Keywords: Brand Social Responsibility, Brand Commitment, Brand Trust, Brand Identification, Personal Norms

Why do consumers consider social responsible associations in their relationship with brands? The case of organic food brands

Introduction

Corporate Social Responsibility (CSR) has traditionally been conceptualized as a managerial obligation to take measures that protect and improve well-being, both of society and firms' interest [START_REF] Sen | Does doing good always lead to doing better? Consumer reactions to Corporate Social Responsibility[END_REF]. Despite the scientific debate as to the influence of CSR on firms' financial and economic performance [START_REF] Brown | The Company and the product: corporate associations and consumer product responses[END_REF], CSR is generally considered to be an asset for firms, extending its shared value [START_REF] Lindgreen | Corporate social responsibility brand leadership: a multiple case study[END_REF]. More and more companies consider CSR to strengthen the performance of their relations with principle stakeholders in particular with clients (Bhattacharya & Sen, 2004).

Indeed, research has shown that positive associations in terms of CSR increase consumers' trust towards the company (Swaen & Chumpitaz, 2008), leading to more positive evaluations of product quality and increased purchase intentions [START_REF] Du | Reaping Relational Rewards from Corporate Social Responsibility: The Role of Competitive Positioning[END_REF].

Besides, positive associations affect customer satisfaction and loyalty (Bhattacharya & Sen, 2004).

Thus, more and more practitioners consider social responsibility as a central element of brand differentiation policy [START_REF] Maignan | A stakeholder model for implementing social responsibility in Marketing[END_REF][START_REF] Hildebrand | Corporate Social Responsibility: A Corporate Marketing Perspective[END_REF]) that corresponds to a new market of social responsibility [START_REF] Lindgreen | Corporate social responsibility: An empirical investigation of U.S. organizations[END_REF]. Certain brands such as "Ben & Jerry", "Ecover", "Green Mountain", "Patagonia" and "Björk", go beyond this, positioning themselves as "the socially responsible brand" in their activity sector [START_REF] Lindgreen | Corporate social responsibility brand leadership: a multiple case study[END_REF]. However, most research has only focused on social responsibility holistically, without taking account of its different facets that are likely to condition consumer reactions.

Despite the number of studies demonstrating the influence of social responsibility associations on the consumer-brand relationship [START_REF] Sen | Does doing good always lead to doing better? Consumer reactions to Corporate Social Responsibility[END_REF][START_REF] Lindgreen | Corporate social responsibility: An empirical investigation of U.S. organizations[END_REF]Lacey & Kenett-Hensel, 2010), to our knowledge, no study has yet focused on the process by which these different dimensions of social responsibility influence the consumer-brand relationship. To study this relationship, we examine brand commitment as a central variable of an effective relational marketing policy towards a durable relationship between consumers and brands [START_REF] Morgan | The Commitment-Trust Theory of Relationship Marketing[END_REF]. It seems crucial to identify the mechanisms and condition of this influence to design policies suited to consumers' expectations in terms of social responsibility. The objective of this research is thus to study how and in which conditions these dimensions of social responsibility influence the process of brand commitment.

1.

Conceptual framework 1.1 Social responsibility and consumers 1.1.1. The social responsibility of brands from consumers' viewpoint

In marketing, CSR is generally studied from the point of view of the consumers' perception of the firm's activities and engagements related to its obligations towards society or stakeholders [START_REF] Brown | The Company and the product: corporate associations and consumer product responses[END_REF]. Most studies on the beneficial effects of CSR consider it as a one-dimensional concept [START_REF] Du | Reaping Relational Rewards from Corporate Social Responsibility: The Role of Competitive Positioning[END_REF][START_REF] Abid | Une échelle de mesure de la responsabilité sociétale des marques : application aux consommateurs de marques alimentaires biologiques[END_REF]. However, it is generally accepted that the domains of socially responsible activities are many and various [START_REF] Sen | Does doing good always lead to doing better? Consumer reactions to Corporate Social Responsibility[END_REF]. In this study we adopt a multidimensional conception of social responsibility in order to capture the concept's rich variety (Swaen & Chumpitaz, 2008).

According to [START_REF] Lindgreen | Corporate social responsibility brand leadership: a multiple case study[END_REF], consumers may establish a relationship with a given brand without necessarily knowing the company behind it. [START_REF] Abid | Une échelle de mesure de la responsabilité sociétale des marques : application aux consommateurs de marques alimentaires biologiques[END_REF] propose an approach to consumers' perception of brands' social responsibility by focusing on consumers' specific brand expectations regarding three dimensions : (1) the consumer dimension, or the brand's capacity to propose safe and healthy products and provide correct information as to their ingredients: (2) the environmental dimension, or the brand's capacity to reduce its environmental footprint and (3) the philanthropic dimension, or the brand's participation in the societal and cultural life of its locality and/or its support of important causes such as handicap or child-protection.

1.1.2. Brand social responsibility: a lever of brand commitment

Consumer commitment can be considered as the central variable of any effective relationship marketing strategy1 , resulting in behaviors such as stable preference, loyalty, resistance to temptation from competitors and/or word of mouth [START_REF] Morgan | The Commitment-Trust Theory of Relationship Marketing[END_REF][START_REF] Chaudhuri | Product-class effects on brand commitment and brand outcomes: the role of brand trust and brand affect[END_REF]Lacey & Kennett-Hensel, 2010).

Indeed, since Morgan and Hunt's studies, most research agrees that engagement is a central variable of relational marketing and plays a crucial role in describing and explaining brand repurchase behavior [START_REF] Doney | An Examination of the Nature of Trust in Buyer-Seller Relationships[END_REF], in particular "true" brand loyalty [START_REF] Fournier | Consumers and their brand: Developing relationship theory in consumer research[END_REF]. The present study follows this attitudinal approach to commitment that is widely mobilized in marketing research on consumer-brand relations [START_REF] Terrasse | L'engagement envers la marque: Proposition d'un modèle théorique et application à la comparaison de la fidélité aux marques nationales et aux marques distributeurs[END_REF].

Commitment can be defined as a psychological state that reflects the consumer's desire to continue a relationship with the brand, even if this means making sacrifices [START_REF] Terrasse | L'engagement envers la marque: Proposition d'un modèle théorique et application à la comparaison de la fidélité aux marques nationales et aux marques distributeurs[END_REF].

The influence of perceptions of social responsibility on consumers' commitment towards a company has received a certain amount of support [START_REF] Pérez | The role of CSR in the corporate Identity Banking Service Providers[END_REF]. Recently, research has proved that this influence also exists between consumers and brands (Lacey & Kennett-Hensel, 2010). Nevertheless, the literature explains this influence through different theoretical perspectives.

When we consider social responsibility from the angle of stakeholder theory developed by

Freeman 2 , CSR is considered as a way of improving relations with company stakeholders by offering them advantages due to the firm's policies and activities [START_REF] Post | Managing the Extended Enterprise: The New Stakeholder View[END_REF]. At the heart of stakeholder theory, is the idea that the firm's long term viability depends on how it manages relationships with stakeholders with a view to mutual advantage. According to this approach, the contribution of social responsibility initiatives on the consumer-brand relationship relies on the brand's capability to develop a contract based on consumer relations [START_REF] Maignan | A stakeholder model for implementing social responsibility in Marketing[END_REF].

Based on social exchange theory [START_REF] Blau | Exchange and Power in social life[END_REF], this approach explains the consumer's wish to develop a long-term exchange relationship with the brand after a subjective analysis of the costs and benefits to be drawn from these exchanges and in comparison with alternative solutions (Stafford,2008). The consumer is committed to brands that he considers socially responsible for instrumental reasons, in other words, because these brands' socially responsible initiatives bring them personal benefits.

However, the influence of social responsibility on the firm-consumer or brand relationship has also been studied from the angle of social identity theory and self-categorization [START_REF] Tajfel | An integrative theory of intergroup conflict[END_REF]. This approach explains the influence of social responsibility through questions of identity. According to this approach, socially responsible initiatives have symbolic value for consumers [START_REF] Mcenally | The Evolving Nature of Branding: Consumer and managerial considerations[END_REF] enabling them to recognize themselves in the values conveyed by these initiatives and thus developing their social identity through their consumption. In fact, socially responsible brands appear to be particularly useful vectors for satisfying consumers' needs for self-definition [START_REF] Fournier | Consumers and their brand: Developing relationship theory in consumer research[END_REF].

2

According to stakeholder theory, the firm, in its role as center of decision, takes on a responsibility "towards all those who whether closely or remotely, depend on its existence, development and survival": stakeholders in its activity (employees, consumers, public authorities, shareholders, suppliers etc.) (Freeman, 1984).

We suppose that the basic process of mediation between the different associations of social responsibility and brand commitment depend on the type of benefit (instrumental or symbolic) that these associations produce for consumers.

1.2 Hypotheses development 1.2.1. The influence of social responsibility on consumers' brand commitment from the perspective of social exchange theory.

The "consumer" dimension of social responsibility refers to the brand's capacity to offer safe, healthy products and give correct information as to a products' composition. This dimension has utilitarian benefits for consumers in terms of health or quality (Swaen & Chumpitaz, 2008;[START_REF] Abid | Une échelle de mesure de la responsabilité sociétale des marques : application aux consommateurs de marques alimentaires biologiques[END_REF].

By sending consumers a signal of its capabilities in terms of product quality control and the reliability of information on product components and origin, a brand can be perceived as credible and honest, two central dimensions of trust [START_REF] Mayer | An integrative model of organizational trust[END_REF].

