The aim of this paper is to represent the implementation of service-learning methodology into Croatia's information sciences which contributed to meeting the needs of the rural population's on two Croatian islands. Those needs defined as a digital skills in rural tourism for adults and STEM education of elementary school children are usually not easily accessible in rural areas.

Thirteen students of information sciences from the University of Zagreb have spent a week with rural organization LAG 5 from Korčula implementing robotics workshops for children (Micro:bit and MBot) and teaching web and online advertising for rural entrepreneurs. This kind of collaboration helped develop the rural community's core skills and entrepreneurial capabilities but also motivated students for developing their expertise in teaching, problemsolving, teamwork, taking initiative and innovation. Service-learning proved to be an effective tool for strengthening the partnership and support between universities and rural organizations in which we boost social innovation and explore specific rural development opportunities.

Introduction

This paper aims to present the first example of rural service-learning in Croatia's information sciences. University students met the rural population's needs defined as digital skills in rural tourism -adult education and STEM education of elementary school children.

Our case unfolded as a part of the "Rural 3.0: Service-Learning for Rural Development project (RURASL)". One of the RURASL project's main goals is to help develop the rural community's core skills and entrepreneurial capabilities (for which such development is not easily accessible) and improve the quality of education for sustainable development through servicelearning university-community partnerships.

As part of the course Urban and Rural Service-Learning, within the project RURASL, graduate students of information sciences from the University of Zagreb (Croatia) had the opportunity to apply knowledge and skills gained at the university in solving the real needs of the rural community on the island of Korčula and Pelješac through the experience of servicelearning (SL). Thirteen students have spent a week (40 hours) implementing 4 robotics workshops for children (Micro:bit and MBot workshops) and 2 workshops for older adults (Web for Rural Entrepreneurs and Online Advertising for Rural Entrepreneurs). The number of school children who participated in 4 workshops on robotics was 36, while the number of older adults participating in the workshops was 10.

The rural community partner was a multistakeholder organisation devoted to sustainable rural development of the Peljesac peninsula and islands Mljet, Korcula and Lastovo, Local Action Group "LAG 5". It is a local NGO implementing the European Union LEADER approach, LEADER standing for "Links Between Actions for the Development of the Rural Economy."

The STEM field (science, technology, engineering & mathematics) has recently stood out as necessary for developing the economy both in the world and in Croatia. However, the interest of young people in the STEM field in Croatia is not sufficiently developed, as evidenced by the fact that the Ministry of Science and Education has awarded 3,400 direct state scholarships in recent years to encourage high school pupils to enrol in studies in the STEM field 1 .

Nevertheless, it needs to be emphasised that the STEM field's interest needs to be nurtured by providing access to technology and education from early school age.

As an innovative approach to teaching and learning that brings students, academics, and the community to develop solutions for challenging issues jointly, service-learning can promote STEM education for elementary school children and serve underrepresented populations (Mikelic Preradovic, 2015). By using service-learning, teachers increasingly work to encourage girls and minorities [START_REF] Naukkarinen | STEM: A help or a hinderance in attracting more girls to engineering[END_REF]

Literature review

Service-learning and STEM education

According to Mikelic Preradovic (2019), jobs in the natural sciences, technology, engineering and mathematics (STEM jobs) are paid 29% more than jobs in other fields. These statistics have been confirmed in other EU countries as well, where STEM professionals are on average better paid than other professionals [START_REF] Reynolds | Skills and Jobs in the Internet Economy[END_REF]. Also, in the last ten years, job growth in the STEM area has been three times higher than the growth of other jobs, and it is expected that in the next decade, the number of jobs in the STEM area will continue to 8.0 percent by 2029, compared with 3.7 percent for all occupations (Zilberman & Lindsey, 2021).

Despite the demand for STEM graduates, it is difficult for companies to find qualified candidates for many jobs. Two main factors contribute to this problem: (1) insufficient student enrollment in STEM colleges and (2) a lack of qualified teachers in the STEM field (Eurydice, 2021). In addition to the fact that many students never enter the STEM area due to inadequate school preparation and teachers' poor quality training [START_REF] Ejiwale | Barriers To Successful Implementation of STEM Education[END_REF], many employees in the STEM area are approaching retirement, suggesting additional demand for this profile.

Service-learning and adult education

According to statistics in the Curriculum for the Development of Basic Digital, Mathematical and Reading Skills of Adults (MZO, 2019), 20 to 25% of European adults between the ages of 16 and 65 have a low level of digital literacy (ability to solve problems in technology-rich environments). Furthermore, CEDEFOP (2019) provides data that 37.4% of adults (25 -64 years) in Croatia have a low digital literacy level. Regarding the adoption of digital skills, the motivation to use lifestyle technologies is emphasised (social networks, watching videos, informal communication, etc.). In contrast, a low level of competence in workplace-related skills was observed (problem-solving in a complex technological environment-related primarily to work tasks, knowledge of technologies such as Excel, PowerPoint, web design, etc.). Research shows us there is a substantial gap to be breached in the adoption of digital skills in problem-solving activities, as well as the computer is rarely used to run a personal website [START_REF] Pavić | Digitalne vještine polaznika obrazovanja odraslih. 7. Međunarodni andragoški simpozij Biograd na moru[END_REF].

Approach

Service-learning was used as "a form of experiential education in which students engage in activities that address human and community needs together with structured opportunities intentionally designed to promote student learning and development." [START_REF] Jacoby | Service-learning in today's higher education[END_REF] to meet the rural community's needs of the island of Korcula and Peljesac.

The proof of student "learning" in "service-learning" was specified as the ability to synthesise material adopted from academic literature with real-life experiences to create an effective curriculum for pupils and adult users. The specific course conditions and skills that students should have acquired after the Urban and Rural Service-Learning course are shown in Table 1.

Table 1. The specific course conditions and skills of students after the Urban and Rural

Service-Learning course

Results

Students of information sciences have spent a week (40 hours) implementing their servicelearning solutions on-site in a summer school format. The number of school children who participated in 4 workshops on robotics was 36. The number of older adults who participated in workshops on web advertising and web development (acquiring more advanced digital skills that involve solving problems in a complex technological environment related primarily to work tasks) was 10.

The workshops aimed to develop digital skills among older adults who were members of various associations in LAG 5. People who still do not have a website of their association and who (as small entrepreneurs) are engaged in tourism and apartment rental and need to acquire knowledge and advertising skills on various online platforms.

Upon the completion of the workshops, students have supplemented their workshop materials with explanations and comments from rural partner LAG 5 and, as a result, compiled a manual with examples that are applicable to meet similar rural needs in other EU countries and can be used by university students to conduct similar rural service-learning workshops or by rural residents for individual self-paced learning (Figure 1.) Upon completing the project, each of them completed an open-ended questionnaire that provided them with self-evaluation by analysing the most effective parts of the project and the skills and knowledge acquired by their SL project.

The analysis of the answers identified the components that the students singled out as the most efficient part of the project (Table 2.). Furthermore, what students single out as the most important things they have acquired and learned with this SL project, and what will serve them in the future as well are shown in Table 2. 2. Main project results and benefits for the students which arose after SL implementation.

• Awareness of the role of the public in the dissemination of SL projects.

