Domestic quail (Coturnix japonica domestica), is there such farm animal?

Domestic quail (Coturnix japonica domestica) Around 10% of all table eggs number in the world come from quail and their meat represents about 0.2% of the global poultry production. The domestic quail population involved in production of meat and eggs represents around 11.8% of all productive birds, placing them second after laying hens. China, Spain, France, Italy, Brazil, the USA and Japan are global leading countries in quail farming. Contemporary domesticated Japanese quail, widely used in agricultural practice, originate from wild Japanese breeds (C. japonica). The wild Asian quail is similar to common quail (Coturnix coturnix) and until recently, was accepted as its subspecies (Coturnix coturnix japonica). In the scientific literature, several names for domestic quail are available, so readers can easily become confused as to which species is really meant - wild, domesticated or common quail. In order to avoid confusion of names and to emphasise changes resulting from domestication, the term 'domestic quail' and its corresponding Latin name Coturnix japonica domestica is more appropriate. Within the EU legal framework, domestic quail should be included in parallel to other domestic fowl species.

Introduction

According to the definition of Clutton-Brock (1987), a domestic animal is one that has been bred in captivity for purposes of economic profit to a human community that maintains complete mastery over its breeding, organisation of territory and food supply. Domestication is the process by which captive animals adapt to man and the environment he provides (Price, 1984). Semi-domestic animals are those for which humans have only partial control over breeding, mortality, space use and food supply, and that have not been greatly modified by artificial selection. Semi-domestication is a process of bringing an animal to a semi-domesticated state. To consider an animal as domesticated, it should possess a number of genetic, morphological and behavioural traits different from those of its wild ancestor (Mysterud, 2010). These changes are due to deliberate or accidental human intervention to satisfy their specific needs. These changes make domesticated animals more distant from their wild ancestors, so most of them are designated as subspecies of the wild species, and others may be considered separate species (e.g. cattle, sheep or yak).

In the scientific literature, wild or farmed quail are referred to by several names, which is often confusing for readers. Another problematic aspect is how quail are placed in the EU legal framework. There is confusion between separate Commission Regulations, some of which include the quail under the term 'poultry' and others do not. Worldwide, the farmed quail population is secondary only to chickens, which is indicative of their role in eggs and poultry meat production relative to other poultry.

The aim of the present paper was to examine the role of quail in livestock farming and propose an adequate terminology for use in the scientific literature and the current EU legal framework.

Quail taxonomy

Quail belong to two families: Odontophoridae (New World quail) and Phasianidae (including several genera representing Old World species). Both the common quail (Coturnix coturnix), and the Japanese quail (Coturnix japonica) belong to the genus Coturnix and until recently, were believed to be one species. Unlike wild Japanese quail, where no subspecies are existing, Coturnix coturnix has five subspecies: Eurasian subspecies (Coturnix coturnix coturnix, Linneaus, 1758), Azorean Quail (Coturnix coturnix conturbans), Abyssinian quail (Coturnix coturnix erlangeri), African subspecies (Coturnix coturnix africana, Temminck and Schlegel, 1849) and Cape Verde quail (Coturnix coturnix inopinata). The genus Coturnix comprises eight species, two of which are already extinct. All members of the genus are migratory birds, and often their species ranges overlap. This hinders the differentiation of some subspecies (Minvielle, 2009). A serious concern for maintaining genetic purity in wild quail is hybridisation between common and domestic quail by hunters and farmers in the wild (Deregnacourt, 2000; Deregnacourt et al., 2005; Barilani et al., 2005; Huisman, 2006; Charaza et al., 2010). Common x domestic quail hybrids are fertile, exhibit the same nesting and brooding behaviour as wild ones (Deregnacourt et al., 2002), but are substantially unstable compared to the wild form, therefore their nests can be exposed to a greater risk of predator attacks (Puigcerver et al., 2014; Capdevila et al., 2016) and they may lose their migration behaviour (Deregnacourt et al., 2005). A similar crossing has been reported between domestic and wild Japanese quail (Chang et al., 2009), which is not a hybridisation per se, as the species is the same, but results in changes in behavioural and exterior traits compared to the wild Japanese quail population.