Similarly, by committing to reduce or eliminate potentially dangerous chemicals from products, a brand sends consumers a signal of its concern for their health and welfare. Thus, consumers are more likely to perceive the brand as benevolent and so, trust it (Swaen & Chumpitaz, 2008;[START_REF] Abid | Une échelle de mesure de la responsabilité sociétale des marques : application aux consommateurs de marques alimentaires biologiques[END_REF].

Brand trust can be defined as "a psychological state that reflects a set of accumulated presumptions concerning the credibility, integrity and benevolence that the consumer attributes to the brand" [START_REF] Gurviez | Proposition d'une échelle de mesure multidimensionnelle de la confiance dans la marque[END_REF]. Trust rests on a mechanism of attributing characteristics, motives and intentions to the exchange partner, making it possible to evaluate that partner's potential, assuming that behavior will be predictable and conform to promises [START_REF] Doney | An Examination of the Nature of Trust in Buyer-Seller Relationships[END_REF].

The influence of perceptions of social responsibility on consumer trust has already been supported to a degree in the literature (Swaen & Chumpitaz, 2008). Trust appears as an indispensable prerequisite for creating and maintaining long-term relationships between the consumer and the brand [START_REF] Morgan | The Commitment-Trust Theory of Relationship Marketing[END_REF]. Brand trust reduces fear of malevolent or opportunistic behaviors and leads to positive expectations that the brand will continue to fulfill its obligations in the future (Swaen & Chumpitaz, 2008). The importance of trust as a central mediator between brand activities and consumer commitment has also been widely demonstrated in the relational marketing literature [START_REF] Morgan | The Commitment-Trust Theory of Relationship Marketing[END_REF][START_REF] Fournier | Consumers and their brand: Developing relationship theory in consumer research[END_REF][START_REF] Chaudhuri | Product-class effects on brand commitment and brand outcomes: the role of brand trust and brand affect[END_REF].We therefore postulate that:

H1: Brand trust mediates the relationship between the consumer dimension of social responsibility and brand commitment.

1.2.2. The influence of social responsibility on brand commitment from the perspective of social identity and self-categorization theory.

The initiatives of social responsibility that refer to the brand's capacity to reduce its environmental footprint, to participate in the societal and cultural life of its local region of activity, to convey symbolic aspects whereby brands develop their identity [START_REF] Du | Reaping Relational Rewards from Corporate Social Responsibility: The Role of Competitive Positioning[END_REF]. These aspects humanize the brand [START_REF] Fournier | Consumers and their brand: Developing relationship theory in consumer research[END_REF] by revealing identity dimensions such as "values" that appear to be fundamental and durable compared to other classic identity traits [START_REF] Du | Reaping Relational Rewards from Corporate Social Responsibility: The Role of Competitive Positioning[END_REF]. When consumers perceive that they share the same values as those conveyed by the brand, they are likely to identify with it.

Brand identification refers to the "inclusion" of the brand in one's own concept of self [START_REF] Bagozzi | Antecedents and purchase consequences of customer participation in small group brand communities[END_REF]. According to social identity theory [START_REF] Tajfel | An integrative theory of intergroup conflict[END_REF], besides personal identity, the concept of self includes a social identity, that is, a form of psychological attachment to an organization apparent when organizational members attribute certain organizational characteristics to themselves [START_REF] Dutton | Organizational images and member identification[END_REF]. By

identifying themselves with a brand, consumers express and reinforce their own identity [START_REF] Ahearne | Antecedents and Consequences of Customer-Company Identification: Expanding the Role of Relationship Marketing[END_REF] by attributing the brand's positive characteristics to themselves. Brand identification thus provides consumers with psychological benefits by strengthening self-esteem, making them more inclined to develop attitudes and behaviors favorable to the brand [START_REF] Donavan | Environmental influences in corporate brand identification and outcomes[END_REF].

The influence of evaluations of social responsibility on consumers' brand or firm identification has already received a certain amount of support [START_REF] Lichtenstein | The effect of corporate social responsibility on customer donations to corporate supported nonprofits[END_REF][START_REF] Du | Reaping Relational Rewards from Corporate Social Responsibility: The Role of Competitive Positioning[END_REF]. Bhattacharya & Sen (2004) declare that brand identification motivates consumers to commit with the brand's objectives. To feel in harmony with themselves and avoid feeling guilty in their purchases, consumers commit with brands that they perceive as sharing the same values as them. Brand commitment thus results from a feeling of moral obligation to act in accordance with their values by continuing to buy brand products that share these values. The influence of brand identification on consumer commitment has been demonstrated empirically by recent studies [START_REF] Ahearne | Antecedents and Consequences of Customer-Company Identification: Expanding the Role of Relationship Marketing[END_REF]. We thus postulate that:

H2: Brand identification mediates the relationship between the environmental dimension of social responsibility and brand commitment H3: Brand identification mediates the relationship between the philanthropic dimension of social responsibility and brand commitment

The role of consumers' personal norms

It is commonly admitted that consumers' expectations in terms of social responsibility are far from being uniform; they vary from one consumer to another (Bhattacharya & Sen, 2004).

We postulate that these expectations are likely to condition consumer reactions towards the symbolic dimensions of social responsibility.

Research on organizational identification 3 suggests that people are more likely to identify with an organization when they perceive congruence between that organization's identity and their own self-concept [START_REF] Bagozzi | Antecedents and purchase consequences of customer participation in small group brand communities[END_REF]. According to cognitive coherence theory, consumers seek balance and harmony between their beliefs and feelings on one hand and their consumer behavior on the other [START_REF] Mcguire | Some internal psychological factors influencing consumer choice[END_REF]. Consequently, consumers avoid choosing brands that are incompatible or dissonant with their own system of beliefs and values; they rather commit to the brands that are most congruent with their self-concept and their personality [START_REF] Aaker | The Malleable Self: The Role of Self-Expression in Persuasion[END_REF].

We can thus suppose that consumers' reactions to the symbolic dimensions of social responsibility depend on the congruence they perceive between the brand's values as revealed by its environmental and societal commitments, and their own. These perceptions of congruence are likely to vary from one consumer to another depending on consumers' personal norms regarding social responsibility.

Personal norms can be defined as personal expectations based on interiorized values [START_REF] Schwartz | Normative Influences on Altruism[END_REF]. They translate into a feeling of moral obligation to act in accordance with these values [START_REF] Schwartz | Normative Influences on Altruism[END_REF]. This moral obligation generally stems from specific personal norms [START_REF] Grankvist | Values and eco-and fair-trade labelled products[END_REF].

Personal norms, in terms of social responsibility, appear as the principal explanation of individuals' predispositions to support causes [START_REF] Stern | New Environmental Theories: Toward a coherent Theory of environmentally significant behavior[END_REF]. Consumers whose self-concept includes high personal norms of social responsibility will thus perceive greater congruence between themselves and the brand, whether in terms of common attributes or shared prototypes, than consumers whose personal norms of social responsibility are low.

3 For a revue, see Bergami, M., &Bagozzi, R. P. (2000). Self-categorization, affective commitment and group self-esteem as distinct aspects of social identity in the organization.Britsh Journal of Social Psychology, 39, 4, 555-577.

Since identification is at least partly motivated by people's need to maintain a coherent and positive self-image, [START_REF] Dutton | Organizational images and member identification[END_REF], consumers are more likely to identify with a brand when its social responsibility efforts are perceived as corresponding to their own personal norms. They will thus tend to identify and engage with brands that they judge favorably on the symbolic dimensions of social responsibility.

We thus postulate that:

H4 : Consumers' personal norms moderate the indirect link between the environmental dimension of social responsibility and brand commitment via brand identification such that this mediation is stronger (weaker) when respondents' personal norms are high (low).

H5 : Consumers' personal norms moderate the indirect link between the philanthropic dimension of social responsibility and brand commitment via brand identification such that this mediation is stronger (weaker) when respondents' norms are high (low).

Insert Figure 1 about here

Material and Methods

To test our research model, we carried out a survey among regular consumers of national brands4 of organic foods sold through mass retail. Indeed, organic food is one of the fastest growing industries in socially responsible consumption5 . The consumption of organic products is based both on instrumental motives, related to health protection, quality and to more ethical concerns such as preserving the environment and/or philanthropic activities [START_REF] Magnusson | Choice of organic foods is related to perceived consequences for human health and to environmentally friendly behaviour[END_REF].

Moreover, the choice of national brands sold only in supermarkets enabled us to control the effect of the image of the retail brands, specialized stores versus supermarkets 6 ,on the relationship consumers may develop with the stores they frequent. In France, 81% of organic products are today purchased in small and medium sized supermarkets (CSA Agence du Bio, 2016 7 ).

To select the brands likely to be included in our panel, we relied on the results of a national study presenting the 10 most purchased organic foods in France (Kantar, 2012) 8 . A distribution channel accepted to distribute our survey to its virtual community of consumers of organic products "Mieux Vivre avec Auchan" (Live better with Auchan). Auchan referenced a large number of organic food products including the list of national organic food products mentioned above. The questionnaire was sent to about 3541 consumers of the "Mieux Vivre avec Auchan" community.

The first question was about respondents' most regularly purchased brand. By concentrating on "regular" consumers, we could be sure that they had already established a relationship with the brand through their previous behavior and were engaged in purchasing, which is an important prerequisite for brand commitment [START_REF] Morgan | The Commitment-Trust Theory of Relationship Marketing[END_REF][START_REF] Terrasse | L'engagement envers la marque: Proposition d'un modèle théorique et application à la comparaison de la fidélité aux marques nationales et aux marques distributeurs[END_REF]. We then asked respondents to fill in the questionnaire with reference to the brand they bought most often.

We obtained 312 complete responses, representing a response rate of about 9%. Among the responses, we selected only those relative to the brands included in our list of ten national brands. Moreover, we evaluated consumer involvement in their purchase of the brand by using a global measure. Only respondents with a high level of involvement were retained.