• Attitudes towards children and the elderly population,

• Awareness of the importance of SL for community development, but also personal development • Students did most of the work when divided the tasks and performed them individually. The principle in which everyone is responsible for their task prevents the diffusion of responsibility that occurs when it is not defined whose task it is • Workshops with children in schools that brought great joy to those children who discovered their passion for STEM

• Having fun figuring out how to present robotics to children

Benefits for the students

• Respect for others, but also yourself

Most efficient part of the project

• Increasing children's interest in the STEM area • Applying the acquired knowledge in real situations and testing that knowledge in teaching adult users

Discussion

Rural service-learning enabled university students to understand how STEM and digital competencies can benefit the community they live in and themselves.

In his evaluation, after the course ended, the teacher singled out the following set of crucial skills that students need when engaging in rural service-learning:

• skills for responsibility towards the rural community,

• working with children and/or adult users,

• connecting with professors and superiors in non-teaching situations,

• adapting communication techniques to children,

• communication and presentation skills,

• oral expression and public speaking,

• presentation of STEM topics in a popular science way,

• liberation from fear of public speaking,

• gaining experience working in a team,

• trying and learning something new,

• applying the knowledge acquired at the college in practice more innovatively and excitingly,

• acquiring knowledge about project implementation,

• understanding what it is like to work on a project, what is expected from a student, what his/her tasks are, how to function in group work and apply everything in real life while transferring their knowledge to someone else.

Finally, the following challenges faced by students were identified by the teacher during the implementation of rural service-learning:

• adaptation of materials to different age groups,

• a lot of revision and redesign at each stage,

• reconciling with the fact that good ideas are not always feasible,

• adapting oneself to the new situation,

• accepting that it is not necessary that everyone has to do everything, but that a good division of labour and communication is essential,

• changing plans in collaboration with teammates,

• harmonising the ideas of team members who have different perspectives,

• flexibility and understanding for other people's obligations,

• agreement and adjustment of deadlines to implement activities that suit all team members.

Implications and limitations

Educational robotics workshops for rural school children served not only to acquire knowledge in STEM areas but also to develop competencies such as cooperation in group work, communication, learning skills, creativity, curiosity, etc.

Nevertheless, university teacher and LAG 5 community partner evaluations of the rural servicelearning described in this paper revealed several challenges, which are in line with the main findings of Visvizi, Lytras and Mudri (2019).

One of the main challenges was students' transportation from the university to the remote rural community (island), which consequently influenced students' time to engage in activities. The other issues were poor Internet access and lack of digital infrastructure in rural area. Finally, the community partner's support for students who were engaged in the summer school also presented a challenge: community partner had to deal not only with issues of student meals, housing and communication with rural residents but also with the requirement to implement service-learning in a much shorter time frame than a traditional semester.

On the positive side, elementary school pupils in this project, using robots, entered the world of programming and robotics by learning how to program a robot to automatically perform actions such as driving in different directions, turning lights on and off, moving along the line, detecting obstacles and more. They have shown an increase in STEM competencies and an affinity for STEM education and careers in the STEM field.

The evaluation results showed that university students felt that SL encouraged them to learn and improve their understanding of the teaching material.

Referring university students to all robots' pedagogical applications helped them become aware that robots are not toys. Solving problems using theoretical concepts enabled them to think about the practical application of theory to other problems. Students also acquired several soft skills such as teaching, problem-solving, teamwork, initiative, innovation.

Students' central challenge was the doubt in their ability to teach robotic and engineering concepts and maintain children's attention.

The workshops' results (and their evaluation) described in this paper show the importance of initiating such trends in rural areas, where access to equipment and extracurricular education is much more limited than in urban areas. However, Croatia represents a country where 42.9%

of the total number of inhabitants lives in a rural area, similar to Estonia, Austria, Slovenia and Romania (Eurostat, 2018) and digital literacy in the rural area is crucial to their social and economic development (ENRD, 2014).

Service-learning proved to be an effective tool for building partnerships between rural community and university that helped to solve developmental problems through the acquisition of skills. It enabled rural inhabitants to develop their skills while producing graduates who are adaptive to change, equipped for their future challenges, who possess a depth of knowledge in their field of study, but also a breadth of knowledge to address social issues in their communities.

  traditionally neglected in STEM, to consider STEM as a future career. However, there are three critical factors for creating a STEM generation: engagement, motivation, and exposure to STEM subjects (De Loof et al, 2019). Each of these factors can be addressed by implementing service-learning in STEM. Furthermore, recent research on adults' digital skills in adult training and development programs in Croatia (Pavić, 2019) from open public universities in seven Croatian cities shows a relatively low digital skills level which can be increased through service-learning projects, as demonstrated by Hanks & Icenogle (2001) in their USA experiences. They revealed that university students could help adults over 50 develop work-related skills such as self-esteem, resume development, and computer skills. Therefore, students in the course Urban and Rural Service-Learning have devised solutions that meet the rural needs and have implemented them through service-learning in the rural community on the island of Korcula and Peljesac.

  According to Lemons, Carberry, Swan & Jarvin (2011), expanding the core competencies of engineering students and incorporating skills essential to SL (collaboration, customer awareness, and ethics understanding) could result in a population of gender and ethnically diverse engineering professionals. According to them (ibid.), "the positive effects of SL activities are attributed to the processing of experiences through critical reflection, discussion, and interaction with other students and professors". Intel Corporation's survey (2011) showed that teenagers' interest in enrolling in one of the STEM colleges increases when they hear how their choice can benefit the world. Examples of the implementation of service-learning in technology include mobile robots (Bhounsule, Chaney, Claeys & Manteufel, 2017), the Internet of Things (Watson & Ogle, 2013) and intergenerational computing (Wexler, Drury, Coppola, Tschinkel & Thomas, 2011).

  As part of the Urban and rural service-learning course, university students planned and conducted service-learning workshops for elementary school pupils and adults. The workshops "STEM education of school children" enabled university students' participation in the digital-literacy education of pupils, teaching them how to use the principles of robotics and automation as tools for transferring STEM competencies to children.University students (familiar with the basics of robotics and programming) were tasked with introducing elementary school pupils to the micro: bit and MBot educational kits and teaching them to program robots to solve assigned tasks. These activities provided students with an appropriate context for practical learning and teaching and opportunities to synthesise technical and transversal skills. Students were in charge of material development, preparation and implementation of workshops.At the workshop for adults, "Web for Rural Entrepreneurs", students had the task to teach participants what a website is, how it is maintained, what it consists of and finally, how to create it and publish it. The platform they chose was Wix because it is straightforward to use, requires no prior knowledge, and is free of charge.The other workshop for adults, "Online Advertising for Rural Entrepreneurs", aimed to teach online advertising in tourism. Participants were taught to work with tools and platforms such as Facebook, Instagram, Google My Business, Airbnb and Booking.

•

  Develop and implement websites and solutions for advertising on the web and social networks • Develop an extracurricular plan of robotics for elementary school students • Elaborate curriculum plan for the development of digital skills in adult education • Formulate and prove a hypothesis about how the approach and ability to work with technology affects education • Effectively teach pupils and adult users of different cultural and educational backgrounds Student skills • Develop and implement software solutions for micro: bit and mBot robots • Students tried to satisfy the need to introduce the extracurricular program to enrich natural science and technical subjects, develop adults' digital skills, and analyse their contribution to educational and community goals • Each student participated in at least 40 hours of service learning through workshops for the rural community • The experience of service-learning was connected with the course's academic goals • The course involved critical thinking about the relationship between academic content and community work Specific condition

Figure 1 .

