 89% of the Andean population is located in Perú and 10% in Bolivia. During the last 25 years the alpaca population in Perú has been decreasing, from 3.3 million in 1967 to 2.8 (1971), 2.5 (1976) and 2.4 (1980). This reduction was partly due to the agrarian reform of the 1970's.

3.1.2 Wild species

The guanaco is the largest of the indigenous wild artiodactyla in Latin America and the ancestor of the domestic llama. Four geographic subspecies have been described. The first, Lama guanicoe guanicoe Muller, 1776 is found in Patagonia, Tierra del Fuego and Argentina south of 35 degrees latitude. The second subspecies L.g.huanacus Molina, 1782 is said to be restricted to Chile, while the third L.g.cacsilensis Lonnberg, 1913 inhabits the high Andes of Perú, Bolivia and north eastern Chile. The fourth, L.g.voglii Krumbiegel, 1944 is restricted to the eastern slope of the Andes between approximately 21 and 32 degrees south latitude in Argentina. Today there is a trend to only recognize L.g. guanicoe and L.g. cacsilensis (Dennler de la Tour, 1954; Torres 1985 a); however other subspecies might exist which are still unknown.

All subspecies exhibit similar dark brown to beige colouration, white under parts and grey to black faces. Comparative biological data on camelids including guanaco are shown in Tables 5 and 6.

Table 6. Comparative biological data on Latin American camelids

Sources: Sumar (1980); Bustinza and Belon (1982); Calderón y Fernandez Baca (1972); Franklin (1982).

Guanacos live in both migratory and sedentary groups, (Franklin 1982). They are easily hunted and in fact numbers have been drastically reduced in most of their range due to excessive hunting. Thus, guanacos in Perú were declared endangered by the Peruvian Government in 1971; later this status was changed to vulnerable by the World Conservation Union (IUCN) in 1974.

Today, there are 14 national reserves in Argentina, 4 in Chile and 3 in Perú; however they remain unprotected in Bolivia.

Regarding Vicuñas, two geographic subspecies have been described. The first Vicugna vicugna vicugna Molina, 1982, found south of 18 degrees, is larger and lighter in colour than the second, V. mensalis Thomas, 1917. This last one has been regarded as the alpaca ancestor (Wheeler 1984 a and b). Both subspecies are characterized by their short, extremely fine cinnamon to light tan coloured fibre growth, white underparts and gracile form, but only V.v. mensalis exhibits a tuft of long white hairs of the chest.

In contrast to the declining population of guanaco, numbers of vicuña are increasing. From being a species threatened with extinction in 1969 it passed to vulnerable in 1972. This change has been the result of preservation and population recovery projects carried out by the Andean countries during the last 20 years. Efforts for vicuña conservation started in Perú in 1968 establishing the Pampa Galeras National Reserve. Vicuñas are fully protected in Bolivia, Chile and Argentina.

3.2 Cameloids

In addition to the obvious difference, the two-humped variety differ from the one-humped in being woollier, shorter in the leg and darker in colour. It is also adapted to the low winter temperature of Central Asia while the one-humped type is the typical animal of the deserts of north Africa and the middle East.

3.2.1 Types

In most areas camels are multi-purpose animals, the females being used primarily as milk producers, the males being used for transport or draft and both sexes provide meat as a tertiary product. The hair is used only by a few groups of owners.

One-humped camels may be divided, according to habitat, into mountain and lowland types (Table 7). The former is small, compact and coarse-boned. The sole of foot is very hard; a long fur may develop in winter. It can be subdivided into pack or riding subtypes. The lowland camel is larger and the foot tends to be softer, it is subdivided into riverine and desert types.

Table 7. Principal physical characters of lowland and mountain camels

Sources: Wilson (1984).

Alternately the one-humped camel may be divided into baggage and riding types and the former into hill and plains sub-types.

There have been attempts to categorize camels as beef, dairy, dual purpose and racing. However there appears little justification for this classification at present. Racing camels do not constitute separate breeds but are selected from within existing populations after they have shown a particular aptitude for speed. They are not reared primarily as meat producers.

3.2.2 Breeds

There has been little attempt to assign quantitative production parameters to the breed description. Recently, an attempt to using a more quantitative approach has been made. It uses six morphological and biological characteristics of habitat, function and geographical distribution to describe the 48 main breeds of the one-humped camel. The classification assigns camels to nine regions and sub-regions in three main groups and eight (or nine) sub-groups. This classification may be found in Blanc and Ennesser (1989). Physical size based on a series of linear measurements was used in this analysis as the main means of identifying breeds. However, it is doubtful, if the majority of workers interested in breeds would limit their number to 48 main ones or even limit themselves to the classifying variables employed.