6 Specialized shops such as Biocoop have a more positive image than traditional retail distributors like Auchan or Leclerc.

7

French Agency for the development and the promotion of organic farming, (CSA-Consumer Science and Analytics), http://www.agencebio.org/comprendre-le-consommateur-bio 8

For reasons of confidentiality, we cannot present the whole list. For example, this list included brands like "Bjorg", "Jardin Bio'" and "Les 2 vaches".

Strong involvement ensured in-depth treatment of information thus rendering brand consumers' evaluation more pertinent [START_REF] Terrasse | L'engagement envers la marque: Proposition d'un modèle théorique et application à la comparaison de la fidélité aux marques nationales et aux marques distributeurs[END_REF]. The final sample comprised 299 respondents of whom 55% were women with an average age of 37 years.

Instruments of measure

To check for social desirability bias, we measured it with a 10-item scale by [START_REF] Strahan | Short, Homogeneous Versions of the Marlowe-Crowne Social Desirability Scale[END_REF] adapted from the [START_REF] Crowne | A new scale of social desirability independent of psychopathology[END_REF] social desirability scale.

Research on socially responsible consumption has shown that social desirability can influence consumers' responses [START_REF] Roberts | Profiling levels of socially responsible consumer beahaviour: a cluster analytic approach and its implications for marketing[END_REF]. Following the recommendation of DeVellis (1991), we withdrew items strongly correlated with social desirability; in all, 7 items were withdrawn respectively from the scales of identification (3 items withdrawn), commitment (1item withdrawn), personal norms (2items withdrawn) and trust (1 item withdrawn).

Consumer brand identification was measured using the 6 item scale of Escalas & Bettman (2003;2005). After withdrawing items strongly correlated with social desirability, the 3-item scale presented good reliability (α = 0,923).

Consumer brand commitment was measured using the [START_REF] Terrasse | L'engagement envers la marque: Proposition d'un modèle théorique et application à la comparaison de la fidélité aux marques nationales et aux marques distributeurs[END_REF] 9-item scale adapted to the French context of the Gruen& al. (2000) and [START_REF] Bansal | A Three-Component Model of Customer Commitment to Service Providers[END_REF] and scales.

The three dimensions of commitment (normative, affective and continuous) were combined into a single higher order factor. The scale presents a satisfactory reliability coefficient (α =

.891) as do each of its dimensions (α between .858 and.865). We then undertook a second order confirmatory factor analysis to determine the fit validity.The results show an acceptable fit to the data (Χ2 = 83.42, df = 18, CFI = .95, TLI = .955, PAGF = .885, RMSEA = .11) thatis significantly better than the first order model (Χ2 = 390.74, dl = 20, CFI = .744, TLI =

.642, PAGF = .506, RMSEA = .249).

Consumers' brand trust was measured using the [START_REF] Gurviez | Proposition d'une échelle de mesure multidimensionnelle de la confiance dans la marque[END_REF] 8 item scale.

The reliability coefficient for this scale is .931; those of the three dimensions vary between (.899 and .925).We then carried out a second order confirmatory factor analysis to determine the data fit validity of the higher order factor. The results show that it fits the data very well (Χ2 = 6.08, df = 6, CFI = 1, TLI = 1, PAGF = .997, RMSEA = .007), indeed far better than the first order model (Χ2 = 316.03, dl = 9, CFI = .798, TLI = .664, PAGF = .442, RMSEA = .338).

We used the [START_REF] Abid | Une échelle de mesure de la responsabilité sociétale des marques : application aux consommateurs de marques alimentaires biologiques[END_REF] 11 item scale to evaluate respectively the following dimensions: consumer (α =.911); environment (α =.853) and philanthropy (α =.921) of brand social responsibility. Finally, we measured consumers' personal norms using the Harland,

Staats & Wilke (1999) 8 item scale. After eliminating the two items correlated significantly with social desirability, the factor analysis also suggested that we eliminate two other poorly represented items. Finally, the factor analysis on the 4 remaining items enabled us to identify a unidimensional structure that restituted a little over 75% of variance and presented satisfactory reliability (α =.892). All instruments of measure are presented in the appendix.

2.2. Analysis strategy 2.2.1. Test strategy for the mediation hypotheses (H1, H2 and H3)

To test our mediating hypotheses, we used the structural equation method (SEM) to evaluate the mediation since this method makes it possible to evaluate all the relationships simultaneously [START_REF] Zao | Reconsidering Baron and Kenny: Myths and truths about mediation analysis[END_REF]. The SEM allows to test complex mediation models in a single analysis comprising several independent, dependent or mediating variables.

Furthermore, by modeling the concept with latent variables, SEM takes account of errors in the measure of the mediator, which allows to distinguish a "true" indirect effect from one that is an artifact of errors in the variables to be measured [START_REF] Zao | Reconsidering Baron and Kenny: Myths and truths about mediation analysis[END_REF].

For each mediation hypothesis, we evaluate the importance of the indirect effect (a x b) by using a bootstrap procedure. Bootstrapping is a procedure of sample simulation from the initial sample. There is a current consensus in the literature that recognizes the superiority of bootstrapping compared to the Baron and Kenny approach for testing mediation effects [START_REF] Hayes | Introduction to mediation, moderation and conditional process analysis: A regression-based approach[END_REF]. This procedure establishes the existence of a significant indirect effect when the confidence interval excludes 0. To establish the type of mediation (total or partial), we see whether the direct effect (c) remains significant or not. When the mediation is partial (the direct effect is significant), it means that other mediating processes, that have not been measured, are at work. Conversely, when there is total mediation (the direct effect is not significant), we can conclude that this variable alone explains the phenomena at work.

Test strategy for hypotheses of moderate mediation (H4 and H5)

Hypotheses H4 and H5 correspond to a conditional indirect effect, meaning that the moderating variable influences the indirect impact of the independent variable on the dependent variable [START_REF] Hayes | Statistical mediation analysis with a multicategorical independent variable[END_REF]. To test these effects, only simultaneous analysis obtains reliable and robust results [START_REF] Hayes | Introduction to mediation, moderation and conditional process analysis: A regression-based approach[END_REF]. There again, the bootstrap procedure is recommended in so far as it obtains a more precise estimate that does not rely on normality hypotheses, rarely verified for these effects [START_REF] Edwards | Methods for integrating moderation and mediation: A general analytical framework using moderated Path Analysis[END_REF]. To test these effects, we use the macro PROCESS developed by [START_REF] Hayes | Introduction to mediation, moderation and conditional process analysis: A regression-based approach[END_REF] that tests them directly using a bootstrap procedure using SPSS. The macro PROCESS generates a moderate mediation index associated to a confidence interval from the bootstrap.

Results

Table 1 below presents the standard deviations and correlations

Insert Table 1 about Here

Results of the test of the mediation hypotheses (H1, H2& H3)

The results of the tests of the mediating effects show that hypothesis 1 relative to the mediating role of trust between the consumer dimension and commitment is confirmed. The direct effect of the consumer dimension on commitment is statistically insignificant. We can thus conclude that the mediation is total.

Our results also confirm hypotheses 2 and 3 relative to the mediating role of identification between the environment and philanthropy dimensions of social responsibility and brand commitment. There again, the direct effects of the environment and philanthropy dimensions on commitment appear non-significant, thus confirming the total mediations. The tables below summarize the results of the test of mediating effects.

Insert Table 2 & 3 about Here

The global model including the three mediating effects explains almost 70% of the variance of brand commitment. It presents a good fit.

Insert Table 4 about Here

Furthermore, the inclusion of the mediating effects in the model significantly increases the model's fit to the data, as shown in the following table.

Insert Table 5 about Here

Results of the test of the hypotheses of moderate mediation (H4, H5)

Our results are not able to validate hypothesis H4 relative to the conditional effect of personal norms on the indirect effect of the environment dimension on brand commitment.

Although the indirect effects appear significant with different levels of moderator, the index of moderate mediation presents a confidence interval that includes 0. Tables 6 and7 below present these results in detail.

Insert Table 6&7 about Here

Our results confirm to the conditional effect of personal norms on the indirect effect of the philanthropy dimension on brand commitment through identification, validating hypothesis H5. The tables below present the results of the test of indirect effects at different values of the moderator as well as the index of moderate mediation. The index of moderate mediation presents a confidence interval that excludes 0 with respect to the philanthropy dimension. We can deduce that personal norms condition the strength of the indirect effect of the philanthropic dimension on brand commitment through identification. The indirect effect appears significantly stronger when consumers' personal norms are high, as we hypothesized.

Tables 8 and9 present these results in detail.

Insert Table 8&9 about Here

Discussion

Our research is part of the stream dealing with the beneficial effect of social responsibility on the consumer-brand relationship. It extends previous studies by using a dual theoretical framework to explain the processes involved in different associations of social responsibility influence on consumer brand commitment.

Notably, this research confirms the need to consider social responsibility with its different dimensions. By distinguishing the instrumental and symbolic dimensions of brand social responsibility, our study identifies the mechanisms through which the different associations of social responsibility can impact consumers. It seems necessary to go beyond the holistic view of social responsibility in order to understand this effect.

Our results show that the mediating processes at work between the different associations of social responsibility and consumer brand commitment depend on the type of benefits that these associations convey (utilitarian or symbolic). Our research also reconciles two theoretical perspectives that have often been considered separately to explain the benefits of social responsibility in the consumer-brand relationship.