 1 Figure 1. RURASL case-based learning materials: https://rural.ffzg.unizg.hr/wpcontent/uploads/2020/12/Case-base-learning-materials_Croatia_English-version.pdf

•

  Knowledge of writing texts for the education of other colleagues • Critical thinking • Organisational work and clear goal setting • Transfer knowledge to children as the most challenging but most important part of the population to educate, • Awareness of the time complexity of the project,

https://mzo.gov.hr/vijesti/tar-ministrica-divjak-sa-studentima-stem-games/1349


Josefa Bleu

Sandrine Meylan

Jean Clobert

Manuel Massot

Grandmaternal age at reproduction affects grandoffspring body condition, reproduction and survival in a wild population of lizards

Keywords: ageing, grandmaternal effect, lagged effect, inter-generational effects, senescence, Zootoca vivipara

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

Introduction

Age at reproduction is known to affect individual reproductive performance, with effects on both litter size and litter success [START_REF] Clutton-Brock | Reproductive success: studies of individual variation in contrasting breeding systems[END_REF]. First, these results can be explained in terms of the constraints imposed by age. If age is associated with improvements in physiology (e.g., developmental maturation), morphology (e.g., increase in size) or behavior (e.g., knowledge about breeding sites), we expect a positive effect of age in early adulthood (maturation) [START_REF] Forslund | Age and reproduction in birds -hypotheses and tests[END_REF]. In contrast, senescence is characterised by physiological deterioration in aged individuals. This process produces higher mortality rates and/or a decrease in reproductive success [START_REF] Nussey | Senescence in natural populations of animals: widespread evidence and its implications for bio-gerontology[END_REF]. Second, the age-related performance of an organism can reflect adaptive strategies [START_REF] Charlesworth | The relation of reproductive effort to age[END_REF].

Indeed, according to life-history theory, reproductive effort should increase as residual reproductive value declines; this life-history strategy serves to maximise lifetime reproductive success, and we should thus expect a positive effect of age in late adulthood (terminal investment) [START_REF] Pugesek | Increased reproductive effort with age in the California gull (Larus californicus)[END_REF][START_REF] Clutton-Brock | Reproductive effort and terminal investment in iteroparous animals[END_REF][START_REF] Roff | Life history evolution[END_REF].

Interestingly, these constraints and strategies can act at the intra-generational level (e.g., age affects reproductive success of the current generation) or at the inter-generational level with maternal age effects on offspring characteristics [START_REF] Roff | Life history evolution[END_REF]. Age-related selective pressures can even differ between intra-generational and inter-generational level [START_REF] Moorad | Evolution of maternal effect senescence[END_REF]. Even if they are not universal [START_REF] Moore | Is a decline in offspring quality a necessary consequence of maternal age[END_REF], maternal age effects can be observed on offspring traits at birth [START_REF] Massot | An integrative study of ageing in a wild population of common lizards[END_REF], on offspring survival [START_REF] Ducatez | Complex interactions between paternal and maternal effects: parental experience and age at reproduction affect fecundity and offspring performance in a butterfly[END_REF][START_REF] Ivimey-Cook | The diversity of maternal-age effects upon pre-adult survival across animal species[END_REF] or even on offspring reproductive success [START_REF] Bouwhuis | Trans-generational effects on ageing in a wild bird population[END_REF][START_REF] Bouwhuis | Sex-specific pathways of parental age effects on offspring lifetime reproductive success in a long-lived seabird[END_REF][START_REF] Schroeder | Reduced fitness in progeny from old parents in a natural population[END_REF]. If differences between offspring of the F1 generation (maternal age effects) persist and affect their own offspring in the F2 generation, we may observe grandmaternal (GM) age effects. Inter-generational effects can create lagged effects in population dynamics and these effects are now recognised as a source of individual heterogeneity that is important to consider to increase our understanding of population dynamics and persistence [START_REF] Benton | Complex population dynamics and complex causation: devils, details and demography[END_REF]. Inter-generational effects can also have evolutionary consequences. They induce time lags in the response of populations to selection, and this may facilitate or hinder adaptive evolution [START_REF] Kirkpatrick | The evolution of maternal characters[END_REF][START_REF] Mcglothlin | The contribution of maternal effects to selection response: an empirical test of competing models[END_REF]. Importantly, it is crucial to determine whether the effects of age are constant over multiple generations, or cumulative as found in a pioneering study in rotifers [START_REF] Lansing | A transmissible, cumulative, and reversible factor in aging[END_REF], to correctly assess their impact on the population and also to understand whether these effects are indeed adaptive. For instance, adaptive strategies could be selected if age is correlated with environmental variables and thus represents a source of information in itself that could enable to pre-adapt the offspring to the environmental conditions they will encounter. For example, a differential impact of environmental variables (density, resource availability, etc.) on survival rates of young and old individuals could correlate age and environmental conditions. For all these possible consequences, it may be important to measure and quantify GM age effects in natural population.

However, few studies have addressed the persistence of age effects over several generations. We are only aware of a few examples in human and some animals studied in laboratory conditions. In human medicine, it has been shown that GM age is a risk factor for trisomy 21 [START_REF] Papp | Grandmaternal age at birth of parents of children with trisomy 21[END_REF][START_REF] Aagesen | Advanced grandmaternal age on the mother's side -a risk of giving rise to trisomy 21[END_REF][START_REF] Malini | Influence of advanced age of maternal grandmothers on Down syndrome[END_REF], but see also, [START_REF] Allen | Maternal age and risk for trisomy 21 assessed by the origin of chromosome nondisjunction: a report from the Atlanta and national Down syndrome projects[END_REF][START_REF] Kovaleva | Grandmaternal ages at birth of parents of children with Down syndrome in St[END_REF] but not for breast cancer [START_REF] De Haan | Age of mother and grandmother in relation to a subject's breast cancer risk[END_REF] and that grandpaternal age is positively correlated with telomere size in grandoffspring [START_REF] Eisenberg | Delayed paternal age of reproduction in humans is associated with longer telomeres across two generations of descendants[END_REF][START_REF] Eisenberg | Older paternal ages and grandpaternal ages at conception predict longer telomeres in human descendants[END_REF]. In non-human animals, the pioneering study in rotifers [START_REF] Lansing | A transmissible, cumulative, and reversible factor in aging[END_REF] showed that the repeated effects of the same maternal age over multiple generations increasingly influenced grandoffspring lifespan, i.e., with evidence of a cumulative effect over multiple generations. A study on aphids [START_REF] Mackay | Maternal age as a source of variation in the ability of an aphid to produce dispersing forms[END_REF] showed that GM age influenced grandoffspring reproductive phenotype. More recently, studies on fruit flies have shown that GM age influences grandoffspring viability [START_REF] Hercus | Maternal and grandmaternal age influence offspring fitness in Drosophila[END_REF][START_REF] Bloch Qazi | Transgenerational effects of maternal and grandmaternal age on offspring viability and performance in Drosophila melanogaster[END_REF], developmental instability [START_REF] Faurby | The effect of maternal and grandmaternal age in benign and high temperature environments[END_REF][START_REF] Røgilds | Maternal and grandmaternal age effects on developmental instability and wing size in parthenogenetic Drosophila mercatorum[END_REF][START_REF] Kjarsgaard | Effects of temperature and maternal and grandmaternal age on wing shape in parthenogenetic Drosophila mercatorum[END_REF] and short-term memorisation capacities [START_REF] Burns | Transgenerational memory effect of ageing in Drosophila[END_REF]. GM age effects were also found affecting grandoffspring weight in the rice weevil [START_REF] Opit | Influence of maternal age on the fitness of progeny in the rice weevil, Sitophilus oryzae (Coleoptera: Curculionidae)[END_REF] and grandoffspring growth and size at maturation in a Daphnia [START_REF] Goos | Maternal diet and age alter direct and indirect relationships between life-history traits across multiple generations[END_REF]. At the population level, it has also been shown in soil mites that maternal age affects population dynamics for several generations [START_REF] Benton | Maternal effects mediated by maternal age: from life histories to population dynamics[END_REF][START_REF] Plaistow | The influence of context-dependent maternal effects on population dynamics: an experimental test[END_REF]. None of these studies have, however, examined the possibility that such GM age effects occur in natural populations.

