Introduction

The agroecological transition in livestock farming relies on decreasing farm inputs and increasing the use of local feed resources and farm leftovers to meet the needs of the on-farm production procedure and on grazing modalities closely following ecological processes in order to ensure the renewal of resources in the long term (Dumont, Fortun-Lamothe, Jouven, Thomas, & Tichit, 2013; Jouven, Lapeyronie, Moulin, & Bocquier, 2010). Worldwide, in pastoral farming systems of dairy sheep and other ruminant species, feed requirements are covered to a varying degree through grazing of local fodder resources under a range of management systems (FAO, 2001). Grazing on a diversity of fodder plant species on rangelands also aids the conservation of landscapes (D’Ottavio et al., 2016) and limits soil erosion resulting from continuous cropping (Schnabel, Gutierrez, & Contador, 2009) in a way that is not competing with the use of land for other food production. Moreover, grazing of local resources promotes the biological interactions between animals and their biophysical environment, providing an interesting basis to design agroecological production systems (Dumont et al., 2013).

Despite these benefits, pastoral farming systems are disappearing, mostly in disadvantaged areas, but also in favourable sites of North Mediterranean countries, leading to degradation of traditional rural landscapes (Caballero et al., 2009; Hadjigeorgiou, Osoro, Almeida, & Molle, 2005). The disappearance of pastoral activity in many regions in recent decades has revealed the importance of sustainable livestock management for environmental conservation. The absence of pastoralism has had negative consequences on biodiversity and ecosystem services, particularly in marginal areas traditionally used for this activity (Constanza et al., 1997). In many ecosystems with a deep-rooted grazing tradition, such as those of the Mediterranean (Hadjigeorgiou, 2011), the withdrawal of livestock farming activities has led to broad negative changes in the richness and diversity of plant species (Sternberg, Gutman, Perevolotsky, Ungar, & Kigel, 2000). Moreover, the abandonment of pastoral farming has led to modification of various interspecies interactions, affecting negatively, for instance, pollinators, herbivorous insects, parasitoids and birds (Dover, Spencer, Collins, Hadjigeorgiou, & Rescia, 2011; Plieninger, Hochtl, & Spek, 2006). Similarly, this change favoured woody vegetation encroachment, leading to the accumulation of fuel biomass and consequently raising the danger of wildfires (Rosa-García, Celaya, García, & Osoro, 2012). At the same time, ruminants’ farming systems have evolved towards intensive ones and established in more accessible areas, such as lowlands and hilly areas of the mainland. Examples include reports on zero grazing in Castilla-Leon in Spain or sedentary intensive or semi-intensive systems in Greece (Caja & de-Rancourt, 2002; de-Rancourt, Fois, Lavin, Tchakerian, & Vallerand, 2009), the abandonment of winter transhumance under agricultural pressure in the Ebro valley in Spain (Caballero et al., 2009) or the replacement of permanent grasslands by agriculturally improved pastures in Italy (Porqueddu et al., 2017).

In parallel, the evolution of flock composition in breeds and their crosses shows different patterns in Mediterranean regions subjected to intensification of farming systems and decreased use of native grazing resources. For example, in north and central Spain, as well as in Greece, farms with highly productive sheep breeds and their crosses coexist with those of local sheep breeds ¹ (e.g. (Perucho et al., 2018; de-la-Fuente, Gabina, Carolino, & Ugarte, 2006). However, in other North Mediterranean regions, also subjected to intensification of farming systems, local breeds remain the main genetic resources in sheep farming. In the South of France, for instance, the Black-Face Manech and the Red-Face Manech are traditionally raised due to their hardiness in the transhumant systems of the West Pyrenees, although the Red-Face Manech now prevails over the Black-Face Manech in most intensive farming systems of the area (Labatut, 2009; Lauvie, Paoli, & Moulin, 2015). Likewise, in the Crau region in France the Merino breed is used due to its specific transhumance ability, but it is also raised in many different sedentary farming systems, including mixed farming systems with an intensive use of land, based on cereals and hay cropping (Lauvie et al., 2015; Moulin, Pluvinage, & Bocquier, 2004). Moreover, holdings with local sheep breeds are found in many different biogeographical regions in Italy under sedentary or itinerant systems often combined with cropping (Caballero et al., 2009).