Previous studies on the beneficial effects of company social responsibility have highlighted the crucial role of trust, presented as the fundamental mechanism of consumer commitment and loyalty (Swaen & Chumpitaz, 2008) in a context where skepticism and suspicion of firm behavior is rife [START_REF] Bibb | Trust Matters: For organizational and Personal Success[END_REF]. The brand's capacity to protect consumers' health and provide clear information on product ingredients appears to be an attribute of brand reputation. It sends out positive signals [START_REF] Spence | Product selection, fixed costs and monopolistic competition[END_REF] regarding the quality of products and offerings [START_REF] Brown | The Company and the product: corporate associations and consumer product responses[END_REF] as well as firms' capacity to watch over consumers' well-being. The consumer dimension of social responsibility thus influences the consumer-brand relationship according to a classic relational process, based on reciprocity and respect of promises in the exchange. Trust plays a central role [START_REF] Morgan | The Commitment-Trust Theory of Relationship Marketing[END_REF] as a mechanism of consumer commitment concerning the consumer dimension of social responsibility.

However, our research shows that trust is not the only lever of consumer commitment. In fact, consumption is not related only to a functional end but also fulfills a role of expression (Baudillard, 1970). The symbolic aspects of social responsibility provide consumers with means for self-fulfillment and expression of their values and ideals through consumption [START_REF] Ahearne | Antecedents and Consequences of Customer-Company Identification: Expanding the Role of Relationship Marketing[END_REF]. This attractiveness depends on the brand's capacity to satisfy consumers' identification needs (Bhattacharya & Sen, 2004).

More and more research is focused on the way consumers identify themselves with a brand, as well as the consequences of this identification on consumer-brand relationships [START_REF] Lichtenstein | The effect of corporate social responsibility on customer donations to corporate supported nonprofits[END_REF]). Our research extends previous studies by showing that brand identification is the fundamental process of consumer commitment regarding the symbolic dimensions of social responsibility.

Furthermore, our results confirm the role of personal norms that condition the influence of the philanthropic dimension on consumer commitment through identification. It thus seems that the influence of the philanthropic dimension on consumer identification and commitment is greater for individuals with strong personal norms.

The concept of personal norms has been widely mobilized in the literature of socially responsible consumption. It sheds light on why certain consumers are more inclined than others to adopt socially responsible consumer behavior. The moderating effect of personal norms is implicit in previous research on consumer reactions to CSR. This strongly depends on the personal support of the consumer for the social responsibility initiatives conveyed by the firm [START_REF] Sen | Does doing good always lead to doing better? Consumer reactions to Corporate Social Responsibility[END_REF].

Several authors have also evoked the role of consumer affinity with the cause defended [START_REF] Drumwright | Company advertising with a social dimension: The role of noneconomic criteria[END_REF], the importance of the question for the self or personal pertinence [START_REF] Creyer | The impact of corporate behavior on perceived product value[END_REF]. However, the research on the beneficial effects of social responsibility on the consumer-brand relationship has never taken account of the role of personal norms. The concept of personal norms, resulting from the work of [START_REF] Schwartz | Normative Influences on Altruism[END_REF], clarifies this effect by providing a theoretical framework that highlights this effect of individual dispositions.

However, the influence of personal norms on the effects of the environmental dimension was not supported by our analysis. This result can probably be explained by our field of study, namely organic products. Indeed, according to a study conducted in 2015 9 , preserving the environment is the second reason why people purchase organic products, just behind maintaining health. We can imagine that regular consumers of organic food products show 9 13 th barometer of the consumption and perception of organic products in France carried out in 2015 by the "French agency for the development and the promotion of the organic farming" and supported by CSA (consumer science and analytics) http://www.agencebio.org/sites/default/files/upload/documents/4_Chiffres/BarometreConso/barometre_agence_ bio_public.pdf strong individual predispositions to take account of environmental considerations in their consumption.

Finally, our research underlines the major role brands play in firms' social responsibility policy. Social responsibility is often approached at the firm level without taking account of the role of brands. This empirical research validates the suggestions of previous research holding that social responsibility can be applied to brands [START_REF] Lindgreen | Corporate social responsibility: An empirical investigation of U.S. organizations[END_REF][START_REF] Lindgreen | Corporate social responsibility brand leadership: a multiple case study[END_REF].

Managerial implications

Our research attests to the central role played by the marketing function in setting up a social responsibility policy [START_REF] Maignan | A stakeholder model for implementing social responsibility in Marketing[END_REF], particularly in terms of brand policy [START_REF] Lindgreen | Corporate social responsibility: An empirical investigation of U.S. organizations[END_REF]. Social responsibility applied to brands secures brand value in terms of durability [START_REF] Hildebrand | Corporate Social Responsibility: A Corporate Marketing Perspective[END_REF]. Consumers take account of social responsibility criteria while evaluating the brands they buy. Studying social responsibility at brand level is even more necessary in so far as a large number of firms adopt mixed or multi-brand strategies [START_REF] Lindgreen | Corporate social responsibility brand leadership: a multiple case study[END_REF].

Our research shows that commitment mechanisms differ depending on the type of social responsibility initiatives highlighted by brands and their symbolic or utilitarian nature for consumers. Understanding these mechanisms helps to guide managerial decisions in terms of the design and promotion of various socially responsible actions that should encourage consumer commitment and loyalty. Brands positioned on attributes such as product safety and quality should communicate about quality certificates to develop trust and overcome widespread consumer skepticism [START_REF] Bibb | Trust Matters: For organizational and Personal Success[END_REF].

In terms of brands' commitment for social and environmental benefits, the objective should be to increase the attractiveness of brand identity for targeted consumers. For this, the characteristics of the targeted consumers should be studied to make sure that the engagements made by brands are compatible with consumers' values in order to encourage brand identification. The communication should be thought of in terms of activating personal norms.

Following Schwartz' model, personal norms influence behavior under two conditions (1) when people think that their action will have real consequences on others' well-being;

(2) when they can claim responsibility for this action to themselves. To activate consumers' personal norms and thus encourage their commitment, marketing managers should pull these two levers in their communication.

Like any study, this one also presents a certain number of limitations that need to be underlined.

First of all, we chose to test our conceptual model on consumers of only one product category, namely, organic food products. This choice certainly strengthened the internal validity of the research. However, it opens the way to a certain number of questions as to the possible generalization of the results to other product categories. The external validity of this study should thus be reinforced by studying other fields.

Next, we tested out model on consumers of national brands. Our results converge with those of researchers in Europe and the United States [START_REF] Maignan | A stakeholder model for implementing social responsibility in Marketing[END_REF][START_REF] Lindgreen | Corporate social responsibility: An empirical investigation of U.S. organizations[END_REF]. It seems therefore that social responsibility can contribute in a similar way to national brands, as well as to a global, transnational brand strategy [START_REF] Pestre | Local, global and transnational CSR strategies in the multinational corporation[END_REF], or a mixed "glocal 10 " strategy [START_REF] Persais | La gestion de la RSE dans un contexte international : vers une «glocalisation » des pratiques ? Une étude de cas d'entreprises françaises implantées au Mexique[END_REF]. Further research should be conducted in order to test the model on consumers of multinational brands such as "Björk"

or "Patagonia".

The third limit of this research is related to time. Like all studies in the relational stream, our research is transversal and does not take account of temporal aspects in the construction of 10 While previous researches set local strategies against global strategies, Persais (2010) suggests combining both, benefiting from advantages of local and global CSR strategies. a relationship. A longitudinal study thus seems appropriate to account for the role of interactions taking place between the consumer and the brand studied.

Regarding future perspectives, a promising avenue of research is the reaction to inconsistencies between engagements that firms convey and their actual practices when these are discovered to be divergent. In fact, we could imagine that consumer reactions to allegations of irresponsibility will depend on the utilitarian or symbolic nature of their commitments. Furthermore, mechanisms for reinstating trust may differ from levers of reconstructing the attractiveness of the brand identity. These research avenues appear particularly promising.

Finally, other variables may modify the relation between consumers and brands, such as the degree of consumer skepticism or the relative weight of social responsibility initiatives during purchase decisions. Other variables, outside the scope of the present study but identified in the literature on ethical and equitable consumption might also be integrated into future work. We can thus mention consumers' tendency towards collectivism or individualism or their sensitivity to products' origin. Cultural and/or national differences are highly likely to play an important role in evaluating brand social responsibility and its influence on consumers. Social desirability I'm always willing to admit it when I make a mistake I always try to practice what I preach I never resent being asked to return a favor I have never been irked when people expressed ideas very different from my own I have never deliberately said something that hurt someone's feelings I like to gossip at times There have been occasions when I took advantage of someone I sometimes try to get rather than forgive and forget At times I have really insisted on having things my own way There have been occasions when I felt like smashing things

Figure

  Figure 1: Research Model

Table 1 :

 1 Means, Standard Deviations and Inter-correlations ** p< 0,01 ; M = Mean, SD = Standard deviation

Table 2 :

 2 Test of mediation model

			M1		M2	Y
			Trust	Identification	Commitment
			t-test		t-test		t-test
	Consumer SR	.565	9.544***			.053	.609 (ns)
	Environmental SR			.449	8.383***	.097	1.415 (ns)
	Philanthropic SR			.411	7.692***	-.110	-1.627 (ns)
	Trust					.650	7.217***
	Identification					.283	3.472***
	R²		.32		.50	.70
	Note: entries are standardized regression weights *p <.05; ** p<.001; *** p<.0001

Table 3 :

 3 Test of mediation paths : confidence interval of 95% for the indirect effect by bootstrap corrected for bias b:confidence interval of 95% for the direct effect by bootstrap corrected for bias

			Confidence			Confidence
			Interval				Interval
	Hypothesis	Indirect	Lower	Upper	Direct	Lower	Upper Result
		effect				effect
	H1	.368	.246	.516		.053	-.131		.229	Full
									mediation
	H2	.127	.046	.234		.118	-.022		.297	Full
									mediation
	H3	.116	.034	.223		-.110	-.273		.038	Full
									mediation
	Note: aTable 4: Model fit
		χ²	do	χ²n	AGFI CFI	TLI	RMSEA

Table 6 :

 6 Model of conditional indirect effect of environment on commitment through identification

	M		Y
	(Identification)	(Commitment)
	Coef.	T-test	Coef.	T-test

Table 7 :

 7 Test of Indirect conditional effect of environment on commitment through identification at different levels of personal norms(H4)

	Moderator W	Indirect effect	Confidence Interval
	(Personal norms)		Lower Bound	Upper Bound
	-1	.181	.121	.255
	0	.182	.134	.240
	1	.183	.126	.252
	Index of moderated mediation	.001	-.037	.036

Table 8 :

 8 Model of conditional indirect effect of philanthropy on commitment through identificationTableau 9: Test of indirect conditional effect of philanthropy on commitment through identification at different levels of personal norms(H5) when I don't buy organic and environment friendly brands .786 People like me should do everything they can to buy organic and environment friendly brands .905

	M	Y

Morgan & Hunt (1994, p.22) define relationship marketing as "all marketing activities directed toward establishing, developing and maintaining successful relational exchanges".