In the wild, GM age effects are unexpected for several reasons. First, it has often been assumed that too few individuals survive to advanced ages in the wild to be able to conduct robust statistical analyses on ageing [START_REF] Kirkwood | Why do we age?[END_REF]. Second, the high level of environmental variance in nature increases the amount of inter-individual variation (e.g., the variance in resource acquisition is increased), reducing the chance to detect GM age effects.

Third, disappearance/appearance of different phenotypes at the population level (e.g., selection) [START_REF] Forslund | Age and reproduction in birds -hypotheses and tests[END_REF][START_REF] Pol | Age-dependent traits: a new statistical model to separate within-and between-individual effects[END_REF] may obscure age effects. Thus, even if age effects are observed under laboratory conditions that enhance the overall survival of individuals, they may not be observed in natural conditions. Although it has been proven possible to measure maternal age effects in the wild [START_REF] Clutton-Brock | Reproductive success: studies of individual variation in contrasting breeding systems[END_REF][START_REF] Nussey | Senescence in natural populations of animals: widespread evidence and its implications for bio-gerontology[END_REF], sample sizes to investigate age effects on one additional generation (GM effects) are even more constrained. However, three studies that documented parental effects on the number of grandoffspring recruits showed the feasibility to test GM age effects in wild populations [START_REF] Bouwhuis | Trans-generational effects on ageing in a wild bird population[END_REF][START_REF] Bouwhuis | Sex-specific pathways of parental age effects on offspring lifetime reproductive success in a long-lived seabird[END_REF][START_REF] Schroeder | Reduced fitness in progeny from old parents in a natural population[END_REF]. The constraint on sample size can be circumvented by the long-term monitoring of large populations. Recently, this was performed by a study on semi-captive Asian elephants in which it was shown a negative GM age effect on grandcalf survival [START_REF] Reichert | Maternal age at birth shapes offspring life-history trajectory across generations in long-lived Asian elephants[END_REF]. The ultimate challenge for studies that will detect GM age effects in wild populations will be to have enough repeated measurements of the same grandmothers at different ages to discriminate between individual and population level mechanisms (e.g., age-specific selection).

In our study, we tested the effects of grandmaternal age (F0 generation) on the characteristics of grandoffspring at birth (F2 generation), on grandoffspring survival and on the reproduction of granddaughters in a wild population of the common lizard, Zootoca vivipara (Figure 1). In this species, maternal age affects reproductive success and offspring characteristics at birth [START_REF] Massot | An integrative study of ageing in a wild population of common lizards[END_REF]. In particular, there is a positive effect of female age on female offspring body condition and body size at birth, and this was related both to maturation in young females and terminal investment in old females [START_REF] Massot | An integrative study of ageing in a wild population of common lizards[END_REF]. This variation in female offspring quality due to the age of their mother could affect the reproductive outputs of female offspring and lead to GM age effects.

Materials and Methods

Model species and long-term survey

The common lizard (Zootoca vivipara, formerly Lacerta vivipara) is a small grounddwelling lizard that is widely distributed across Eurasia. It is a live-bearing species, except in the extreme south of Europe where oviparous populations have been recorded (Surget-Groba et al., 2001). In the study population, the average litter size is five shell-less eggs, and females give birth to fully formed juveniles that emerge from the foetal membrane within one or two hours after laying. There is no parental care, and juveniles are independent from birth. Lizards reproduce once per year, during the summer, and hibernate from late September to April-May.

Age-specific survival rates increase until maturity (usually at two years in our population), and then decrease with age in males and also in females that showed a high reproductive effort during their first reproduction [START_REF] Massot | An integrative study of ageing in a wild population of common lizards[END_REF]. The data of our study were obtained from a long-term mark-recapture survey conducted each year from 1989 through 2004 in a natural population at Mont-Lozère (1420 m a.s.l., 44°23'03''N, 3°52'40''E, Cévennes National Park, southeastern France). This population and the long-term survey protocol have been described elsewhere (e.g., [START_REF] Massot | Processes at the origin of similarities in dispersal behaviour among siblings[END_REF][START_REF] Galliard | Cohort variation, climate effects and population dynamics in a short-lived lizard[END_REF][START_REF] Massot | An integrative study of ageing in a wild population of common lizards[END_REF][START_REF] Bleu | Reproductive allocation strategies: a long-term study on proximate factors and temporal adjustments in a viviparous lizard[END_REF]. Briefly, pregnant females are captured each year and kept in the laboratory for threefour weeks until parturition. No sample bias is expected because capture probabilities of adult females do no vary with age [START_REF] Massot | An integrative study of ageing in a wild population of common lizards[END_REF]. At birth, offspring are measured (snout-vent length, to the nearest mm), weighed (to the nearest mg), sexed and marked by toe-clipping with no effect on subsequent recapture and survival probabilities [START_REF] Massot | Density dependence in the common lizard: demographic consequences of a density manipulation[END_REF]. This protocol allows us to determine the maternal lineages, the age at reproduction, litter size, litter success, offspring body size and body mass at birth, and offspring sex. Lizards that are not born in the laboratory can be aged if they are captured in the field as juveniles or subadults, and this allows us to age many grandmothers. Offspring are recaptured at the end of the reproductive season (in September, i.e. before hibernation) and the summer of the next years (in June-July) to estimate their survival. Litters do not always result in live offspring, they can contain stillborns, unfertilised eggs or abortive embryos. We defined litter size as the total litter size, and litter success as the proportion of live offspring in a litter.