In Somalia, a recent typology of three breeds has included ease of milking and rapidity of weight gain as parameters in the breed description (table 8).

Table 8. Characteristics of three Somali camel breeds.

Sources: Mohamed Ali Husein (1987).

Other recent attempts to classify breeds are shown in table 9. In the USSR all one-humped camels are of the Arvana breed (Turkmen=purebred). This is described as a dairy, transport and riding type whose qualities have been achieved through a long period of selection.

Table 9. Some camel breeds of arab Africa.

Source: Wardeh et al (1991).

3.2.3 Hybrids

The most usual hybrid is the bactrian x dromedary. The F1 products show heterosis in respect of body size, hardiness, endurance and longevity. Some bactrian characters, such as the hairy beard and legs are retained, and the single hump is longer and not as well developed as in the dromedary. This cross is a strong draft animal, whose wool yield tends towards that of the bactrian. The milk yield and the butterfat content of the hybrid are intermediate between the two parents.

4 Priority breeds for conservation and improvement

Given the situation outlined above, the following breeds of Lamoids have been identified for immediate development with criteria for their choice.

4.1 Huacaya

• Breed represents 90% of the world's Alpaca population.
• It is well adapted to the rigors of life of the high Andes.
• In addition to high quality fibre, they produce other valuable goods for industrial and artisan production.
• They are vital for the Andean economy especially for small farmers since 90% of all Huacayas are under their control.
• The breed is not efficiently utilized, its improvement can greatly benefit the people living in the High Andes. Recent experience, although limited, indicates that they can be reared in areas beyond the Andes and their capacity to utilize marginal pasture lands makes them an important potential resource on a world-wide scale.

4.2 Suri

• About 100% of the world's population of this breed is located in the southern Peruvian Altiplano, which unfortunately has declined drastically over the past 20 years.
• Suri alpaca once represented over 25% of the total alpaca population in Perú but represents only 5% today. In numbers of animals this is a decline from 700,00 down to just 130,000. Unless measures are taken to preserve and protect this germplasm now, suri alpaca could disappear by the year 2000.
• One primary cause relates to producers who unknowingly select against Suri through indiscriminate cross breeding with the Huacaya alpaca.

4.3 Qara

• This breed represents about 70–80% of the world's llama population.
• They have been selectively bred for use as pack animals and meat producers. However transport by truck and rail is replacing the llama and therefore there is a trend for llama numbers to decline.
• At present all llamas are under the control of small farmers. Their grazing habits, mostly on bunch grasses, and their resistance to hydrologic stress make them an important resource for arid zones and mixed grazing with sheep and/or alpacas.

While important progress has been made at research Centers in Perú, Bolivia and Argentina, much remains to be done before the full productive potential of alpacas and llamas can be realized. Of particular value are efforts to improve and standardize the production qualities of both Huacaya and Suri fibre and Qara meat and development of the necessary technology for implementing this knowledge.

To conserve and improve breeds, governmental physical facilities and flocks of llamas have been built in Argentina (Abrapampa) and Bolivia (Patacamaya and Turco) and of alpacas Huacaya in Perú (La Raya). However very little genetic progress have been made; inbreeding and lack of adequately defined breeding objectives are current problems affecting the vast majority of flocks especially those of the small farmers. Knowledge of the relevant genetic parameters is scarce. Conservation and development programmes of these breeds should aim at supporting the existing activities and place emphasis on the analysis of records and its use for selection.

As to the Suri alpaca, support should be given to assure its conservation, to develop and maintain a breeding flock, to conduct inter-disciplinary research and extend information to educate producers in the importance of maintaining the breed.

Regarding camels, it is not appropriate to make suggestions about which breeds should be given support for immediate development. Given the limited information that has been available, some injustices could be done. It is hoped that in the future, through the ongoing and new attempts to describe breeds objectively, breed selection for development programmes will be easier.

REFERENCES

Blanc, C.P. and Y. Ennesser, 1989. Approche zoogeographique de la différenciation infraspécifique chez le dromedaire Camelus dromedarius Linné, 1776 (Mammalia: Camelidae) Rev. Elev. Med. Vet. Pays Trop. 42: 573-57.

Bustinza, V. and J. Belon. 1982. Produccíon de Vicuñas de Kala Kala. Allpaká, 21–30.