We eliminated retailers' brands

Organic food reach a market penetration rate of 96.3% in 2013 "Kantar World panel".

.922 I'm ready ( e) to do everything possible to continue using this brand .783 Brand identification I use this brand to communicate who I am to other people.

.926 I consider this brand to be "me" (it reflects who I consider myself to be or the way that I want to present myself to others).

.914 I think this brand helps me become the type of person I want to be.

.


Mudra Khare

Dhiraj Sinha

Rita Zgheib

Ibrahima Sory

Jean-Christophe Traore

Didier Lagier

Pierre-Edouard Raoult

Fournier

Sory Ibrahima Traore

Jean-Christophe Lagier

Didier Raoult

Fournier Pierre

Pierre-Edouard Fournier
email: pierre-edouard.fournier@univ-amu.fr

Genomic description and characterization of Nigeribacterium massiliense gen. nov., sp. nov., isolated from the human gut

Keywords: sp. nov Culturomics, taxonomy, genomics, taxono-genomics, Nigeribacterium massiliense

scientifiques de niveau recherche, publiés ou non, émanant des établissements d'enseignement et de recherche français ou étrangers, des laboratoires publics ou privés.

Introduction

The human microbiome consists of a diversified array of microorganisms that include predominantly bacteria, viruses and fungi, but also archaea and protozoa. The main human habitats of microorganisms are the gastrointestinal system, oral cavity, genital organs, respiratory tract and skin [START_REF] Lloyd-Price | The healthy human microbiome[END_REF]. The microbiota cell count in a human being is estimated to be ~10 13 -10 14 , with around 10:1 microbial cells to human cells [START_REF] Sender | Are We Really Vastly Outnumbered? Revisiting the Ratio of Bacterial to Host Cells in Humans[END_REF]. Most of these microorganisms inhabit the colon [START_REF] Sender | Revised Estimates for the Number of Human and Bacteria Cells in the Body[END_REF], followed by the oral, skin and urogenital tracts [START_REF] Gupta | Geography, ethnicity or subsistence-specific variations in human microbiome composition and diversity[END_REF]. The gastrointestinal microbiota is mainly composed of bacteria that belong to the phyla Firmicutes, Bacteroidetes and Actinobacteria [START_REF] Tap | Towards the human intestinal microbiota phylogenetic core[END_REF]. Currently, for the study of the human gut microbiota, stool samples are the most representative [START_REF] Bilen | Taxonogenomics description of Parabacteroides timonensis sp. nov. isolated from a human stool sample[END_REF]. In the past two decades, metagenomics has enabled exploring the composition of the gut microbiota very rapidy as well as extensively but this strategy has several limitations. The DNA extraction efficiency may vary among microorganisms; the results of these analyses are only snapshots of the real composition of the gut microbiota; this technique does not discriminate among living and dead microorganisms and does not allow appreciating their metabolic activity; and the interpretation of sequences allocated to unknown species is difficult and does not allow creating new species [START_REF] Wang | Application of metagenomics in the human gut microbiome[END_REF]. Because of these limitations and the need to obtain isolates from human-associated microorganisms, culturomics was developed. Culturomics is a strategy based on the diversification of culture conditions to mimic as much as possible each bacterial species' natural growth conditions [START_REF] Lagier | Culturing the human microbiota and culturomics[END_REF]. To date, culturomics, combined to taxono-genomics [START_REF] Ramasamy | A polyphasic strategy incorporating genomic data for the taxonomic description of novel bacterial species[END_REF], has enabled the characterization and creation of more than 500 human-associated bacterial species [START_REF] Lagier | Culturing the human microbiota and culturomics[END_REF]. Taxono-genomics was developed to incorporate into the taxonomic description of new bacterial taxa their peptidic profile using MALDI-TOF MS [START_REF] Seng | Ongoing revolution in bacteriology: Routine identification of bacteria by matrix-assisted laser desorption ionization time-offlight mass spectrometry[END_REF] and their genomic characteristics [START_REF] Bilen | Taxonogenomics description of Parabacteroides timonensis sp. nov. isolated from a human stool sample[END_REF][START_REF] Fournier | From culturomics to taxonomogenomics: A need to change the taxonomy of prokaryotes in clinical microbiology[END_REF][START_REF] Lagier | The rebirth of culture in microbiology through the example of culturomics to study human gut microbiota[END_REF].

Using taxono-genomics, we herein report the description of strain Marseille-P1302 that was cultivated from the stool of a child suffering from Kwashiorkor. On the basis of its characteristics, we conclude that strain Marseille-P1302 belongs to a new species in a new genus for which we propose the name Nigeribacterium massiliense gen. nov., sp. nov.

Materials and Methods

Sample collection

The bacterial strain was isolated in 2014 from the stool of a 2-year-old child suffering from Kwashiorkor, a form of acute severe malnutrition in Niger. The patient was not taking any antibiotics at the time of sampling and the fecal sample was frozen at -80 C and was transported to Marseille, France, for diagnostic purposes. The child's parents provided an informed and signed consent. The study was validated by the ethics committee of the Institute Federatif de Recherche (Marseille, France) IFR48 under agreement number 09-022. All methods were performed in accordance with the relevant guidelines and regulations.

Strain isolation and identification

Culturomics was used to explore the microbial diversity of the child's gut microbiota [START_REF] Lagier | Microbial culturomics: Paradigm shift in the human gut microbiome study[END_REF]. One g of stool was pre-incubated at 37°C in anaerobic blood culture bottle (bioMérieux, Marcy l'Etoile, France) for 2 days. Then, the culture medium was spread on a 5% sheep bloodenriched Columbia agar plate (bioMérieux) and incubated at 37°C in aerobic condition.

Identification of the obtained colonies was attempted using MALDI-TOF-MS as previously described [START_REF] Seng | Ongoing revolution in bacteriology: Routine identification of bacteria by matrix-assisted laser desorption ionization time-offlight mass spectrometry[END_REF] using a Microflex spectrometer (Bruker Daltonics, Leipzig, Germany). The resultant spectrum was compared to the spectra contained in the Bruker database. If the score was >2.0, the strain was considered as identified at the species level, but if the score was <2 the strain was considered as unidentified. Strains considered as unidentified were further characterized by 16S rRNA gene amplification and sequencing using the Fd1 -rP2 universal primers [START_REF] Drancourt | 16S ribosomal DNA sequence analysis of a large collection of environmental and clinical unidentifiable bacterial isolates[END_REF]. If the 16S rRNA sequence similarity with the phylogenetically closest species with a validly published name was lower than 98.65% or 95%, the tested strain was considered as belonging to a new species and/or a new genus, respectively [START_REF] Erko | Taxonomic parameters revisited: tarnished gold standards[END_REF][START_REF] Kim | Towards a taxonomic coherence between average nucleotide identity and 16S rRNA gene sequence similarity for species demarcation of prokaryotes[END_REF].

Growth condition and Bacterial Morphology

The optimal growth conditions of strain Marseille-P1302 were tested on 5% sheep bloodenriched Columbia agar (bioMérieux) at various temperatures (20, 25, 28, 37, 42 and 56°C), and also in various atmospheres (aerobic, anaerobic and microaerophilic). Gram-stain, motility, sporulation, catalase and oxidase tests were conducted according to the previously described protocols [START_REF] Wormser | Manual of Clinical Microbiology[END_REF]. For transmission electron microscopy (TEM), a colony was collected from agar and immersed into a 2.5% glutaraldehyde fixative solution. Then, a drop of a suspension was directly deposited on a poly-L-lysine coated microscopic slide for 5 minutes and treated with 1% phosphotungstic acid aqueous solution (pH 2.0) for 3 minutes to increase the TEM image contrast. The slide was gently washed in water, air dried and examined in a tabletop TEM (Tecnai G20). Acquisitions are shown in Figure 2.

Isolation and sequencing of genomic DNA

After the lysozyme incubation at 37℃ for 30 minutes Genomic DNA (gDNA) of strain Marseille-P1302 was extracted using the EZ1 biorobot (Qiagen, Hilden, Germany) with the EZ1 DNA Tissue kit. The elution volume was 50µL. The genomic DNA was quantified by a Qubit assay with the high sensitivity kit (Life technologies, Carlsbad, CA, USA) at 105.7ng/µl.