Dataset

The full dataset contains 579 grandoffspring from 135 litters, with sampled grandmothers and mothers from two to eight years old (Tables S1 andS2 in Supporting Information). However, in this dataset, some litters were not independent because they were produced by the same grandmother or mother. More precisely, 25 grandmothers and 26 mothers appeared two times in the dataset, five grandmothers and three mothers appeared three times, four grandmothers and one mother appeared four times and one grandmother appeared five times. The identity of a grandmother or a mother can be repeated for several reasons. Firstly, some litters were produced by different mothers but shared the same grandmother: in eight cases the grandmother had the same age (mothers were from the same litter) and in seven cases the grandmother had different ages (mothers from two different years). The last case is of particular interest because it could allow longitudinal analyses, however it was not very frequent. Secondly, some litters were produced by the same mother at different maternal ages (and thus the same grandmother at the same GM age). This was the most frequent case, with 26 cases where the mother was captured two different years, three cases three different years and one case four different years.

Statistical analyses

We tested the effects of grandmaternal (GM) age on grandoffspring characteristics at birth and on the reproduction of granddaughters with mixed and linear models using R 4.1.1 statistical software (R Core Team, 2021). The significance of the fixed effects was tested using F statistics for linear mixed models and a chi-square test for logistic regression [START_REF] Bolker | Generalized linear mixed models: a practical guide for ecology and evolution[END_REF]. We checked the homoscedasticity and the normality of the residuals of the linear mixed models. The collinearity of the explanatory variables was also checked using variance inflation factors.

Firstly, we analysed the effects of GM age on grandoffspring body size at birth (snout-vent length) and on grandoffspring body condition at birth. We defined body condition as the residuals of a linear regression of body mass against body size. The regression was performed for each sex separately because of sexual size dimorphism in juvenile common lizards (Le [START_REF] Galliard | Ontogenic sources of variation in sexual size dimorphism in a viviparous lizard[END_REF]. We used the 'lmer' function from the packages 'lme4' (version 1.1-27) [START_REF] Bates | Fitting linear mixed-effects models using lme4[END_REF] and 'lmerTest' (version 3.1-3) [START_REF] Kuznetsova | lmerTest Package: Tests in Linear Mixed Effects Models[END_REF] to compute linear mixed models with the Satterthwaite's method to compute degrees of freedom and F-statistics.

The explanatory variables of the full models were: GM age, maternal age (M age), offspring sex and the first-order interactions. Because age effects are often curvilinear (e.g., [START_REF] Massot | An integrative study of ageing in a wild population of common lizards[END_REF], the quadratic terms of the GM age and M age were also included in the model. The random effects were the year of maternal reproduction and maternal identity (offspring from the same litter are not independent and some mothers may appear different years in the dataset). The identity of the grandmother was not included as a random effect because the variance explained by this effect was close to 0. We believe that it might not be relevant to include this effect as we did not have enough repetitions of grandoffspring from the same grandmother but different mothers to estimate this properly. We also did not have individual estimates of longevity of mothers and grandmothers to include as covariates to control for selective disappearance related to maternal and grandmaternal ages (e.g., [START_REF] Hayward | Reproductive senescence in female Soay sheep: variation across traits and contributions of individual ageing and selective disappearance[END_REF].

Secondly, we analysed reproduction of granddaughters at the age of two years old, i.e.

at their first reproduction [START_REF] Massot | An integrative study of ageing in a wild population of common lizards[END_REF]. We analysed granddaughter litter size and granddaughter litter success with logistic regression (generalized linear models). For litter size, we used a Conway-Maxwell-Poisson regression to take into account underdispersion of these count data (dispersion parameter = 0.26) (Sellers & Premeaux, 2020). More precisely, we used the 'glm.cmp' function from the package 'mpcmp' (version 0.3.6). For litter success, we used a logistic regression with quasi-binomial error to take into account overdispersion of these proportion data (dispersion parameter = 1.66). We used the 'glm' function (base package) and "Anova" from the car package (version 3.0-11) [START_REF] Fox | An R Companion to Applied Regression[END_REF] to compute type 3 anova. Because of the small sample sizes, we tested only the interaction between GM age and M age and we did not add random effects (the identity of only one mother and two grandmothers were repeated in the dataset). The explanatory variables of the full model were: GM age (linear and quadratic terms), M age (linear and quadratic terms), and granddaughter body size. We included granddaughter body size because reproduction depends on body size in female common lizards [START_REF] Massot | Density dependence in the common lizard: demographic consequences of a density manipulation[END_REF][START_REF] Massot | An integrative study of ageing in a wild population of common lizards[END_REF].

Survival analyses

Grandoffspring were recaptured at the end of the summer before their first hibernation and each summer of the next years. The resulting capture-mark-recapture data allowed us to estimate the survival probabilities. Survival estimates were obtained while taking into account the capture probabilities using a capture-mark-recapture method based on the open population model of Cormack-Jolly-Seber [START_REF] Clobert | A general approach to survival rate estimation by recaptures or resightings of marked birds[END_REF]. This model produces apparent survival estimates resulting from mortality and emigration. Because we have shown that emigration is rare in our study population, our estimates may be assimilated to true survival rates [START_REF] Massot | Density dependence in the common lizard: demographic consequences of a density manipulation[END_REF]. We used the program E-surge version 2.2.3 to fit models (Choquet et al., 2009b).

We tested whether survival and capture probabilities were dependent on GM age and M age (linear and quadratic terms), and the interaction between the two. The full model of survival probabilities also included as factors the season (autumn/summer), the year and the sex in interaction with age classes (variation in survival between age classes are reported in [START_REF] Massot | An integrative study of ageing in a wild population of common lizards[END_REF]. Concerning capture probabilities, the full model included the season, year and sex factors in addition to the variables of GM age and M age (linear and quadratic terms, and their interaction). We first selected a model for capture probabilities (while keeping the full model for survival probabilities) and then a model for survival probabilities. The model selection was based on the quasi-likelihood Akaike information criterion (QAICc), this criterion accounts for the overdispersion of data that can be induced by missing factors [START_REF] Burnham | Model Selection and Multimodel Inference: a practical informationtheoretic approach[END_REF]. The goodness of fit of the Cormack-Jolly-Seber model to the data was checked with the program U-CARE version 2.3.4 (Choquet et al., 2009a).

Age-structured matrix model

In order to approximate the GM age effect on granddaughter fitness, we implemented an age-structured matrix model with the ULM software version 6.0 [START_REF] Legendre | ULM, a software for conservation and evolutionary biologists[END_REF]. We modelled the life cycle of the common lizard from a 9-age class Leslie matrix, and we calculated the asymptotic multiplication rate as the dominant eigenvalue of the matrix.

The asymptotic multiplication rate was used as a proxy of fitness. As usual for this kind of modelling, our model was based solely on females. This approach assumes that male abundance is always high enough to ensure reproduction of most females [START_REF] Bessa-Gomes | Discrete two-sex models of population dynamics: on modelling the mating function[END_REF]. This assumption is relevant to our studied population where all sexually mature females (females of three years of age and older, and two years old females with a body size larger than a threshold) reproduce [START_REF] Massot | An integrative study of ageing in a wild population of common lizards[END_REF]. We fixed the female age at maturity at two years as usually observed [START_REF] Massot | An integrative study of ageing in a wild population of common lizards[END_REF]. The proportion of females at birth was fixed at 50% (Le [START_REF] Galliard | Female common lizards (Lacerta vivipara) do not adjust their sex-biased investment in relation to the adult sex ratio[END_REF] to adjust fecundities to the numbers of females produced. The values of age-specific fecundity of granddaughters and survival for the first winter were the estimates from this study (Figure 1). The ULM software allows to compute the age-specific multiplication rates and to measure the multiplication rates sensitivity to change in the different vital parameters tested [START_REF] Legendre | ULM, a software for conservation and evolutionary biologists[END_REF][START_REF] Caswell | Matrix population models: construction, analysis, and interpretation[END_REF].