Cajal, J.L. 1983. Situación del Guanaco y Estrategia de Conservación de los Camélidos en la Argentina. Buenos Aires, Subsecretaria de Ciencia y Tecnología.

Cajal, J.L., A.A Recer, J.C. Pujalte. 1981. La Reserva Provincial de San Guillermo y sus Asociaciones Ambientales. Buenos Aires, Programa de Recursos Naturales Renovables.

Calderon, W. and S. Fernandez Baca. 1972. Peso Vivo y Rendimiento de canal en la alpaca. Rev. Inv. Pec. (IVITA) 1:5–9.

Carpio. M. and Z. Solari. 1982. Diámetro de la Fibra en el Vellon de la Vicuña. Informes de trabajos de Investigación en Vicuñas 1:54–102 Lima, UNALM.

Condorena, A.C. 1983. Concepto del Sistema Estabilizado como Teoría de Organización y Producción en la Crianza de Alpacas, ed. N. Condorena. Cusco, pp.219.

Dennlen de la Tour, G. 1954. The Guanaco. Oryx 2:273–279.

FAO. Production Year Books, FAO, Rome, Italy.

Flores Ochoa, J. 1977. Pastores de Alpacas de los Andes. In: Pastores de Puna Uywamichig Punarunakuna, (J.A. Flores Ochoa, ed.) pp. 15–22, Instituto de Estudios Andinos, Lima.

Franklin, W.L. 1982 Biology, Ecology and Relationship to man of the South American Camelids. In: Mammalian Biology in South America Symposium (M.A. Mares and H.H. Genways eds.) pp. 457–489. University of Pittsburgh, Pittsburgh.

Franklin, W.L. 1987. My Two Decades with American Camels. International Wildlife Sept-Oct: 34–43.

Hofman, R.K., K.C. Otte., C.F. Ponce del Prado y M.A. Ríos 1983. El Manejo de la Vicuña Silvestre. 2 Tomos Eschborn, GTZ.

Mohamed Ali Husein, 1987. Emic Notes on Camel breeds in Somalia (Camel Forum Working Paper N^o 17) Somali Academy of Sciences and Arts Mogadishu, Somalia.

Novoa, C. and J. Wheeler. 1984. Llamas and Alpacas In: Evolution of domesticated Animals (I.L. Mason ed.) pp. 116–128, Longman London.

San Martín F. 1987. Comparative Forage Selectivity and Nutrition of South American Camelids and Sheep. Ann Arbor, Univ. Microfilms International.

Sumar J. 1980. La Llama, Recurso Genético de los Andes. Anales de la III Reunión Cientifica Anual. Asoc. Peruana Prod. Animal. Lima.

Thomas, R.B. 1973. Human adaptation to a High Andean Energy System. Univ. Park, occasional Papers N^o7, Department of Anthropology, Pennsylvania State University.

Torres, H. 1985a. Guanaco, Distribución y Conservación del Guanaco. UICN/CSE Grupo Especialista en Camelidos Sudamericanos. Informe Especial N^o2 Cambridge, Cambridge University Press.

Torres, H. 1985b. Conservación de la Vicuña en Chile. Arica, Corporación Nacional Forestal.

Velasco, J. 1980. Mejoramiento Genético de Alpacas. Anales III Reunión Científica Anual. Soc. Peruana de Prod. Animal. Lima.

Vidal, O. 1967. La Crianza de la Llama y algunas caracteristicas de su fibra. Tesis Ing. Zootecnista, Univ. Nac. Agraria La Molina. Lima, Perú.

Villarroel, J. 1963. Un estudio de la fibra de alpaca. Anales Científicos, UNA La Molina, Perú Vol I.

Vallenas, A.; J.F. Cummings and J.F. Munnell. 1971. A Gross study of the Compartmentalized stomach of two New-World Camelids, the Llama and Guanaco. Journal of Morphology 134:399–424.

Wardeh, M.F.; Zaied, A.A. and H.S. Horier, 1991. Camel breed types in Arab Africa in: Wardeh, M.F., Wilson R.T. and Zaied, A.A. (eds), Proceedings of the International Conference on Camel production and improvement, 10–13 December 1991. Tobruk, Libya. Arab Center for the Study of Dry Zones and Arid Lands; Damascus Syria pp. 78–86.

Wheeler, J.C. 1984a. On the origen and early development of Camelid Pastoralism in the Andes. En: J. Clutton-Brock y C. Grigson, Compiladores, Animals and Archaeology, Vol 3, early Herders and their flocks pp.395–410. Oxford, Bar International Series 202.