Quail domestication

The modern domesticated Japanese quail which are widely used in animal husbandry practice originate from the wild Japanese quail (Coturnix japonica, Temminck and Schlegel, 1849), as confirmed by modern genetic studies (Chang et al., 2007). Rearing wild birds in captivity and their domestication are different things. The analysis of available literature showed that wild quail have been reared as fighting, song and decorative birds from ancient times in East and Southeast Asia, while the real domestication had occurred in Japan within a very short period by the end of the 19th and beginning of the 20th century.

According to Mizutani (2003) first written testimonies about rearing Japanese quail as song birds have been found in Japan and date back to the 18th century, while other point to the 12th century (Kovach, 1974). Other researchers (Chang et al., 2005) showed that wild quail were reared as song birds in China as far back as 770-476 BC, and were used as sacrificial animals even earlier. Male quail are used for fighting in some parts of Asia (Kuo, 1960; Mills et al., 1997; Chang et al., 2005), according to a long-standing tradition. There is evidence that this sport has been practiced by ancient Greeks and Romans (Magnin, 1838; Brugas and de Witte, 1868; Morgan, 1975). Most probably, the catching and rearing in captivity of Asian quail occurred in parallel in China and Japan, and possibly in other Asian countries where the birds were distributed. According to Crawford (1990) quail domestication occurred the 11th century in Japan, China or Korea and it is generally assumed that already domesticated quail were imported in Japan from China or Korea around that time (Howes, 1964; Wakasugi, 1984; Crawford, 1990; Chang et al., 2005). There are written testimonies that Japanese quail eggs and meat were used in eastern food and medicine from the 17th century onward (Genchev, 2014). According to Howes (1964) true domestication has occurred during the last century.

The more serious achievements of selection of Japanese quail as a productive bird species date back to 1910-1941 (Mills et al., 1997). At that time, a population with substantially better egg-laying and meat traits than other captive quail was created in Japan (Donchev and Angelov, 1971). Quail from this population had a similar appearance to their wild ancestors, but had higher egg production and a lack of brooding instinct. By virtue of purposeful selection of the species, egg production has dramatically increased and now approaches that of chickens.

Prolonged variable selection has resulted in greater genetic distance from wild ancestors. Differences are associated with a number of changes - morphological, behavioural and productive. The main morphological distinctions affect appearance, such as body size and weight (Mills et al., 1997; Chang et al., 2009), plumage colour (Cheng and Kimura, 1990; Mizutani, 2003; Tsudzuki, 2008), and other phenotypic traits (Chang et al., 2001; 2009). Differences between wild and domestic quail behaviour have been noticed, including their migratory (Deregnacourt et al., 2005), sexual (Mills et al., 1997; Chang et al., 2009) nesting, brooding (Nichols, 1991), and other responses (Nichols, 1991; Chang et al., 2009). Prolonged selection on different productive traits in domesticated quail positively affects their egg laying capacity (Chang et al., 2009), sexual maturity (Mizutani, 2003; Cheng et al., 2010), egg size (Wilson et al., 1971; Chang et al., 2009), fertilisation and hatchability (Wilson et al., 1971) compared to those in wild Japanese quail.

Quail in industrial poultry farming

Japanese quail are widely geographically spread as domesticated bird species. They are used in farming systems for production of meat and eggs (Baumgartner, 1994; Chang, 1998; Minvielle, 1998; Minvielle, 2004; Vali, 2008; Genchev, 2014; Lukanov and Genchev, 2018), and, from around 60 years ago, as laboratory animals (Padgett and Ivey, 1959; Reese and Reese, 1962; Kovach, 1974; Sano et al., 1995; Minvielle, 2004; Huss et al., 2008; Baer et al., 2015; Pavlova et al., 2018). Last but not least, this is a decorative bird species (both wild and domestic) reared by amateur poultry breeders on a global scale (Lukanov et al., 2018). In some Asian countries (Pakistan, Afghanistan, China etc.) male quail are used for fighting (Mills et al., 1997).