The ULM software can be downloaded from http://www.biologie.ens.fr/~legendre/ulm/ulm.html.

Results

Body size and body condition at birth of grandoffspring

Table 1 shows the results of the analysis on body size. Body size at birth of F2 individuals was dependent on maternal age (F1 generation) and grandoffspring sex. F2 females were bigger than F2 males and the effect of maternal age was quadratic with a maximum for mothers of five years old (Figure S1). However, there was no significant effect of GM age (F0 generation).

Body condition at birth of F2 individuals was dependent on maternal age (F1 generation) and GM age (F0 generation) (Table 2). Body condition at birth was dependent on maternal age in interaction with sex. F2 males had lower body condition than F2 females from older mothers (Figure S1). Body condition of F2 individuals was also significantly related to GM age in interaction with sex (Table 2). Body condition decreased with GM age for F2 males (estimate ± s.e. = -1.27 ± 1.07) but not for F2 females (estimate = 0.60 ± 1.10). As a consequence, F2 males had lower body condition than F2 females from older grandmothers (Figure 2).

Survival of grandoffspring

We studied M age (F1) and GM age (F0) effects on survival probabilities of grandoffspring (F2 individuals). Figure 3 shows survival probabilities of grandoffspring during their first hibernation, and the range of 95% confidence interval of survival probabilities for all the combinations of grandmaternal and maternal ages. Survival probabilities of grandoffspring had large confidence intervals mainly in the combinations of the oldest grandmothers and mothers (rigth panels of Figure 3). Grandoffspring survival was not confidently estimated in these combinations of the oldest ages as we could expect from their small sample sizes (Table S2). Figure 3 (left panels) reports only survival probabilities of grandoffspring the most confidently estimated. Overall, we found an effect of both M age (linear effect) and GM age (quadratic effect) (log-likelihood ratio tests: X 2 1 = 6.1 P = 0.013 for M age effect, X 2 1 = 8.1 P = 0.005 for GM age quadratic effect; detailed analysis in Table S3). The maternal and grandmaternal effects were additive. Male and female offspring of the oldest mothers showed the highest survival rates (left panels of Figure 3). For the quadratic GM age effect, the survival rates of male and female grandoffspring were the highest for the three years old grandmothers (left panels of Figure 3).

Reproduction of granddaughters

Twenty-two pregnant granddaughters were recaptured at the age of two years old, the age at which females may first reproduce in this population [START_REF] Massot | An integrative study of ageing in a wild population of common lizards[END_REF]. Litter size and litter success of granddaughters (F2 females) were dependent on GM age (F0 females) (Tables 3 and4). The reproductive performances of granddaughters increased with GM age (Figure 4). Granddaughter litter size was also positively correlated with granddaughter body size (Table 3). Granddaughter litter size and success were not significantly related to maternal age (Tables 3 and4).

Age-structured population model

We calculated the net effect of GM age effects by estimating the multiplication rate (λ) of granddaughters produced by GM of different ages with an aged-structured matrix model. This measure is an approximation of the fitness of granddaughters since it takes into account both the GM age effects on survival and fecundity. GM age has a non linear effect on λ, with maximum values for three and four years old GM (Figure 5). λ is clearly smaller for the granddaughters produced by grandmothers older than four years old (Figure 5). The sensitivity was 0.71 for juvenile survival, 0.05 for fecundity at age 2, 0.03 for fecundity at age 3, and less than 0.02 for fecundity at older ages. This indicates that most of this pattern is driven by the GM effects on grandoffspring survival.

Discussion

A recent review of animal populations has illustrated that offspring quality often depends on maternal traits [START_REF] Moore | A mother's legacy: the strength of maternal effects in animal populations[END_REF]. The literature on maternal effects expanded greatly over the last three decades and showed that these effects are common and affect a diversity of traits. Specifically, a series of previous studies in the common lizard found that this live-bearing species exhibits multiple maternal effects. For example, offspring body size and body condition were dependent on maternal age [START_REF] Massot | An integrative study of ageing in a wild population of common lizards[END_REF], offspring growth and survival were influenced by temperature during gestation [START_REF] Lorenzon | The contribution of phenotypic plasticity to adaptation in Lacerta vivipara[END_REF], offspring dispersal responded to food availability during gestation [START_REF] Massot | Climate warming, dispersal inhibition and extinction risk[END_REF], and litter size was related to rainfall during vitellogenesis [START_REF] Bleu | Reproductive allocation strategies: a long-term study on proximate factors and temporal adjustments in a viviparous lizard[END_REF]. The commonness of maternal effects in the common lizard might explain why we found grandmaternal age effects on four out of the five parameters tested, with GM age effects on body condition at birth, survival and two reproductive parameters. Interestingly, these grandmaternal age effects were independent of maternal age effects. GM age and M age had only additive effects on grandoffspring body condition and survival. We found only a GM age effect on granddaughter reproduction. We found no interaction between GM age and M age effects for all the tested parameters (i.e., no M age x GM age interaction in Tables 1 to 4, and Table S3). Therefore, we did not detect a cumulative effect of maternal age effects over multiple generations as in the pioneering study of [START_REF] Lansing | A transmissible, cumulative, and reversible factor in aging[END_REF].

Our study illustrates that the effects of age at reproduction can persist for up to two generations in a wild population, with age effects on grandoffpsring body condition, survival and reproduction. Age effects may be explained by individual consequences of ageing (maturation, senescence and terminal investment) or by the age-specific disappearance/appearance of different phenotypes at the population level (e.g., selection) [START_REF] Forslund | Age and reproduction in birds -hypotheses and tests[END_REF][START_REF] Pol | Age-dependent traits: a new statistical model to separate within-and between-individual effects[END_REF]. In this study, we did not have enough data to perform longitudinal analyses with repeated measurements of the same grandmothers at different ages. We also did not have the possibility to use individual estimates of longevity of mothers and grandmothers as covariates to control for selective disappearance related to maternal and grandmaternal ages (e.g., [START_REF] Hayward | Reproductive senescence in female Soay sheep: variation across traits and contributions of individual ageing and selective disappearance[END_REF]. For this reason, we analysed only the patterns at the population level and we cannot formally distinguish between individual and population effects. A previous study in the common lizard supports the hypothesis of responses at the individual level for at least a part of the grandmaternal age effects we observed. This study performed analyses on longitudinal data with repeated measurements of the same mothers at different ages [START_REF] Massot | An integrative study of ageing in a wild population of common lizards[END_REF] and found relationships between maternal age and offspring traits at birth (body size and body condition). However, we cannot know if these individual responses at the first generation span the next generation. We also cannot rule out the possibility that these individual responses were modified or replaced by a differential mortality among offspring of the first generation (hypothesis of selective disappearance at the population level).