Wheeler, J.C. 1984b. La domesticaión de la alpaca (Lama pacos L.) y la Llama (Lama glama L.) y el desarrollo temprano de la ganadería autóctona en los Andes centrales. Boletín de Lima 36: 74–84.

Wing, E.S. 1986. Domestication of Andean Mammals. En: F. Vuilleumier y M. Monasterio, Compiladores, High Altitude Biogeography, pp. 246–264. Oxford. Oxford University Press.

Wing, E.S. 1977. Animal Domestication in the Andes. En: C.A. Reed, Compilador, Origins of Agriculture pp. 387–859. The Hague, Mouton.

Wilson, R.T. 1984. The Camel, Longman. London UK.

Wilson, R.T. 1991. A critical analysis of the research literature on the one-humped camel. In: Proceedings of the First Research Coordination Meetings of the Coordinated Research Programme of the Joint FAO/IAEA Division of Nuclear Techniques in Food and Agriculture. Institut Agronomique et Veterinaire Hassan II BP 6202 Rabat-Instituts Maroc.

A GLOBAL REVIEW OF THE GENETIC RESOURCES OF POULTRY

R.D. Crawford¹

1 Introduction

Status of the world's poultry genetic resources differs in several aspects from that of livestock. Indigenous poultry stocks contributing to food supplies in developing countries are only poorly described in the world literature. Chickens are best known as a food source. But there is only a sparse literature about other species such as turkeys, guinea fowl, domestic ducks, muscovy ducks and geese which also make a major contribution to food supplies. Accordingly, it is premature and perhaps impossible to recommend conservation and development of particular indigenous breeds since for the most part breeds have not been identified. Instead recommendations must be based on a broader view stressing particular poultry species and types.

2 Global Situation of Poultry Species and Types²

2.1 Poultry species.

Nomenclature for domesticated birds can be confusing. The most serious confusion surrounds the term ‘poultry’. In some countries it is used as a general term to include all domesticated avian species. In other countries it refers only to chickens. And in some countries it refers to all except waterfowl species. In this report the term ‘poultry’ will be used in the general sense to include all domesticated avian species.

¹ Department of Animal and Poultry Sciences, University of Saskatchewan, Saskatoon, Saskatchewan, S7N OWO, Canada.

² References for material quoted in this paper are Crawford, 1990 and National Research Council, 1991.

There are six poultry species that are a significant food resource in developing countries:

2.1.1 Chickens

Chickens are ubiquitous. They surpass all other domestic animal species as a source of protein in the human diet. There are three general types of food-producing chickens:

1. industrial stocks: bred by multinational corporations for mass production of eggs and meat. The production birds are terminal crosses. They monopolize urban cash economy markets in all developed countries, and increasingly in developing countries.
2. middle-level stocks: usually dual purpose (eggs and meat) stocks that have reasonably good production performance under reasonably good management conditions. They have disappeared from most developed countries.
3. indigenous stocks: the local unimproved poultry of developing countries. Production performance is usually poor, but they are widely held to have evolved adaptation to local environments. Their importance is in subsistence farming and village food production outside of a cash economy.

2.1.2 Turkeys

Turkeys as an indigenous form are ubiquitous in Latin America, but they are generally absent elsewhere. Industrial turkeys monopolize meat production in developed countries. Middle-level turkeys have been replaced by industrial stocks and only a few remain anywhere.

2.1.3 Guinea Fowl

Guinea fowl have major importance as a food source throughout developing countries of Africa; some of the birds are domestic, some are feral, and some are wild. They have only very minor importance elsewhere, although they are gaining favor in Europe as a specialty food product.

2.1.4 Domestic Ducks

Domestic ducks (Anas platyrhynchos) have immense importance as a food source in Asia, especially in the southeast. Eggs have more importance than meat. Some of the stocks are indigenous in type. Others are middle-level and are used locally by specialist producers. The species has only minor importance in Africa, Latin America, and the Near East. Industrial meat stocks are used in developed countries, but there are no major industrial egg production stocks.

2.1.5 Muscovy Ducks

Muscovy ducks (Cairina moschata) are the least known and understood of all poultry species. They can produce a sterile hybrid, the mulard, with domestic ducks. Muscovies were domesticated in Latin America where they remain ubiquitous. They are also prevalent in Africa and Asia. They are important in subsistence farming and village economies. Industrialization has begun in Europe.