We thus hypothesized that the use of local breeds may not systematically ensure the preservation of pastoral practices in Mediterranean dairy sheep farming systems. In order to test this hypothesis, we explored holdings using different livestock breeds, as well as the evolution of their feeding systems with emphasis on pastoral practices. In particular, we focused on two production territories where dairy sheep farming systems were based on the use of different livestock breeds or their crosses, while they were subjected to similar trends towards intensification. Each case study brought up features from different situations to illustrate our argumentation.

Material and methods

Results

Comparative evolution of feeding systems and crossbreeding practices in Thessaly

Figure 1 illustrates changes in the feeding systems (in terms of supplementation, grazing and crops for animal consumption) and the breeds introduced in the flocks of 14 dairy sheep farms in Thessaly after the first crossbreeding with a highly productive breed (first yellow square of each line). Among these numerous changes of breeds and feeding systems, several were justified by farmers after a failure of the new breed to adapt to the existent feeding system. Starting from these specific cases, we analysed the different drivers of evolution of breeds and feeding systems in the short and long-term.

Introducing highly productive breeds displayed a lack of adaptation to existing feeding systems

A lack of adaptation of newly introduced, highly productive breeds (or their crosses) in the flock to existing feeding strategy and grazing conditions was mentioned by nine farmers. These inconsistencies between breeds and feeding systems took three forms: (i) the breed/crossbreed was deemed not hardy enough to function under existing grazing conditions; (ii) the feeding strategy implemented for the breed was considered economically unsustainable; and (iii) the various breeds, raised simultaneously in the flock, had different feeding requirements, but it was not practical to implement a different feeding practice for each of them (Table 3).

Three types of farmers’ responses (adjustments) to this lack of adaptation were identified. The first type was the modification of the feeding/grazing strategy towards decreasing the role of native resources in the applied production system, represented by red stars in $. The second type consisted of the rejection of the highly productive breed in use and/or the testing of a new breed: farmers adopted an exploratory behaviour towards locally available breeds, leading to several changes in breeding animals in a short time period, according to availabilities offered by the market and consultation with other farmers in the region and elsewhere. The third type of farmers’ response was the reintroduction of breeding animals from the local hardy Karagouniko breed in the herd, represented by brown triangles in Figure 1.

In some farms, the failure of the highly productive breeds to adapt to the feeding system in place was not explicitly mentioned by the farmer, but feeding systems were modified after their introduction. Some of these changes in feeding systems occurred in the general frame of the intensification of farm production means, i.e. the purchase of highly productive breeding animals (yellow squares in Figure 1) came with an investment in feed quantity and quality, housing, equipment and task mechanization in order to maximize the expression of the specific breed’s productive potential. The introduction of a highly productive breed and the decrease of grazing and/or the change in diet quantity or quality generally occurred simultaneously or within a short time period after breed introduction (example of changes in farms 1, 2, 3, 4, 5 or 6 during the decade 2000-2010, Figure 1).

Other changes in feeding systems were driven by the following four factors: (i) a change in land access (example of farm 8 since 2003, Figure 1, (ii) economic or climatic perturbations, (iii) a modification of the role of sheep farming in the family’s income and (iv) the increase in average flock size. They occurred at different moments regardless of the change of breed (examples of changes in farms 9 and 13 in 1995 and 1997-1998, respectively). The above mentioned drivers of change could be combined (one perturbation implies farm intensification). For example, in mixed farming holdings, traditionally based on cash crops and sheep farming (farms 3,5 and 6 in Figure 1), the combination of decreasing cotton prices (since the 2000’s) and the decoupled subsidies for cotton crops (since 2006) led to an increasing role of sheep farming in the family income, together with the search for new alternatives to cotton crops. According to farmers’ economic possibilities, subsequent changes in sheep farming activities were immediate (farms 5 and 6) or progressive (farm 3) and included the testing of different, highly productive breeds combined with high supplementation levels and the replacement of cotton crops by crops destined for animal feeding.