Yet, whatever the levels of action, this study shows that variation of the age at reproduction had effects lasting for up to two generations. Although it would be interesting to understand the mechanisms, finding an overall GM age effect is a first and necessary step. This shows that GM age effects are a reality in wild populations that can affect population dynamics, and that they are not just a human specificity or a laboratory curiosity. This is consistent with a study on semi-captive Asian elephants [START_REF] Reichert | Maternal age at birth shapes offspring life-history trajectory across generations in long-lived Asian elephants[END_REF] in which it has been shown that grandoffspring of old grandmothers had reduced survival [START_REF] Reichert | Maternal age at birth shapes offspring life-history trajectory across generations in long-lived Asian elephants[END_REF]. Our study found also that male and female grandoffspring of old grandmothers had reduced survival. Maternal effects were also observed in elephants. Offspring of older mothers had reduced survival and higher reproductive success. In addition, it was suggested a quadratic effect of maternal age on offspring body condition. Because offspring of older mothers had reduced survival, the authors discussed the hypothesis that the differential mortality between F0 and F1 generations led to the GM age effect at the F2 generation [START_REF] Reichert | Maternal age at birth shapes offspring life-history trajectory across generations in long-lived Asian elephants[END_REF]. As in our study, it appeared challenging to discriminate between individual and population level mechanisms.

In our study, we analysed long-term GM effects on grandoffspring survival and reproduction. The effect of GM age on survival was globally negative with a decrease of grandoffspring survival when grandmothers were older than four years old (Figure 3). For the fecundity of grandaugthers, the effect of GM age was positive. Litter size and litter success of granddaughters increased with increasing GM age (Figure 4). To understand the combined effects of GM age on survival and fecundity, we compared the multiplication rates of granddaughters produced by grandmothers of different ages. The overall pattern shows a non linear trend with a maximum of fitness for the granddaughters of grandmothers of three and four years old (Figure 5). Even if it is difficult to conclude on the adaptive value of the GM effects detected since we do not know if they are at the population or individual levels, these GM age effects seem concordant with individual responses related to maturation in young grandmothers and senescence in old grandmothers.

In conclusion, we found evidence that the age of grandmothers at reproduction (F0 generation) has effects on grandoffspring (F2 generation) in a wild population. These effects that persist for up to two generations induce time lags in the response of populations to selective pressures, and may facilitate or hinder adaptive evolution [START_REF] Kirkpatrick | The evolution of maternal characters[END_REF][START_REF] Mcglothlin | The contribution of maternal effects to selection response: an empirical test of competing models[END_REF]. Our study provides two useful avenues for future research. First, it will be important to understand the mechanisms underlying the grandmaternal age effects, especially determining if they are individual or/and population responses. Second, it would be interesting to test whether the age of the grandfathers at reproduction may also have effects in wild populations, as found in humans [START_REF] Eisenberg | Delayed paternal age of reproduction in humans is associated with longer telomeres across two generations of descendants[END_REF][START_REF] Eisenberg | Older paternal ages and grandpaternal ages at conception predict longer telomeres in human descendants[END_REF]. S3) as illustrated here in grandoffspring females (panel A) and males (panel B). Left panels show the survival rates estimated with a range of 95% confidence interval less than 0.5. Right panels show the range of 95% confidence interval of survival rates for all the combinations of grandmaternal and maternal ages, with the range limit for the value of 0.5 indicated by the dashed line.

Figure legends Figure 1 .

 legends1 Figure legends

Figure 2 .

 2 Figure 2. Grandmaternal age effect on grandoffspring body condition at birth.

Figure 3 .

 3 Figure 3. Grandmaternal age and maternal age effects on grandoffspring survival

Figure 4 .

 4 Figure 4. Grandmaternal age effects on granddaughter reproduction at two years old.

Figure 5 .

 5 Figure 5. Age-dependent multiplication rates of granddaughters. Multiplication rates

  Figure 1

Acknowledgments:

We thank all students and colleagues who helped collecting data over the years, the Parc National des Cévennes and the Office National des Forêts for field work facilities. This work was supported by the Agence Nationale de la Recherche (grant 07-

Data availability statement: Data deposited in the Dryad Digital Repository: https://doi.org/10.5061/dryad.9ghx3ffjv, (Bleu et al., 2021).

Conflict of interests:

The authors declare no conflict of interest.

Authors' contributions: All authors conceived the ideas. MM collected the data. JB analysed the data and led the writing of the manuscript. All authors contributed critically to the drafts and gave final approval for publication.

WIREs Computational Statistics, e1533. https://doi.org/10.1002/wics.1533 Surget-Groba, Y., Heulin, B., Guillaume, C.-P., Thorpe, R. S., Kupriyanova, L., Vogrin, N., … Smith, N. (2001).

Intraspecific phylogeography of Lacerta vivipara and the evolution of viviparity. Molecular Phylogenetics and Evolution,18(3)


Ionel Muguruel

Dimitra Jitea

George Gaki

Serafeim Vlahos

Irina Felekis

James Herzon

Maite Moran

Tristan Puig De Morales

Yanka Berchoux

Marija Kazakova-Mateva

Roglić

Herzon

Jitea Im

Puig De Morales

The assessment of the education needs and gaps for the sustainable development of the EU peripheral rural areas

établissements d'enseignement et de recherche français ou étrangers, des laboratoires publics ou privés.

INTRODUCTION

Peripheral rural areas have been largely studied in the last decades due to their contributions to biodiversity preservation in the EU, despite the fact that there is no agreed common EU definition for them [START_REF] Jonard | Delimitations of rural areas in Europe using criteria of population density, remoteness and land cover[END_REF][START_REF] Féret | Framework providing definitions, operational typology and review of EU strategies for rural areas[END_REF]. Currently, the OECD typology has more statistical importance and not real public policy implications. The OECD typology is mainly based on population densities and it is highly sensitive to the size of the geographic units and the classification thresholds. There were some attempts to improve the OECD typology using peripherality and a land cover index such as to define peripheral rural areas [START_REF] Jonard | Delimitations of rural areas in Europe using criteria of population density, remoteness and land cover[END_REF]. In that approach a peripheral rural area is located "45 minutes' travel time to reach an urban centre with at least 50 000 inhabitants" and it is "an open space commune if at least 75% of its area is covered by forest, agricultural or natural areas".

Nevertheless, these definitions remain too broad because they do not integrate the physical natural constraints and limitations that farmers and territories must face in some specific areas like mountains, other areas facing natural or other specific constraints or protected areas like natural parks or Natura 2000 sites. These areas have specific particularities in terms of farming practices and activities that support the conservation of the last biodiversity rich spaces in Europe. Some of them are recognised as being High Nature Value (HNV) areas supported under the second CAP pillar by agri-environment measures. Usually, such areas also achieve the geographical and land cover limits established above. But what is important from the educational perspective, is that graduates and stakeholders working in those areas (mountain, HNVf, natural parks or Natura 2000 sites) must deal with recognised problems (intensification or abandonment) that alter biodiversity. For these reasons, in the present study, a region was considered to be a peripheral rural area if it faces structural weakness due to the agri-environmental constraints created by mountains, HNV farming practices, natural park or Natura 2000 sites.

The intellectual output 1 (IO1) is the starting point of the Erasmus+ project RUR'UP on "Innovative education for sustainable development in peripheral rural areas''. It summarises the findings of two tasks implemented between October 2020 and March 2021 in 7 EU Member States (Figure 1).