2.1.6 Geese

Geese are a domestic bird of temperate climates. Except for China and southeast Asia, they are not characteristic of developing countries. Goose production there and in developed countries rests on indigenous and middle-level stocks. Industrial production has not yet begun.

3 World inventory of poultry genetic resources.

1. Knowledge about existing poultry genetic resources of the world is only fragmentary. It lags far behind that for livestock species, especially cattle and sheep. Breeders of industrial poultry will not reveal specific information about their foundation and grandparent lines, although broad generalities about their procedures are general knowledge. Middle-level poultry stocks of developed countries have been recorded to a variable extent, especially in Europe and North America, but those of developing countries have not. The biggest lack is that of information on indigenous poultry stocks of developing countries. There is essentially no information at all in the English literature about poultry genetic resources of Latin America, and very little concerning stocks of the Near East. Information from Africa and Asia has been increased through recent projects sponsored by FAO and others, particularly concerning indigenous chickens of India and domestic ducks of southeast Asia.

2. The European Association of Animal Production (EAAP)/FAO Global Animal Genetic Data Bank thus far has not included information on poultry resources. Descriptors suitable for encoding data on poultry have now been prepared for the major poultry species. Data recording should begin soon, if it has not already done so.

3. FAO should be reminded that in 1989 the World's Poultry Science Association (WPSA) presented a resolution to the FAO Expert Consultation on Animal Genetic Resources; the resolution was accepted by the consultation and was included in its proceedings (FAO, 1990). The resolution asked that FAO continue and increase its work in conservation and management of poultry genetic resources globally. WPSA offered to help FAO in any way possible in conducting this work. WPSA represents a powerful resource of professional people from nearly all countries. Total membership exceeds 5000. It is waiting for an initiative from FAO. For instance, it could provide priceless assistance in preparing a world inventory of poultry genetic resources, whereby FAO might provide leadership and coordination, and WPSA could provide field staff and information.

4 Distribution, utilization, vulnerability of poultry species.

4.1 Chickens

Chickens are ubiquitous. Globally they contribute more animal protein (eggs and meat) to the human diet than do any other domestic animal species. Indigenous chicken stocks are an important feature of village and subsistence farmers in virtually all developing countries. Their food production performance is usually poor, partly because of inadequate nutrition and management, disease exposure, and lack of selection for production traits. However it is widely presumed (without much solid evidence) that the indigenous stocks are genetically adapted for survival and reproduction in the local environment, and hence they are to be preferred. There is also increasing anecdotal information that eggs and meat from indigenous chickens are much preferred by consumers to those from industrial stocks.

Replacement and substitution of indigenous chicken stocks by both middle-level and industrial stocks from developed countries has been in progress for many years. At the village and subsistence farm level substitution has been mostly by middle-level dual-purpose stocks. The general result has been an increase in egg and meat production, but concurrently also a greatly increased mortality and morbidity due to disease and management shortcomings. It is a reasonable guess that in countries with a long history of improvement programmes, local indigenous stocks will have had considerable genetic influence from middle-level stocks. Where a commercial industry has developed near urban centres, the stocks in use are industrial from multinational corporations. It is unlikely that these are being used to replace indigenous stocks in rural areas, but they replace indigenous birds totally where there is a cash economy poultry industry near urban centres.

There is need for conservation action to ensure continuance and improvement of indigenous stocks in rural areas. Success in improving production performance through genetic selection has been demonstrated many times already, in Egypt, India, Iran, and Malaysia.

4.2 Turkeys

Turkeys have restricted distribution. They are ubiquitous in Latin America, where they were domesticated. But they are generally absent from other developing areas of the world. They play an important role in food production at village and subsistence farm level throughout Latin America. Curiously, their role is that of providing cash income rather than that of providing food for their owners. Production performance is generally poor because of inadequate nutrition and management, and because they probably have had little selection for production performance. Mortality and morbidity are high.

It should be clearly recognized that the Latin American indigenous turkeys are very different in genetic origin from the turkeys utilized in developed countries (and in the commercial turkey industry within Latin America). Turkeys were domesticated in Mexico from Meleagris gallopavo gallopavo. They spread throughout Central and South America and have persisted as indigenous turkeys. Their plumage is mostly black. Meanwhile, those taken to Europe and subsequently to eastern North America hybridized with another wild subspecies, Meleagris gallopavo silvestris, to produce the bronze turkey, forerunner of all commercial turkeys in developed countries. Two types of commercial birds persist from the bronze hybrids. Industrial turkeys, all of them white, are bred by only a few multinational corporations; the birds have astounding meat production performance, but they have been bred beyond the limits of normal reproductive fitness; they can no longer mate naturally and must be bred by artificial insemination. A very few stocks of middle-level turkeys persist, some of them bronze and some of other colors. These have moderately good meat production performance while retaining natural mating ability.