Worldwide, the main production of quail eggs is concentrated in East Asia and Brazil, and meat in Europe, the USA and China (Baumgartner, 1994; Minvielle, 1998; 2004; Bertechini, 2012). Although the estimation of the quail population involved in industrial farming is difficult, it has progressively increased over the last few decades. At the beginning of the millennium, about 1 billion Japanese quail were reared globally (Xu et al., 2003; Chang et al., 2007). According to Duval et al. (2012) over nine billion quail are produced each year in East Asia (China, Japan, Hong Kong). China is the biggest producer of quail meat and eggs producing 146,000 to 190,000 tonnes of meat and about 90 billion eggs on an annual basis (da Cunha, 2009). For comparison, in the early 1990s, China produced about seven billion eggs annually, and Chinese quail meat production was behind Spain, France and the USA (Minvielle, 2004). Another serious player on the world's quail retail market is the European Union (mainly Spain, France, Italy and Portugal) with about 100 million reared quail (www.ciwf.org.uk), mainly used for meat production. The USA dominates meat-type quail farming with about 19 million fattening birds (da Cunha, 2009). The continuously increasing production of eggs and meat in Brazil with more than 11.5 million quail should not be underestimated (Bertechini, 2012). By the early 2000s, the industrial quail farming population in Japan decreased to seven million layers (Mizutani, 2003), that, according to Minvielle (2004) produce yearly about 1.8 billion quail eggs. Despite the serious increase in the number of quail reared in Australia in 2001-2002 up to 6.5 million (da Cunha, 2009), in 2010 the population decreased to 3.5 million birds (Department of Agriculture, Australia, 2010, cited by Farooq, 2014). Other countries with well-developed quail production are Canada, Egypt, Saudi Arabia, India, Estonia, Russia, Singapore, Venezuela, Peru, Columbia and Bolivia. It can be summed up that the global production of quail meat and eggs is concentrated in several countries - China, Spain, France, Italy, Brazil, the USA and Japan. Quail produce about 10% of all table eggs in the world, or around 1.21.3 million tonnes, leaving chickens as a dominating species with production of about 72 million tonnes eggs in 2016 (Conway, 2017). On the basis of available data, world quail meat production is estimated to be 200,000 to 240,000 tonnes, that is, about 0.2% of the world poultry meat produce (Figure 1).

Despite the large number of reared birds, the small size of quail and their eggs negatively influences quantitative parameters for production. The FAO database contains data on chickens, turkeys, ducks, geese and guineafowl, pigeons and others. According to FAOSTAT the global numbers of pigeons and other birds for 2016 was about 28.5 million (FAOSTAT, 2018). Taking into account the estimated number of quail reared in the world, it is clear that quail were not included at all in the statistics, despite being included in the list of domestic animals (FAO, 2000). There was a statement that, in 2016, chickens accounted for some 91% of the world's poultry population, followed by ducks (5%), turkeys (2%), other poultry, such as geese and guineafowl, made up the remaining 2% (FAO, 2018). The lack of exact data for production from domestic quail as a farmed bird species neglected its role in poultry farming. The available data suggested that the domestic quail population involved in global production of poultry meat and eggs comes second after that of domestic chickens (Figure 2). This clearly confirms the role of domestic quail for human consumption and the need for throwing more light on this avian species.

Figure 1 Global poultry meat production by species
(FAOSTAT, 2018 and estimated data for quail).

Figure 2 Average share of the domestic birds, slaughtered annually worldwide
(FAOSTAT, 2018 and estimated data for quail).