Table 3: Components of the feeding system and the traits of the breed involved in the different forms of inconsistencies mentioned by farmers in Thessaly

Inconsistency (number of farms)	Component of the feeding system at stake	Traits of the breed at stake (number of mentions)
Lack of hardiness (n=8)	Grazing conditions (Climatic conditions)	Sensitivity to thermal stress (n=1)
	Grazing conditions (Duration of grazing/nature of the foraging resource)	Feeding mode leading to sensitivity to mastitis and ruminal pathologies (n=1)
	Grazing conditions (Presence of pathogens)	Sensitivity to vector-borne diseases (n=2)
	Grazing conditions (Distance and topography)	Walking ability (n=3)
	Grazing conditions (Open-field pastures)	Grazing behaviour (n=2)
Feeding costs (n=5)	Feed supply (quantities at the multiyear scale)	Longevity (n=2)
	Feed supply (quantities at the yearly scale)	Feeding requirements (n=3)
Competition between breeds (n=1)	Feed supply (quantities at the yearly scale)	Feeding intake (n=1)

Introduction of Karagouniko local breed for hardiness was not necessarily associated with maintenance of pastoral components

The reintroduction of the local Karagouniko breed was operated by farmers in three of the nine farms for which inconsistencies between flock genetic composition and feeding system were mentioned. In farmers’ statements, the characteristics of the highly productive breeds/crossbreeds forcing such changes in the flock genetic composition were the following: high sensitivity to thermic stress (n=2), high cost of feeding (n=1), high cost of animal health care or high sensitivity to mastitis (n=2), inefficient performance on pasture (n=1), a lack of adaptation to transhumance (n=1), and difficulty in hand milking (n=1). Likewise, the results from the T1 interviews indicated that the dairy sheep farmers of Thessaly utilised local breeds in their holding in order to improve the hardiness of the flock (in terms of adaptation to pastoral conditions) and rejected highly productive breeds for their incapacity to do so. These abilities and the breeds used or rejected for their corresponding characteristics are presented in Table 4.

Table 4: Breeds’ traits motivating the introduction or rejection of a specific breed in the flock: the case of traits linked to pastoral farming. Hardiness (or a similar term) was used by farmers to characterize the overall ability of the breed to withstand the constraints of its raising conditions.

Reasons for introducing or rejecting a specific breed (traits linked to pastoral farming)	Percentage of sampled farmers mentioning the trait (n=42 farmers)	Breeds used ( in bold ) or rejected (normal font)
Ability to handle transhumance	14% (n=6)	Kalarritiko breed
Low sensitivity to cold and high humidity	50% (n=21)	Karagouniko breed and local populations Chios, Lacaune, and Frizarta/Friesian breeds
Low sensitivity to heat	45% (n=19)	Karagouniko and Awassi breeds Frizarta/Friesian and Assaf breeds
Hardiness	48% (n=20)	Karagouniko , Kalarritiko , Piliou and local populations Chios and Lacaune breeds
Behaviour compatible with grazing	26% (n=11)	Karagouniko breed Lacaune breed

However, in the medium and long-term, the pastoral components of the feeding systems of those three farms were not necessarily maintained. In the first farm (farm 5), maintaining grazing on communal grasslands, in order to keep feeding costs low, was part of the farmer’s strategy according to his statement. Reintroduction of Karagouniko purebred animals aiming to keep a hardy flock, several years after the first crossbreeding, allowed the farmer to maintain this grazing practice for the following years (farm 5 in Figure 1). In a second farm (farm 6 in Figure 1), reintroducing the Karagouniko breed, through crossing, was motivated by the low capacity of the highly productive breeds to cope with climatic constraints. However, the newly composed crossbred flock remained disappointingly sensitive to thermal stress, as well as to health risks on communal grasslands, which led the farmer to decrease grazing a few years after the introduction of the local breed. In a third farm (farm 11 in Figure 1) using native grasslands (including summer pastures through transhumance) was part of the farmer’s strategy but, ultimately, the maintenance of this practice competed with workload management. In this case, crossbreeding with the local breed temporarily delayed the stoppage of transhumance. However, the introduction of machine milking led this farmer towards a sedentary system (this is not presented in the figure, but it was planned as a short-term project by the farmer at the year of the interview). It is also interesting to mention that the modalities of the introduction of the local breed in these three farms were limited to the introduction of breeding males in the specific year that the problem was observed and it was not followed by a concrete crossbreeding plan with the introduced local breed.