There is no danger of industrial stocks replacing indigenous turkeys in rural areas, especially because the former reproduce only by A.I. But industrial production has probably already made major inroads into meat production near urban areas. Middle-level turkeys are being sold into Latin America, especially from Europe, and because of superior meat production they may be a threat to persistence of the indigenous birds.

4.3 Guinea Fowl

Guinea fowl are kept throughout the world, but only in developing countries of Africa do they have major importance as a food source. Both eggs and meat are utilized, and in some areas these products are preferred to those from chickens.

A problem peculiar to considerations of guinea fowl is that domestication status is only poorly understood. Some of the birds are domestic, but eggs and meat are also harvested from feral and wild populations. The relative proportions of each are not known. Literature on the species is scanty. Their production performance has been most studied in Nigeria.

Because guinea fowl are so little used throughout the world, there is no immediate threat of substituting indigenous stocks with improved stocks from elsewhere. However industrialization is underway in Europe, particularly in France, and it is beginning in other countries too. The needs presently are to define whether exploited stocks are domestic, feral, or wild; to identify and evaluate domestic stocks from various geographic areas, with a view to greater distribution and utilization of superior ones; and to attempt genetic improvement of stocks.

4.4 Domestic Duck

Domestic duck, Anas platyrhynchos, has immense importance as a food source in village economies of Asia, especially in southeast Asia. The species has only minor importance in Africa, Latin America, and the Near East. In some Asian countries only duck eggs are utilized; in other areas there is also strong demand for duck meat.

Domestic duck populations of Asia have been intensively studied in some countries but not in others. There appear to be two levels of production: small scale production by villagers and subsistence farmers using unimproved indigenous stocks; and large scale production of eggs and meat by specialists using the equivalent of western middle-level stocks that have undergone some genetic improvement through natural and artificial selection, and that receive careful and expert management.

Industrial production of ducks in developed countries is dominated by one major breeding company, and there are several smaller companies. Their main product is meat from the Pekin breed; they have lesser concern over egg producing stocks. The industrial stocks have entered developing countries and are claimed to be performing well. It remains to be seen whether they will persist and whether they will displace indigenous and local middle-level stocks. There is need for further inventory and assessment of indigenous and local middle-level stocks, and for selection improvement programmes.

4.5 Muscovy Duck

Muscovy duck, Cairina moschata, is the least known and understood of all poultry species. It was domesticated in Latin America where it remains ubiquitous. It is also found in all equatorial countries of Africa and Asia, and it is particularly prevalent in southeast Asia.

In Latin America and Africa, muscovy ducks are an important poultry species in subsistence farming and village economies. They are used for both egg and meat production, and fulfill a minor role in household insect control. They are successful in part because of their aggressive behavior and superior naturally reproducing abilities.

In Asia the species is also used in subsistence farming and village economies. But more importantly it is used for commercial production by specialists for quantity production of meat, either as a pure species, or as a hybrid with domestic ducks. Production of the sterile hybrid, or mulard, is particularly important in Taiwan and some other local areas.

Muscovy ducks have only recently been exploited in developed countries, particularly in France, where genetic selection has been successful in improving growth rate and meat production. Improved stocks will undoubtedly soon enter developing countries, especially in southeast Asia's specialized duck industry. Meanwhile there is need for inventory which exists nowhere, for assessment of local stocks, and for selection projects to increase food yield.

4.6 Geese

Geese are a domestic bird of temperate climates. Except for China and southeast Asia, they are not a characteristic species of developing countries. Domestic geese used in Asia resemble the Chinese and African breeds of western countries; they were domesticated from the swan goose, Anser cygnoides of eastern Asia. Western breeds were domesticated from the wild greylag, Anser anser; they do not perform well in hot climates.

The domestic geese of China and southeast Asia can be classed as indigenous. But some stocks have presumably undergone both natural and artificial selection for enhanced reproductive and growth performance; these stocks are held by local commercial producers, and could be regarded as middle-level. It should be noted that even in developed countries, breeding and production of geese have not been industrialized, although genetic base is becoming monotypic using mostly the Emden breed.

Thus far there is no indication that geese of China and southeast Asia are being replaced by stocks from elsewhere, probably because industrial exploitation has not yet begun. However there is need for inventory and assessment, and for selection projects to enhance production performance.