For genome sequencing, a mate-pair library was prepared with 1.5µg of gDNA using the Nextera mate-pair Illumina guide (Illumina, San Diego, CA, USA). The gDNA was simultaneously fragmented and tagged with mate-pair junction adapters. The pattern of fragmentation was validated on an Agilent 2100 BioAnalyzer (Agilent Technologies, Santa Clara, CA, USA) with a DNA 7500 labchip kit. The fragmented DNA ranged in size from 1.5 kb up to 11 kb with an optimal size at 6.662 kb. No size selection was performed and 376.1 ng of tagmented fragments were circularized. The circularized DNA was mechanically sheared to small fragments with an optimal of 529 bp on the Covaris device S2 in T6 tubes (Covaris, Woburn, MA, USA). The library profile was visualized on a High Sensitivity Bioanalyzer LabChip kit (Agilent Technologies) and the final concentration library was measured at 4.162 nmol/l. After a denaturation step and dilution at 15pM, the library was loaded onto the flow cell and then onto a MiSeq sequencer (Illumina). Automated cluster generation and sequencing run were performed in a single 39-hour run in a 2x251-bp format. Total information of 8.9 Gb was obtained from a 1023 K/mm 2 cluster density with a cluster passing quality control filters of 89.6 %. The 1 178 689 reads were filtered according to the read qualities. The sequence assembly protocol was followed as previously described [START_REF] Anani | Colibacter massiliensis gen. nov. sp. nov., a novel Gram-stain-positive anaerobic diplococcal bacterium, isolated from the human left colon[END_REF]

Phenotypic and Biochemical features

Biochemical properties of strain Marseille-P1302 were tested using API ZYM and API 50CH strips (bioMérieux) according to the manufacturer's instructions. Minimal inhibitory concentrations (MICs) of amoxicillin, vancomycin, ciprofloxacin, trimethoprimsulfamethoxazole, rifampin erythromycin and colistin were determined according to EUCAST recommendations using E-test gradient strips (bioMerieux) [START_REF] Alcock | CARD 2020: Antibiotic resistome surveillance with the comprehensive antibiotic resistance database[END_REF][START_REF] Macià | Antimicrobial susceptibility testing in biofilm-growing bacteria[END_REF].

Cellular fatty acid methyl ester (FAME) analysis was performed by Gas Chromatography/ Mass Spectrometry (GC/MS). Two samples were prepared with approximately 80 mg of bacterial biomass per tube harvested from several culture plates. Fatty acid methyl esters were prepared as described by Sasser in 2006 [START_REF] Sasser | Identification of Bacteria by Gas Chromatography of Cellular Fatty Acids[END_REF]. GC/MS analyses were carried out as described before [START_REF] Dione | Genome sequence and description of Anaerosalibacter massiliensis sp[END_REF]. Briefly, FAMEs were separated using an Elite 5-MS column and monitored by mass spectrometry (Clarus 500 -SQ 8 S, Perkin Elmer, Courtaboeuf, France). Spectral database search was performed using MS Search 2.0 operated with the Standard Reference Database 1A (NIST, Gaithersburg, USA) and the FAMEs mass spectral database (Wiley, Chichester, UK).

Genome annotation and comparison

The whole genome as well as 16S rRNA sequences (accession numbers CCYM00000000.1 and LK985392.1, respectively) of strain Marseille-P1302 were submitted to GenBank (www.ncbi.nlm.nih.gov). The 16S rRNA sequence from strain Marseille-P1302 was compared to those of other bacteria present in GenBank using BLASTn. Since for further analysis we have used whole genome sequences for the comparative analysis and these were not available for Brevilactibacter sinopodophylli strain TEYR-7 T and Propionicicella superfundia strain BL-10 T . 16S rRNA sequence extracted from Brevilactibacter sinopodophylli strain KCTC 33808 T and Propionicicella superfundia strain DSM 22317 T , respectively, were also used in the 16S phylogenetic analysis. Strain KCTC 33808 is also a type strain of Brevilactibacter sinopodophylli, while strain DSM 22317 is type strain of Propionicicella superfundia. A complete 16S rRNA-based phylogenetic tree was constructed by using the MEGA 7 software [START_REF] Kumar | MEGA X: Molecular evolutionary genetics analysis across computing platforms[END_REF].

Genome annotation of strain Marseille-P1302 was obtained with the help of the PROKKA pipeline that includes several tools such as Aragorn, Barrnap and Prodigal to identify tRNAs, tmRNAs, rRNAs and coding sequences (CDS), respectively [START_REF] Seemann | Prokka: Rapid prokaryotic genome annotation[END_REF]. The predicted protein sequences were compared to the Clusters of Orthologous Groups (COGs) database using BLASTP (E value of 1e-03, coverage 0.7 and identity percent 30%). For the identification of antimicrobial resistance markers, plasmids and virulence genes, the contigs were screened using CARD [START_REF] Seemann | Prokka: Rapid prokaryotic genome annotation[END_REF][START_REF] Jia | CARD 2017: Expansion and model-centric curation of the comprehensive antibiotic resistance database[END_REF], Resfinder [START_REF] Prasertsee | Core genome sequence analysis to characterize Salmonella enterica serovar Rissen ST469 from a swine production chain[END_REF], ARG-ANNOT [START_REF] Brambila-Tapia | Identification of beta-Lactamases and beta-Lactam-Related Proteins in Human Pathogenic Bacteria using a Computational Search Approach[END_REF], PlasmidFinder [START_REF] Carattoli | In Silico detection and typing of plasmids using plasmidfinder and plasmid multilocus sequence typing[END_REF] and VFDB [START_REF] Chen | VFDB: A reference database for bacterial virulence factors[END_REF], respectively, in the ABRicate pipeline (https://github.com/tseemann/abricate) [START_REF] Arnott | 5],12: I:-sequence type 34, New South Wales, Australia[END_REF] with a minimum identity of 60% and coverage of 20%. For the evaluation of sequence similarity at the genomic level between strain Marseille-P1302 and closely related species, we used the Genome-to-Genome Distance Calculator (GGDC) web server (http://ggdc.dsmz.de) to calculate digital DNA-DNA hybridization values values (dDDH) [START_REF] Meier-Kolthoff | Genome sequence-based species delimitation with confidence intervals and improved distance functions[END_REF][START_REF] Meier-Kolthoff | When should a DDH experiment be mandatory in microbial taxonomy?[END_REF]. The recommended formula 2 was considered for the interpretation of results. Moreover we also used the GGDC for the determination of the G+C content to compare the tested strains. The average nucleotide identity at the genome level was also estimated using the OAU tool that uses USEARCH algorithm for fast similarity search [START_REF] Yoon | A large-scale evaluation of algorithms to calculate average nucleotide identity[END_REF][START_REF] Edgar | Search and clustering orders of magnitude faster than BLAST[END_REF]. Morpheus software used for creating the heat map for ANI values (Morpheus, https://software.broadinstitute.org/morpheus/). The GView tool [START_REF] Petkau | Interactive microbial genome visualization with GView[END_REF] was used for the visualisation of genomic features.

Results and Discussion

Strain identification and phylogenetic analysis

Strain Marseille-P1302 was isolated in 2014 and its growth was observed on 5% sheep bloodenriched Columbia agar at 37°C (bioMérieux) in aerobic atmosphere.

The MALDI-TOFF-MS spectrum of the novel isolate obtained a score <1.9, suggesting that the corresponding species was not represented in the reference database, and could belong to a new species. The spectrum was added to our database. The highest 16S rRNA sequence identity of strain Marseille-P1302 was 94.97% with Brevilactibacter flavus strain VG341 T (MG799849. [START_REF] Lloyd-Price | The healthy human microbiome[END_REF] following to this 93.41% and 93.37% with Brevilactibacter sinopodophylli strains KCTC 33808 T and TEYR-7 T (NR_158077.1), 93% and 92.7% with Propionicicella superfundia strains DSM 22317 T and BL-10 T (NR-043609.1), 91.01% with Microlunatus phosphovorus strain NM-1 T (NR_074642.2) and 93.03% with Propioniciclava tarda strain WR061 T (NR_112669.1) (Figure 1). As these values are lower than the 95% threshold for delineating a new genus as suggested by Stackebrandt and Ebers [START_REF] Erko | Taxonomic parameters revisited: tarnished gold standards[END_REF], strain Marseille-P1302 was considered as representative of a putatively new genus within the family Propionibacteriaceae in the phylum Actinobacteria.

Phenotypic and chemotaxonomic features

The growth of strain Marseille-P1302, was observed on 5% sheep blood-enriched Columbia agar at 37°C (bioMerieux). Optimal growth of strain Marseille P1302 was obtained in aerobic atmosphere, but weak growth was also observed in microaerophillic and anaerobic conditions.