Local purebred flocks in a diversity of feeding systems in Corsica

Currently, different feeding systems can be observed among local purebred flocks of Corsica (Corsican sheep breed), ranging from the most pastoral to the most intensive in feeding inputs and land use and including both transhumant and sedentary flocks.

Figure 2 presents a classification of five types of feeding systems (FS1 to FS5) according to the role of grazing in covering the energy requirements, the type of grazed pastures and the farmers’ strategy with respect to the provision of hay. Most of the pastoral systems (characterized by less harvested feed energy used compared to that of grazed native resources) are found in southwestern and central Corsica (FS1 and FS2), the most intensive feeding systems (in terms of feeding inputs and workload for fodder production) are located in the eastern coastal lowlands (FS5), and the intermediate feeding systems are found in all locations of the island (FS3 and FS4). This current situation of local breed dairy sheep farming conditions in Corsica reflects a trend towards the securing of feeding systems, either by on-farm fodder production in areas with favourable agroecological conditions, or by hay purchased at the market. The diversity of land use in Corsica, for the 206 dairy sheep farms using the Corsican breed, has also been documented byPerucho et al. (2020)

Adaptation to variations of fodder on offer through replacement and culling management in local purebred flocks of Corsica

In addition to the use of local breeds in purebred flocks, several of the interviewed farmers performed specific breeding practices in order to maintain the same feeding system in spite of the variations in climatic conditions and subsequent fodder offer. These breeding practices consisted of adjusting the demography of the flock according to inter-annual variations of fodder on offer. In some pastoral systems of southern and central Corsica (farms 1, 2, 3 and 4; included in FS1 and FS2, Figure 3, this consisted of increasing the replacement rate in years with good forage offers (FORAGE VAR/REP RATE, line 4 of the table in Figure 3) and of reversing the practice in years of low forage offers. Moreover, in pastoral systems FS1 and FS2, the observed replacement rates were generally low (less than 20%, REP RATE, see line 2 of the table in Figure 3). Another adaptation of the flock demography consisted of increasing the culling rate in order to limit the number of animals that feed during the lambing period of the following year (farms 11, 12, and 13, FORAGE VAR/CULL RATE, line 3 of the table in Figure 3). This practice was observed in farming systems relying partly on cultivated forage (FS3) in areas of the island impacted by a dry microclimate (e.g. north western lowlands), resulting in variable fodder production from one year to another. These examples, although concerning only a small number of farmers of the sample, demonstrate that the use of a local breed might be combined with other breeding practices so that the farmer is able to maintain the same feeding system from one year to another.

Discussion

Our results indicate that the practice of crossbreeding with highly productive breeds in Thessaly is often connected with the intensification of feeding by increasing the quantity of feed inputs and modifying the diet with the aim of improving its quality. Similarly, the use of local breeds is usually linked with the applied pastoral practices and prevailing climatic conditions. Boyazoglu and Hatziminaoglou (2005) described the long-term evolution of feeding systems in the European part of the Mediterranean basin, characterised by a decrease in transhumance and the use of grazing areas, while the population of small ruminants maintained its genetic composition. On the other hand,Couix, Gaillard, Lauvie, Mugnier, and Verrier (2016) showed that, in dairy cow holdings, the replacement of the Holstein breed by dual purpose local breeds in the western part of France was associated with an overall evolution of the farming systems towards decreased production costs, including the costs associated with the feeding system. In this study, a change in breed was often followed by the adoption of pasture-based feeding systems.Samdup, Udo, Eilers, Ibrahim, and Zijpp (2010) analysed the adoption of crossbreeding with different exotic, highly productive breeds in a range of cattle farming systems in Bhutan (from extensive to intensive livestock farming systems) and the impact of crossbreeding on such farming systems four years after its implementation. They observed a lower adoption rate of crossbreeding in extensive and semi-intensive farms with the farms either keeping the local breed or rejecting one specific exotic breed (among the two introduced) due to its high feed requirements. The authors also mentioned that during lactation, crossbred cows were stall fed, resulting in reduced grazing in forest and natural grasslands in comparison to local cattle (Samdup et al., 2010). The same conclusions were drawn in a study in Ethiopia (Roschinsky, Kluszczynska, Sölkner, Puskur, & Wurzinger, 2015), where most of the cattle farmers adopting crossbreeding with exotic breeds changed their grazing and feeding management towards restricted grazing and the purchase of compound feeds or household by-products, while feed shortages and feed prices were the drivers for rejecting the practice of crossbreeding.Caballero et al. (2009) also mention the decline of some indigenous breeds in Spain, Italy and Greece and the abandonment of extensive livestock farming in marginal areas. Apart from diachronic studies, studies dealing with farmers’ preferences for breeds’ traits emphasize the key role of local breeds’ adaptive traits in pastoral systems (Kosgey, Rowlands, Arendonk, & Baker, 2008; Tamou, Boer, Ripoll-Bosch, & Oosting, 2018) and the preference for exotic breeds in agropastoral systems benefitting from best pastures and climatic conditions (Konig et al., 2015). These results agree with the first part of our results associating, on the one hand, crossbreeding for higher productivity with feeding intensification and, on the other hand, local breeds with lower feeding inputs in pastoral systems.