5 Summary of Situation by Geographic Area

5.1 Africa.

The principal poultry species contributing to food supplies in developing countries of Africa are chickens and guinea fowl. Chickens have received considerable assessment and development attention. Guinea fowl have received almost none.

Guinea fowl are a major source of eggs and a lesser source of meat throughout developing Africa, and especially in west Africa. There is a distinct consumer preference for guinea fowl products over those from chickens. Relative consumption of products from the two species is not known. Unfortunately, knowledge concerning the birds is very scanty. It is known that some of the stocks are domestic, some are feral, and there is harvesting of products from wild populations, but proportions of each are unknown. There is no inventory of any sort. Except for some published research from Nigeria on production performance, nutrition, and management, there is almost no research information in the English literature. There is little danger of replacement by improved stocks from elsewhere since industrialization of guinea fowl has only just begun in Europe. However because of the importance of guinea fowl as a food source, efforts should be directed toward increasing knowledge about them - inventory and assessment, including storing of information in the EAAP/FAO Global Animal Genetic Data Bank, and toward improvement of existing stocks to enhance their use.

Chickens are ubiquitous and a major contributor to food supplies in subsistence farming and village economies. Indigenous stocks are poorly known, but there is scattered literature from some countries which includes FAO-sponsored studies. Improvement programmes in Egypt and Nigeria have demonstrated progress from genetic selection. There is anecdotal information to indicate that products from indigenous chickens are much preferred to those from improved commercial chickens. Industrial stocks have replaced indigenous near urban centres to monopolize the cash economy sector, but they are unlikely to have much influence in rural areas. In contrast, middle-level stocks have had a long history of introduction to developing countries; they probably have had some influence on indigenous populations.

5.2 Asia

The principal poultry species contributing to food supplies in Asia are chickens, domestic ducks, and muscovy ducks; geese have minor importance in China and in southeast Asia. Chickens and domestic ducks have received a lot of development attention but muscovy ducks have been generally overlooked.

Chickens have immense importance as a food source. Indigenous chicken populations have had prolonged study, especially in India and in Malaysia. They have been better characterized than stocks in other continents. Their situation is similar to that of indigenous chickens elsewhere: industrial stocks have replaced all other types in commercial production units near urban centres, and products of industrial stocks prevail in cash economy markets; middle-level stocks continue to be used in upgrading schemes in rural areas, and they have undoubtedly altered some indigenous stocks permanently; other indigenous populations remain undisturbed. Continuing inventory, assessment, and improvement programmes for indigenous stocks should be encouraged.

Domestic ducks are the major poultry species in some areas of Asia, particularly in the southeast. Eggs are the principal food product, with meat having much less importance. Indigenous stocks have had intensive study as a result of projects and activities of FAO, SABRAO, national governments, and other agencies. Industrial stock from the world's only major primary breeder of meat ducks has entered developing countries; there are no major industrial breeders of egg stocks. Asia has had its own middle-level stocks for centuries, kept by specialists for egg production primarily; anecdotal information implies that their production performance is excellent; it was these stocks that founded the egg production duck stocks of the western world. There is need for continuing inventory and assessment, including storage of information in the EAAP/FAO Global Animal Genetic Data Bank. There is value in considering more extensive use of the best Asian middle-level stocks, and in improving the local indigenous stocks in subsistence and village economies.

Muscovy ducks are widely used in Asia both at the subsistence farming/village level and in the local poultry industry. The birds are kept as a pure species in villages, and are produced as purebreds and as hybrids by industry specialists. The preferred hybrid is from a muscovy male mated to a domestic duck female by hand-mating or artificial insemination; fertility and hatchability are low but demand and price are such that hybrids are generated in large numbers. Census data seldom distinguish between muscovy and domestic species; neither do descriptive writings. Accordingly there is no information on muscovy duck numbers, populations, performance, etc. This void should be corrected.

5.3 Latin America

There are three major food-producing species in Latin America - chickens, turkeys, and muscovy ducks. There is a huge and rapidly growing commercial poultry industry in Latin America using industrial stocks and the infrastructure of developed countries; this industry provides for the food needs of urban centres. Although poorly documented, there have undoubtedly been repeated importations of middle-level stocks from elsewhere which have diluted (and sometimes replaced) the indigenous poultry of rural areas and which will have contributed to commercial production in urban areas. Unfortunately, almost nothing is recorded in the English literature about indigenous birds of these three species - no inventories or descriptive data, no assessments, and no reports of selection projects. Almost all of the available genetic resources literature from Latin America pertains to cattle and sheep.