Quail as a laboratory animal model, pet and show bird

Domesticated Japanese quail are often used as laboratory animals (Huss et al., 2008; Cheng et al., 2010; Andraszek et al., 2014; Baer et al., 2015). Due to its small body size, robustness, fast maturation, laying ability and cage adaptive housing, they are known as the 'Drosophila of the avian laboratory' (Padgett and Ivey, 1959; Reese and Reese, 1962). Their genome has been fully sequenced (Nishibori et al., 2001; Kawahara-Miki et al., 2013), which makes them suitable for genetic studies, replacing the domestic chicken (Andraszek et al., 2014). Domesticated Japanese quail are used as laboratory animal models for studies in developmental biology, animal science, veterinary and human medicine (Huss et al., 2008; Tserveni-Goussi and Fortomaris, 2011) and genetic and epigenetic investigations (Minvielle, 2004; Huss et al., 2008; Andraszek et al., 2014; Baer et al., 2015), including as a transgenic bird model (Shin et al., 2008; Poyntner et al., 2009). All this put them on the top of the bird laboratory list.

Domesticated Japanese quail are popular as an ornamental, exhibition and pet bird in many areas in the world (Lukanov et al., 2018). Their small size, fast growing and maturing, easy care, taming and different plumage coloration makes them preferable for a lot of poultry fans. Some quail breeder associations and clubs have been established in Europe, such as Sondervereines der Japanischen Legewachteln (Germany), DFKP-Club (The Netherlands), improving their potential as a show bird. The European association of poultry, pigeon, cage bird, rabbit and cavy breeders separate the domesticated Japanese quail from the wild ones, naming them 'laying quail', and are preparing a European standard (ESC-P, 2018).

Poultry or game bird, where is the Japanese quails place?

In poultry farming, several species are reared as productive fowl, for example chicken (Gallus gallus domesticus, Linnaeus, 1758), turkey (Meleagris gallopavo, Linnaeus, 1758), geese (Anser anser domesticus, and Anser cygnoides domesticus, Linnaeus, 1758), duck (Anas platyrhynchos domesticus, Linnaeus, 1758), Muscovy duck (Cairina moschata domestica), pigeon (Columba livia domestica), Japanese quail (Coturnix japonica, Temminck and Schlegel, 1849), guineafowl (Numida meleagris, Linnaeus, 1758), as well as ostriches (Struthio camelus domesticus, Linnaeus, 1758), the common pheasant (Phasianus colchicus, Linnaeus, 1758) and Virginia quail (Colinus virginianus, Linnaeus, 1758). Less important, from economical point of view, are the other ratites reared in farms - emus, rheas and cassowaries (FAO, 2000). As can be seen from species' names, three domestic bird species are implemented in industrial production of meat and/or eggs without a clear distinction between them and their respective ancestors. These include Japanese quail, guineafowl and turkeys. Similarly, in species designated as semi-domesticated (e.g. the common pheasant and the Virginia quail), no differentiation exists between captive and wild birds, which is logical as no major differences are present. Unlike these, domestic quail, turkeys and guineafowl are unlikely survive in the wild because of physiological, morphological and ethological changes occurring consequently to domestication and selection for productive traits. In the scientific literature, a clear distinction from the wild form is made only with respect to turkeys by adding the adjective 'domestic' (Hart et al., 1999; Radzimirska, 2003; Maxwell, 2008; Pazdzior et al., 2012; Dalloul et al., 2010) and/or using a trinomen Meleagris gallopavo domesticus/domestica (Hejlicek and Treml, 1995; Radzimirska, 2003; Pazdzior et al., 2012; Olias et al., 2014). In some sources, including those referring for other domestic fowl species, the terms 'variety' and 'forma' are used to designate the subspecies, similar to the botanical trinomial nomenclature, but this is not considered proper for zoological use (ICZN, 2000). While there are no serious taxonomical changes in the name of guineafowl, the name for Japanese quail was changed from Coturnix coturnix japonica to Coturnix japonica, placing the bird in a different species from the common quail Coturnix coturnix (McGowan, 1994; Clements, 2002). Even some modern literature sources still mention the bird as Coturnix coturnix japonica (Kul and Seker, 2004; Antarasena et al., 2006; Scholtz et al., 2009; Alkan et al., 2010; Duval et al., 2013; Franqueza et al., 2016; Akdemir et al., 2018), and domesticated Japanese quail is reported as Coturnix coturnix (Orsi et al., 2017; Cox et al., 2018). Another serious concern is the free usage of the word combination 'Japanese quail' without specifying whether the wild or domesticated Japanese quail was meant. This, of course, does not decrease the worth of cited reports, but the lack of a uniform term is confusing for readers. Regardless of abovementioned, there is a positive tendency for increasing use of the term 'domestic quail' or 'domestic Japanese quail' in the scientific literature (Mills et al., 1997; FAO, 2000; Chang et al., 2009; Orsi et al., 2017; Nunome et al., 2017; Sun et al., 2017; Shalini et al., 2018).