However, these two associations are not systematically observed, and farmers do not necessarily abandon raising local breeds following an intensification process. The example of Corsica, where one local breed is raised under feeding systems with different levels of intensification, and the one of Thessaly, where a variety of genetic types can be found at different stages of the intensification process, suggest that additional studies are needed in order to more convincingly conclude whether coevolution of local breeds and feeding systems is regionally univocal. The data collected through interviews should be compared with the physiological and behavioural responses of highly productive sheep breeds and their crosses in pastoral farming systems in North Mediterranean countries. Methods to predict breeds’ suitability to environmental conditions are proposed, in this sense, by (Lozano-Jaramillo, Bastiaansen, Dessie, & Komen, 2018) and (Marshall, 2014), but these concern South Mediterranean or tropical systems. In the case of North Mediterranean countries, there are fewer pastoral components and the climatic conditions are milder, but the need for adaptive potential of the animals to similar environments is increasing in importance due to intensifying climatic changes (Hoffman, 2013). In this sense, learning processes regarding grazing (Meuret & Provenza, 2014) and genetic selection should also be considered when assessing the suitability of North Mediterranean breed types. Finally, our results illustrate that defining the “pastoral dimension” of North Mediterranean systems implies considering a set of pastoral components assessed not in terms of absolute value, but also in relative value: pastoral components of one farm are described with regard to the other farming systems of the area. Accordingly, detailing constraints of feeding systems by pastoral components might be useful in the assessment of breed suitability.

The fact that feeding systems in an area can evolve independently from animal genetic resources used there, is not analysed per se in current research, but rather is suggested by the diversity of factors known to affect the maintenance of pastoral components and the management of animal genetic resources. The evolution of small ruminant feeding systems towards decreased pastoral components is well documented in different Mediterranean countries (de-Rancourt et al., 2009). The drivers of this decrease, as depicted in recent literature, echo the findings of the present study: the farmers wish to alleviate the labour-consuming tasks associated with the feeding system, which leads to a decrease in grazing (Aubron, Noël, & Lasseur, 2016). Other factors such as the farmers’ mind-set and their social environment, the absence of markets for products based on grazing, as well as land fragmentation have been shown to influence the decision to graze or not to graze in European dairy cow holdings (Dasselaar, Hennessy, & Isselstein, 2020). In Greece, the economic context (in terms of prices and subsidies) favoured the choice of purchasing feed outside of the farm (Hadjigeorgiou, 2011; Stefanakis, Volanis, Zoiopoulos, & Hadjigeorgiou, 2007; Volanis, Stefanakis, Hadjigeorgiou, & Zoiopoulos, 2007), together with other factors such as the low quality of spontaneous forage material, the difficulty to access communal rangelands and an inadequate rangeland management system. Other examples from small ruminant farming in Greece also illustrate that the legislative framework can force the settlement of nomadic farmers through specific requirements for facilities to comply with milk and stock hygiene standards, animal welfare and manure management (Hadjigeorgiou, 2011). Finally, the trend towards agricultural intensification in easily accessible productive lands led to land use changes through abandonment of mountainous areas in southern Europe and shrub encroachment in many remote areas formerly used for grazing (Caballero et al., 2009; MacDonald et al., 2000). This phenomenon is highly dependent on EU agricultural and natural conservation policies (Tzanopoulos et al., 2011).