The colonies of strain Marseille-P1302 were β-haemolytic. On blood-enriched agar, colonies were smooth and transparent, with a diameter of 0.2-0.4mm. Bacterial cells were non motile, non sporulating and coccoid with a diameter of 0.8-1.0µm (Figure 2). Bacterial Cells were Gram-positive (Figure 3) and exhibited catalase but no oxidase activity (Table1). An API ZYM strip (bioMérieux) showed the presence of leucine arylamidase, valine arylamidase, naphthol-AS-B1-phosphohydrolase, β-galactosidase, α-glucosidase and βglucosidase activities whereas negative reactions were obtained for alkaline phosphatase, esterase (C4), esterase lipase (C8), lipase (C14), cystine arylamidase, trypsin, α-chymotrypsin, acid phosphatase, α-galactosidase, β-glucuronidase, N-acetyl-β-glucosaminidase, αmannosidase and α-fucosidase. By comparison with compared species, strain Marseille-P1302 differed in a combination of alkaline phosphatase, esterase (C4), esterase lipase (C8) and Nacetyl-β-glucosaminidase activities. The phenotypic comparison with other related strains is detailed in Table 1 [START_REF] Zhang | Amycolatopsis albispora sp. Nov., isolated from deep-sea sediment[END_REF][START_REF] Lee | Microbacterium mangrovi sp. nov., an amylolytic actinobacterium isolated from mangrove forest soil[END_REF][START_REF] Dekio | Proposal of new combination, cutibacterium acnes subsp. Elongatum comb. nov., and emended descriptions of the genus cutibacterium, cutibacterium acnes subsp. acnes and cutibacterium acnes subsp. defendens[END_REF][START_REF] Ramasamy | Genome sequence and description of Aeromicrobium massiliense sp[END_REF][START_REF] Tamura | Luteococcus japonicus gen. nov., sp. nov., a new gram-positive coccus with LL-diaminopimelic acid in the cell wall[END_REF][START_REF] Udwary | Genome sequencing reveals complex secondary metabolome in the marine actinomycete Salinispora tropica[END_REF][START_REF] Lee | Tessaracoccus flavescens sp. nov., isolated from marine sediment[END_REF][START_REF] Zhang | Nocardioides gilvus sp. nov., isolated from Namtso Lake. Antonie van Leeuwenhoek[END_REF][START_REF] Nakamura | Microlunatus phosphovorus gen. nov., sp. nov., a new gram-positive polyphosphate-accumulating bacterium isolated from activated sludge[END_REF][START_REF] Wenning | Brevilactibacter flavus gen. Nov., sp. nov., a novel bacterium of the family propionibacteriaceae isolated from raw milk and dairy products and reclassification of propioniciclava sinopodophylli as brevilactibacter sinopodophylli comb[END_REF][START_REF] Bae | C, energy production and conversion; E, amino acid transport and metabolism; F, nucleotide transport and metabolism; G, carbohydrate transport and metabolism; H, coenzyme transport and metabolism; I, lipid transport and metabolism; P, inorganic ion transport and metabolism; Q, secondary metabolites biosynthesis, transport and catabolism; X, mobilome: prophages, transposons; A, RNA processing and modification; B, chromatin structure and dynamics; J, translation, ribosomal structure and biogenesis; K, transcription; L, replication, recombination and repair; D, cell cycle control, cell division, chromosomal partitioning; M, cell wall/membrane/envelope biogenesis; N, cell motility; O, posttranslational modification, protein turnover, chaperones; T, signal transduction mechanisms; U, intracellular trafficking, secretion, and vesicular transport; V, defense mechanisms; W, extracellular structures; Z, cytoskeleton; R, general function predicted only; S, function unknown[END_REF].

Strain Marseille-P1302 was resistant to vancomycin, ciprofloxacin, erythromycin and trimethoprim-sulphamethoxazole but was susceptible to amoxicillin, rifampin and colistin. FAME analysis revealed that the major cellular fatty acids found in strain Marseille-P1302 was 12-methyl-tetradecanoic acid (89%), followed by 12-methyl-Tridecanoic acid (4%) and Hexadecanoic acid (2%). These fatty acids were principally saturated structures, half of them presenting branches. Minor amounts of unsaturated fatty acids were also detected (Table 2).

Genomic characteristics and comparison

The genome size of strain Marseille-P1302 is 2,934,258-bp long with a 70.38% G+C content.

A total of 2704 coding sequences, 9 rRNAs and 46 tRNAs were identified. The gene distribution in functional categories is presented in Table 3. Nine antibiotic resistance genes were detected, with identities ranging from 66.81% to 82.09%. According to CARD (comprehensive antibiotic resistance database) gene Streptomyces_cinnamoneus_EF-Tu_mutants_conferring_resistance_to_elfamycin showing maximum coverage and identity percentage of 99% and 87.02% respectively, and it is responsible for the elfamycin resistance.

While the other database such as Resfinder and ARG-ANNOT showing otr(A)_1 and (Tet)otrA respectively with very low coverage value of 15.06% and identity percentage of 74.03% . Both the genes are responsible for the oxytetracycline and tetracycline resistance. The VFDB database showing the eight virulence genes among the only one gene relA has maximum coverage with identity of 68.03% and related to bacterial stringent response and involved in quorum sensing. Details of these resistance genes and virulence factors are presented in Table 4. No plasmid was detected.

Comparison between the draft genomes of strain Marseille-P1302 and other phylogenetically closely related strains is presented in Table 5. The genome size (Mb) and number of proteincoding genes of strain Marseille-P1302 (2.93 and 2704 respectively) is smaller than those of all compared strains except Cutibacterium acnes strain ATCC 6919 T (2.49 and 2308, respectively). Similarly, the lowest G+C content value was recorded for Cutibacterium acnes strain ATCC 6919 T (60%) while strain Marseille-P1302 had a value of 70.38%, which is within the range of all other strains (67% for Luteococcus japonicus strain DSM 10546 T to 77.32% for Aeromicrobium massiliense strain JC 14 T ). The distribution of genes into COG categories is shown in Figure 5. Strain Marseille-P1302 showing 2168 functional orthologous genes distributed in different categories of COG while the lowest number of orthologous genes (1785) found in strain Cutibacterium acnes strain ATCC 6919 T and the highest number of orthologous genes (6313) were carried by Amycolatopsis albispora strain WP1 T . The dDDH values of strain Marseille-P1302 with other closely related strains ranged from 17.9% to 22.8% (Table 6). The lowest value was recorded with Microbacterium mangrove strain MUSC115 T and the highest value with Cutibacterium acnes strain ATCC6919 T (Table 6). OrthoANI values of strain Marseille-P1302 with other closely related strains varied from 59.09% with Microbacterium mangrovi strain MUSC115 T to 75.05% with Brevilactibacter sinopodophylli strain KCTC 33808 T . Values observed among other compared species were in the same range (Figure 6).

Based on these phenotypic and genomic characteristics, we consider strain Marseille-P1302 as belonging to a new genus and a new species, for which we propose the names Nigeribacterium gen. nov. and Nigeribacterium massiliense gen. nov., sp. nov., respectively. Strain Marseille-P1302 T (= CSUR P1302) is the type strain of Nigeribacterium massiliense gen. nov., sp. nov., the type species of the genus Nigeribacterium gen. nov.

Description of Nigeribacterium gen. nov.

Nigeribacterium (ni.ge.ri.bac.te.ri'um. N.L. neutr. gen. n, combination of Niger, the country where the patient from whom the type strain was isolated, lived, and bacterium). Bacterial cells are aerobic, Gram-positive, non motile, non-spore-forming and cocci-shaped. Cells do not have any oxidase activity but are catalase-positive. Nigeribacterium massiliense gen. nov., sp. nov. is the type species of the genus Nigeribacterium gen. nov.

Description of Nigeribacterium massiliense gen. nov., sp. nov.

Nigeribacterium massiliense (mas.il.i.en'se. L. Gen. neutr. n. massiliense, of Massilia, the Latin name of Marseille, the place where the strain was isolated). In addition to the description of the genus, cells have a diameter of 0.8µm to 1µm. Colonies grown on 5% sheep blood-enriched Columbia agar are small, smooth and transparent, varying in size from 0.2mm to 0.4mm in diameter(Figure 2). Growth is observed at 28 and 37°C, but optimal growth occurs at 37°C in aerobic atmosphere. Cells show weak growth in anaerobic and microaerophilic conditions.

Using an API ZYM strip, leucine arylamidase, valine arylamidase, naphthol-AS-B1phoshohydrolase, β-galactosidase, α-glucosidase and β-glucosidase activities are observed, but negative results are obtained for alkaline phosphatase, esterase (C4), esterase lipase (C8), lipase (C14), cystine arylamidase, trypsin, α-chymotrypsin, acid phosphatase, α-galactosidase, βglucuronidase, N-acetyl-β-glucosaminidase, α-mannosidase and α-fucosidase activities. Using an API 50CH strip, cells are able to metabolize D-galactose, D-glucose, D-fructose, Nacetylglucosamine, esculine, D-maltose, D-lactose, D-saccharose, D-turanose and D-arabitol (Table 1). The most abundant fatty acids are 12-methyl-tetradecanoic acid (89%) while some traces of 12-methyl-Tridecanoic acid (4%) and Hexadecanoic acid (2%) are also detected (Table 2). The genome is 2,934,258-bp long, its G+C content is 70.38% with genes distributed in different COG categories shown in Figure 4 and in Table 3.

The type strain, Marseille-P1302 T , isolated from the fecal sample of a patient from Niger and suffering from Kwashiorkor, was deposited in the Collection de Souches de l'Unité des Rickettsies (CSUR, WDCM 875) and Deutsche Sammlung von Mikroorganismen und Zellkulturen (DSM) collections, under accession numbers CSUR P1302 and DSM 29084, respectively. The 16S rRNA and genome sequences are available in GenBank under accession numbers LK985392.1 and NZ_CCYM01000003., respectively.

  Using an API 50CH strip (bioMérieux), after[START_REF] Seemann | Prokka: Rapid prokaryotic genome annotation[END_REF] hours of incubation at 37°C, strain Marseille-P1302 was able to metabolize D-galactose, D-glucose, D-fructose, N-acetylglucosamine, esculine, D-maltose, D-lactose, D-saccharose, D-turanose and D-arabitol. In contrast, glycerol, erythritol, D-arabinose, L-arabinose, D-ribose, D-xylose, L-xylose, D-adonitol, Methyl-βDxylopyranoside, D-mannose, L-sorbose, L-rhamnose, Dulcitol, Inositol, D-mannitol, Dsorbitol, Methyl-αD-mannopyranoside, Methyl-αD-glucopyranoside, Amygdaline, Arbutine, Salicine, D-celiobiose, D-melibiose, D-trehalose, Inuline, D-melezitose, D-raffinose, Amidon, Glycogene, Xylitol, Gentiobiose, D-lyxose, D-tagatose, D-fucose, L-fucose, L-arabitol, Potassium gluconate, potassium 2-cetogluconate, potassium 5-cetogluconate were not metabolized (Table1).