Indigenous turkeys are an unusual case. They are ubiquitous in Latin America in subsistence farming and village economies, but they are not primarily a food source for the owners; instead they provide a source of cash income from urban users. They are genetically unique and different from all other domestic turkeys. The indigenous birds are presumed to derive exclusively from the original domestication of Meleagris gallopavo gallopavo that occurred in Mexico. All other domestic turkeys are presumed to derive from hybridization of M.g. gallopavo with M.g.silvestris that occurred in eastern North America in the 18th and 19th centuries. The Latin American indigenous turkeys have had virtually no study. Their attributes, status, and performance relative to the gallopavo-silvestris hybrids are unknown. This void of information should be filled.

Muscovy ducks were domesticated in northern South America prior to the Spanish conquest. They have spread throughout Latin America as an indigenous poultry stock in subsistence farming and village economies. They provide a source of meat, and perhaps of eggs; they have had a historic role in household insect control. Hybrids with domestic ducks seem not to be important as a product. Muscovy ducks have not been commercialized in Latin America thus far, although that could happen soon using stocks and infrastructure developed in Europe. There has been almost no study of the species in its centre of origin. There is need for inventory, assessment, and improvement projects.

Chickens are ubiquitous in Latin America, as they are elsewhere. They have been there for centuries; there is ongoing academic debate on whether they were there before the Spanish conquest, and if so, where they came from. Indigenous stocks have been replaced by middle-level and industrial birds near urban areas. Middle-level stocks have undoubtedly diluted the indigenous stocks in rural areas. Meanwhile the indigenous stocks have had almost no study. This void should be corrected before there is further erosion.

5.4 Near East

As for Latin America, there seems to be very little published information on the poultry genetic resources of the Near East. It is presumed that the predominant poultry species is the chicken, with domestic ducks playing a much less important role. The status of chickens is probably the same as that in other developing areas - industrial stocks monopolizing urban food supplies, and middle-level stocks diluting indigenous stocks in rural areas. Indigenous stocks themselves are only poorly known. There are published studies from Iran, but recent political events may have altered the situation dramatically. Little information is available from other countries of the area. It is presumed that domestic ducks have secondary importance in appropriate areas of the Near East, but information is lacking. Because of the absence of information, little can be recommended except to stress the need for inventory and assessment surveys.

6 Priorities for FAO Work in Poultry Conservation and Management

6.1 Inventory and assessment of poultry genetic resources.

Because of the paucity of basic information on indigenous poultry genetic resources in all continents of the developing world, it is imperative that first priority be given to a world inventory and assessment, and assembling of that information in a data bank. Capability for data bank entry and storage is now available through the EAAP/FAO Global Animal Genetic Data Bank. The World's Poultry Science Association has offered to collaborate with FAO in any way possible; a logical collaboration would be that of assembling the needed body of data.

6.2 Specific stocks targeted for conservation and improvement.

Again because of paucity of basic information, it is not appropriate or even possible to identify specific breeds for attention. Instead it is urged that species and broad types, which have thus far been ignored, be targeted for study and development by FAO. The following priorized list is suggested:

6.2.1 Indigenous turkeys of Latin America

Inventory and assessment of local and regional resources throughout the area; conservation and selection of superior stocks for future use.

6.2.2 Guinea Fowl of Africa

Inventory and assessment of indigenous stocks to identify and distinguish them from feral and wild food sources; conservation and selection of the most likely stocks for future utilization.

6.2.3 Muscovy ducks of Latin America and Asia

Inventory and assessment of indigenous stocks, and of middle-level stocks used for purebred and hybrid (mulard) production in Asia; conservation and improvement of superior stocks for future utilization.

6.2.4 Domestic ducks of Asia

Continued assembling of information; conservation and improvement of superior stocks, both indigenous and middle-level, for future use.

6.2.5 Chickens

Inventory and assessment of indigenous stocks in Africa, Asia, and Latin America; conservation and improvement of superior stocks for future utilization.

7 References

Crawford, R.D., editor, 1990. Poultry Breeding and Genetics. Elsevier Science Publishers, Amsterdam and New York.

FAO, 1990. Animal genetic resources. A global programme for sustainable development. FAO Animal Production and Health Paper 80. Food and Agriculture Organization of the United Nations, Rome, Italy.

National Research Council. 1991. Microlivestock:Little-Known Small Animals with a Promising Economic Future. National Academy Press, Washington, D.C., U.S.A.