Likewise, the diversity of factors affecting the evolution of animal genetic resources in flocks has to be taken into account. According to (FAO, 2015), the reported main causes of genetic erosion in 23 countries in Europe and the Caucasus (in response to open-ended questions) mostly consisted of (i) breeds not profitable/competitive or that have poor performance (48% of the countries), (ii) intensification of production or decline of traditional production systems or small-sized farms (39% of the countries) and/or (iii) the introduction/increased use of exotic breeds (35% of the countries). Indeed, the introduction of exotic breeds through uncontrolled crossbreeding (failure or absence of crossbreeding programmes) greatly compromised the conservation of local breeds (Leroy et al., 2016). More than 20% of the above mentioned countries also mention problems in breed management linked to weak or absent management policies, programmes or institutions (FAO, 2015). Indeed, except for their adaptive traits, local breeds in European countries are also kept (being the first motivation) for their links with tradition and their importance in society and, unequally among countries and breeds, for the economic incentives and conservation programmes that benefit them (Gandini et al., 2010). In this context, the successes or failures faced by national genetic management programmes and the associated involvement of researchers in such management (Leroy, Baumung, Boettcher, Scherf, & Hoffman, 2016) directly impact the evolution of local breeds’ populations. In Greece, for example, the state-run genetic management of local breeds has suffered several interruptions in financing, impacting the implementation of data recording (Georgoudis & Ligda, 2000). Together with unfavourable dairy policies, the problems in the implementation of breed management programmes have been impacting on, for example, the breeding scheme of the Karagouniko breed in Thessaly (e.g. (Perucho et al., 2019). On the other hand, the financial support, through EU-funded agri-environmental measures, aimed at farmers raising local breeds threatened by extinction, together with the support provided to farmers in mountainous or disadvantaged areas, has succeeded in maintaining local breed populations over time, as is, for example, the case for the pastoral sheep farming systems of the Kalarritiko purebred flocks (National Rural Network, 2019).

Conclusion

The example of long-term changes in dairy sheep farms in Thessaly, Greece, indicated that dairy sheep farmers sometimes use local breeds in crossbred flocks in order to improve flock hardiness after a first crossbreeding with highly productive breeds. In this strategy grazing practices can be maintained and feeding costs reduced together with other health costs. In some cases, the shift to new exotic breeds was also considered as an alternative to local breeds in order to improve flock hardiness. However, any additional factors impacted the management of feeding systems, such as local trends towards intensification, access to production means (e.g. land, workforce, capital) and workforce management. This resulted in an evolution of feeding systems (decrease of grazing on native grasslands – increase of supplementation) not necessarily in accordance with the initial reason for the introduction of the local breed in the flock (the maintenance of pastoral components of the feeding system). Likewise, although the Corsican sheep breed is considered as hardy, its use could be maintained in different feeding systems including systems engaged in an intensification process. Finally, farmers in Corsica chose to act on flock structure to adapt to several constraints linked to the forage offer in their local purebred flocks. This result indicated that the use of a local breed in purebred or crossbred flocks should be combined with several other practices or conditions so that raising local breeds remains closely related to the pastoral activity.

By demonstrating a non-univocal co-evolution between local breeds and farming systems, this study highlights the need to better characterize local and exotic breeds’ abilities as well as farmers’ strategies to cope with different perturbations of their environment. This knowledge will help maintain pastoral systems in production territories and the livestock breeds associated with these territories.

Conflict of interest statement

The authors declare that no conflict of interest exists.

Author contributions

The authors confirm contribution to the paper as follows: study conception and design: Lola Perucho, Anne Lauvie, Charles-Henri Moulin, Jean-Christophe Paoli, Christina Ligda; data collection: Lola Perucho; analysis and interpretation of results: Lola Perucho, Ioannis Hadjigeorgiou, Anne Lauvie, Charles-Henri Moulin, Jean-Christophe Paoli, Christina Ligda; draft manuscript preparation: Lola Perucho; manuscript revision: Lola Perucho, Ioannis Hadjigeorgiou, Anne Lauvie, Charles-Henri Moulin, Jean-Christophe Paoli, Christina Ligda.