Quail farming in EU and its legislation

Another interesting point is the lack of clear regulations pertaining to the production and quality of domestic quail meat and eggs in the European Union. This is probably due to an existing discrepancy about the designation of the domestic quail - both farm or game species. The following fowl species are included in Commission Regulation No 543/2008 (EC, 2008a): domestic fowl (Gallus domesticus), turkeys (Meleagris gallopavo dom.), ducks (Anas platyrhynchos dom.), Muscovy ducks (Cairina muschata), Mallard ducks (Cairina muschata x Anas platyrhynchos), geese (Anser anser dom.) and guinea fowl (Numida meleagris domesticus). According to the definition of EC No 798/2008 (EC, 2008b) and EC No 158/2009 (EC, 2009), 'poultry' refers to fowl, turkeys, guinea fowl, ducks, geese, quail, pigeons, pheasants, partridges and ratites (ratitae), reared or kept in captivity for breeding, the production of meat or eggs for consumption, or for restocking supplies of game. This definition encompasses other game species, such as partridges and pheasants. A similar situation exists for stock egg production, quality evaluation, storage and marketing. In the existing regulations, only chicken eggs are considered, omitting the fact that there are other species producing eggs for consumption, e.g. the domestic quail. EC No 589/2008 (EC, 2008c) defines the term 'eggs' as 'eggs in shell i.e. other than broken, incubated or cooked eggs, that are produced by hens of the species Gallus gallus and are fit for direct human consumption or for the preparation of egg products'. Despite the lack of clear regulations on quail eggs, they partly follow the rules applied for chicken stock eggs. Quail eggs, however, are not subject to grading by weight, marking is impossible due to specific coloration of the eggshell, candling and exact determination of the air cell size are impossible. The widescale introduction of domestic quail in industrial poultry breeding should be reflected in such legislation and the latter should take account of the specific features of the product, its production, quality evaluation, storage, packaging and marketing. Only a comprehensible legal base referring to this bird species will prevent a double standard among producers and will largely avoid the possibility of subjective interpretation of existing legislation on the part of controlling bodies.

Conclusions

In conclusion, domesticated Japanese quail should be referred to as 'domestic quail' (Coturnix japonica domestica), in its distinction from the wild Japanese quail (Coturnix japonica). Within the EU legal framework, domestic quail should be regarded as a domestic fowl species, as they are used for production and are including in intensive rearing systems. Similar terminology could be applied to other domestic birds to distinguish them from their wild ancestors.

H. Lukanov, Department of Animal Science - Monogastric and Other Animals, Faculty of Agriculture, Trakia University, Stara Zagora, Bulgaria
World's Poultry Science Journal, Vol. 75, December 2019

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Domestic quail (Coturnix japonica domestica), is there such farm animal?