Figure 1 .

 1 Figure 1. 16S rRNA-based phylogenetic tree highlighting the position of strain Marseille-

Figure 2 .

 2 Figure 2. Transmission electron microscopy of strain Marseille-P1302 T using a Tecnai G20

Figure 4 .

 4 Figure 4. Graphical circular map of the chromosome from strain Marseille-P1302 T . From the inside out the circle represents the COG categories, GC skew, G+C content and CDS.

Figure 5 .

 5 Figure 5. Distribution of genes from strain Marseille-P1302 T and closely related species into COGs functional categories. Category abbreviations are as follows: C, energy production and conversion; E, amino acid transport and metabolism; F, nucleotide transport and metabolism; G, carbohydrate transport and metabolism; H, coenzyme transport and metabolism; I, lipid transport and metabolism; P, inorganic ion transport and metabolism; Q, secondary metabolites biosynthesis, transport and catabolism; X, mobilome: prophages, transposons; A, RNA processing and modification; B, chromatin structure and dynamics; J, translation, ribosomal structure and biogenesis; K, transcription; L, replication, recombination and repair; D, cell cycle control, cell division, chromosomal partitioning; M, cell wall/membrane/envelope biogenesis; N, cell motility; O, posttranslational modification, protein turnover, chaperones; T, signal transduction mechanisms; U, intracellular trafficking, secretion, and vesicular transport; V, defense mechanisms; W, extracellular structures; Z, cytoskeleton; R, general function predicted only; S, function unknown.

Figure 6 .

 6 Figure 6. OrthoANI values obtained when comparing strain Marseille-P1302 T to phylogenetically closely related strains.

Table 1 .

 1 Differential characteristics of strain Marseille P1302 and related species

	Characteristics	1	2	3	4	5	6	7	8	9	10	11	12	13
	Gram stain	+	+	+	+	+	+	+	+	+	+	+	+	+
	Production of
	Catalase	+	+	+	+	+	+	na	+	-	+	+	-	+
	Oxidase	-	-	-	-	-	+	na	-	+	+	+	-	+
	Utilization of
	L-Arabinose	-	na	na	na	-	+	na	na	na	na	-	na	na
	D-Galactose	+	na	-	na	na	+	+	+	na	na	-	na	+
	D-Glucose	+	+	na	+	+	+	na	na	na	na	-	na	+
	D-Fructose	+	na	+	na	na	+	na	na	na	na	+	na	+
	N-acetylglucosamine	+	na	+	na	-	na	na	na	na	na	-	na	+
	Esculine	+	na	na	-	+	+	na	na	+	na	+	na	+
	D-Maltose	+	na	na	-	na	+	na	na	na	na	-	na	+
	D-Arabitol	+	na	+	na	na	na	na	na	-	na	-	na	+
	Enzymatic activities
	Alkaline Phosphatase	-	+	na	-	-	+	+	+	-	+	-	-	na
	Esterase (C4)	-	+	na	na	+	-	+	+	+(w)	+	+	na	na
	Esterase Lipase (C8)	-	+	na	na	+	+	+	+	+	+	+	na	na
	Lipase (C14)	-	-	na	+	-	-	-	-	-	-	-	na	na
	Leucine arylamidase	+	+	na	-	+	+	+	+	+	+	+	na	na
	Valine arylamidase	+	+	na	na	w	-	+	+	-	+	+/-	na	na

Table 2 .

 2 Cellular fatty acid composition of strain Marseille-P1302 T 491

	520	a Mean peak area percentage; TR, trace amount
	521
	522

Table 3 .

 3 Number of genes in strain Marseille-P1302 T associated with the 25 general COG functional categories.

	Code	Value	% Value a	Description
	J	149	5.51	Translation
	A	1	0.03	RNA processing and modification
	K	150	5.54	Transcription
	L	141	5.22	Replication, recombination and repair
	B	0	-	Chromatin structure and dynamics
	D	26	0.96	Cell cycle control, mitosis and meiosis
	Y	0	-	Nuclear structure
	V	34	1.25	Defence mechanism
	T	71	2.62	Signal transduction mechanism
	M	98	3.62	Cell wall/membrane biogenesis
	N	2	0.07	Cell motility
	Z	0	-	Cytoskeleton
	W	0	-	Extracellular structures
	U	21	0.77	Intracellular trafficking and secretion
	O	80	2.95	Posttranslational modification, protein turnover, chaperones
	X	0	0	Mobilome: prophages, transposons
	C	151	5.58	Energy production and conversion
	G	213	7.87	Carbohydrate transport and metabolism
	E	218	8.06	Amino acid transport and metabolism
	F	73	2.69	Nucleotide transport and metabolism
	H	85	3.14	Coenzyme transport and metabolism
	I	79	2.92	Lipid transport and metabolism
	P	95	3.51	Inorganic ion transport and metabolism
	Q	33	1.22	Secondary metabolites biosynthesis, transport and catabolism
	R	257	9.5	General function prediction only
	S	190	7.02	Function unknown
	others	537	19.85	Not in COGs

a: the value based on the total number of protein coding genes in the annotated genome

Table 4 .

 4 Details of resistance genes, plasmids and virulence factors detected in strain Marseille-P1302 T 538

	Database	Genes	Coverage % Identity %
	CARD
	(comprehensive
	antibiotic
	resistance
	database)
		Streptomyces_cinnamoneus_EF-Tu_mutants_conferring_resistance_to_elfamycin	99.58	82.09
		otr(A)	15.36	74.03
		novA	21.36	67.83
		Streptomyces_rishiriensis_parY_mutant_conferring_resistance_to _aminocoumarin	17.38	73.27
		Streptomyces_rishiriensis_parY_mutant_conferring_resistance_to _aminocoumarin	30.63	66.81
		mtrA	70.93	75.2
		Streptomyces_rishiriensis_parY_mutant_conferring_resistance_to _aminocoumarin	94.6	73.91
		novA	84.42	66.83
		novA	31.04	68.2
	Resfinder
		otr(A)_1	15.36	74.03
	ARG-ANNOT
		(Tet)otrA
			15.36	74.03
	Plasmidfinder	-	-	-

Table 5 .

 5 Genomic characteristics of strain Marseille-P1302 T and other compared species 541

	Species	Strain		Size (Mb)	G+C (%mol)	Gene content	tRNA	rRNA
	Nigeribacterium massiliense	Marseille-P1302	T	2.93	70.38	2704		9
	Amycolatopsis albispora	WP1 T		9.12	72.2	8434		12
	Microbacterium mangrovi	MUSC 115 T		4.41	70.9	4066		3
	Cutibacterium acnes	ATCC 6919 T		2.49	60	2308		9
	Aeromicrobium massiliense	JC14 T		3.31	77.32	3290		5
	Luteococcus japonicus	DSM 10546 T		3.45	67	3270		9
	Salinispora tropica	CNB 440 T		5.18	71.5	4662		9
	Tessaracoccus flavescens	SST 39 T		3.60	68.4	3515		6
	Nocardiodes gilvus	XZ17 T		3.33	68.9	3153		5
	Microlunatus phosphovorus	NM-1 T		5.68	68	5190		3
	Brevilactibacter sinopodophylii	KCTC 33808 T		3.07	70.79	2917		5
	Propionicicella superfundia	DSM 22317 T		3.61	69.99	3295		3
	Brevilactibacter flavus	VG341 T		3.00	67.69	2771		3
	542
	543

Table 6 .

 6 dDDH values obtained by comparing strain Marseille-P1302 T with other closely related species.

		Amycolatopsis	Microbacterium	Cutibacterium	Aeromicrobium	Luteococcus	Salinispor	Tessaracoccus	Nocardiodes	Microlunatus	Propionicicella	Brevilactibacter	Brevilactibacter
		albispora	mangrove strain	acnes strain	massiliense	japonicus	a tropica	flavescens	gilvus strain	phosphovoru	superfundia	sinopodophylii	flavus strain
		strain	MUSC	ATCC	strain	strain DSM	strain	strain SST	XZ17 T (NZ_	s strain NM-	DSM	KCTC	VG341 T (PPCV
		WP1 T (NZ_CP	115 T (NZ_JTDK	6919 T (NZ_CP	JC14 T (CAHG0	10546	CNB	39 T (NZ_CP01	QHKY0000	1 T (NC_0156	22317 T (AUIA0	33808 T (SDMQ01	01000001.1)
		15163)	00000000.1)	023676)	1000001.1)	T (NZ_RKH	440(NC_0	9607.1)	0000.1)	35.1)	1000001.1)	000001.1)
						G00000000.	09380.1)
						1)
	dDDH	18.7	17.9	22.8	18.4	20.3	20.2	19.8	18.6	19.8	19.1	19.8	19.4

Acknowledgments

The study was supported by the Méditerranée Infection foundation, the National Research Agency under the program "Investissements d'avenir", reference ANR-10-IAHU-03 and by Région Provence Alpes Côte d'Azur and European funding FEDER PRIMI.

Author contribution statement

MK performed the genomic analysis and phenotypic tests, DS participated in the genomic analysis, RZ participated in the phenotypic analysis, SIT isolated the bacterium, DR, JCL and PEF designed the study and wrote the article. All authors reviewed the manuscript.

Additional information

There are no competing interest.

Figure 3. Gram staining image of strain Marseille-P1302 T

Nocardiodes gilvus strain XZ17 T ; 10, Microlunatus phosphovorus strain NM-1 T , 11, Brevilactibacter sinopodophylli strain KCTC 33808 T , 12, Propionicicella superfundia 488 strain DSM 22317 T , 13, Brevilactibacter flavus strain VG341 T , Abbreviations: +, positive; -, negative; na, not available; w, weak.