Objectives:

1. Assess the content of the potential programs in Bulgaria, Croatia, Greece, Finland, France, Ireland, and Romania (curriculum); 2. Identify the tools used in the educational programs/curricula emphasizing any blended learning approaches, transnational exchanges initiatives and direct practical training provision by practitioners; 3. Provided skills and the number and targeted students group. Task 1.2. Expand the understanding of the education and skills needs of the professionals activating in the peripheral rural areas (farmers, advisory, small industry, NGOs, public sector, etc.)

METHODOLOGY

Review the content of the tertiary education programs

RUR'UP partners used structured internet searches to locate web-based information about bachelors' and masters' programs developed in each partner Member State (Bulgaria, Croatia, Greece, Finland, France, Ireland, and Romania) that potentially target the sustainability of peripheral rural areas. Official governmental lists providing the Higher Education Institutions (HEI) accredited to function in each Member State (eg. Romanian Ministry of Education & Research https://www.edu.ro/institutii-invatamant-superior) were used. From such lists, the institutions that potentially target the peripheral rural areas and those that target related rural programs and curricula were selected. These had to provide education in disciplines relevant to the four High Nature Value (HNV) innovation types -regulatory; technical; social; market -identified in the HNV-Link project (http://www.hnvlink.eu/) as important for the sustainability of such areas. The disciplines were life sciences; agricultural/ agronomic sciences; food sciences; environmental sciences; economic sciences; social & political sciences.

The official websites of the institutions previously identified were reviewed to spot the potential bachelors' & masters' programs that currently address the challenges of the peripheral rural areas. The review investigated two main challenges established by the United Nations as Key Sustainable Development Goals (https://www.un.org/sustainabledevelopment/sustainabledevelopment-goals/ using common definitions:

1. Decent work and economic growth: graduates learn to improve economic resilience of farm and territory through diversification, technological upgrading and innovation, focusing on high value added and labour-intensive products and sustainable tourism based on local culture and products;

2. Life on Land: graduates have classes about sustainable use of resources and halt the biodiversity losses;

The review also used common definitions in identifying the role of the peripheral rural areas in the HEI catchment area and in the identification of the academic degree's levels:

1. Definition of the catchment area: the geographical area from which an institution (University or High Education Institution) attracts a population that uses its services. If there are peripheral rural agricultural areas within 50 km of an institution (peripheral rural agricultural area = mountain, HNVf area, natural park or Natura 2000 site) it is considered that the institution has peripheral rural areas in its catchment area.

2. Academic degree: there are important differences between the EU national education systems. To have common grounds of comparison, it was assessed the number of years after the baccalaureate. Eg: Bachelors' -(BAC +3); Masters' -MSc (BAC +5). The review also aims to identify specific skills using common definitions developed from the Korn Ferry (2017) Competencies development guide:

1. Self-direction: graduates solve case-studies or projects where they manage goals and time and go beyond basic curriculum requirements such as to explore and to expand their own learning; 2. Organization: graduates learn how people and organisations function in agricultural rural areas, focusing mostly on peripheral areas. They have multi-disciplinary knowledge in farm and food technologies, socio-economy and environment.

3. Co-operation: build partnerships and work collaboratively with others to meet shared objectives. Graduates work in group case-studies or projects that involve multiple stakeholders (colleagues; educators; industry partners). There are transnational exchange initiatives that will be reported in the review process.

Negotiation:

Handling conflict situations effectively, facilitating consensus between parties.

Graduates learn to manage conflict by having conflict and negotiation management courses.

5. IT&C: graduates learn Information and Communication Technologie (IT&C) knowledge adapted to the new farming technologies and innovations.

6. Project management: graduates learn project management and study entrepreneurial initiatives in order to be able to generate self-sufficient incomes, develop local high-value products or households' diversification in an environmental friendly approach.

7. Practical knowledge: graduates solve real-life problems and receive direct practical training from practitioners.;

8. Teaching: graduates develop teaching and guidance skills appropriate for advisory and training activities in the peripheral rural areas. They use blended learning techniques which combine online educational materials and opportunities for interaction with traditional place-based classroom methods. 9. Legislation: ability to apply legislation changes in environmental requirements, farm management and market regulations, etc.

10. Acting in a multicultural environment: ability to interact effectively with people from different cultures and to successfully manage intercultural situations. Graduate courses include international exchange programs dedicated to students; they also feature lectures with teachers coming from abroad; it is a program taught in an international language that attracts students from different countries.

Reporting was done in an on-line shared document using a common reporting guide.

Methodology used for better understanding the needs of professionals from peripheral rural areas

RUR'UP project used focus groups or/and surveys to expand the understanding of the educational and skills needs of the practitioners. Primarily these targeted the practical ecosystem previously developed at the local level during the course of the HNV-Link H2020 thematic network, such as farmers' associations, local action groups, AgroClusters, advisory services, educators, trainers' providers, etc. It investigated the needs in several cross-education themes: 1. Social and institutional innovation skills; 2. Regulations and policy needs; 3. Products and markets; 4. Farming practices and management; 5. Soft skills; and, 6. Blended learning approaches making use of the IT&C tools.

Focus-groups technique

The focus groups (FG) research is a qualitative technique that aims at engaging discussions with a group of different stakeholders around a topic of interest. Such meetings can involve between 5 and 12 different stakeholders that are not necessarily a representative sample for a specific population [START_REF] Krueger | Focus groups: a practical guide for applied research[END_REF]. Bigger groups are criticised because they have the tendency to fragment. The discussions should not be longer than two hours. To implement a focus group such as to expand the understanding about the education and skills needs of the professionals active in the peripheral rural areas, all local teams followed a common protocol (Appendix 1).

Survey methodology

The partners that could not organise focus-groups due to the SARS-COV-2 lockdown sent an online survey to at least 11 and up to 20 stakeholders coming from or working in peripheral rural areas (Appendix 2). These came from actor groups of: 1. Farmers and small industry; 2. Local action groups or AgroClusters & farmers' associations representatives; 3. Advisory services, educators, or training providers. 4. Employed or residents in peripheral rural areas; 5. NGO representatives; and, 6. Governmental bodies.

RESULTS

Review the current programs and curricula developed in the EU tertiary education system for the sustainable development of the peripheral rural areas (PRA)

There were identified an important number of HEI in each partner Member State with peripheral rural areas in their catchment territory (Table 1). Although there are an important number of bachelor and master programs in the institutions that potentially target peripheral rural areas, only four member states (Croatia, France, Ireland, and Romania) reported the existence of educational programs devoted to the topics of peripheral rural areas. The number of learning activities and the European Credit Transfer and Accumulation System (ECTS) dedicated to the peripheral rural areas in the overall educational programs are marginal (below 2 %). They mostly cover subjects related to mountain areas and to a lesser extent to other peripheral rural areas like Natura 2000 or natural parks. High nature value farming is only partially covered mostly in terms of technical knowledge related to the management of natural meadows and pastures. The other areas of sustainability needs like socio-economic and environment are mostly missing from the current curriculum (Table 2).

Programs and curriculum developments to meet the United Nations Sustainable Development Goals

The two United Nations Sustainable Development goals under investigation SDG8 -Decent work and economic growth and SDG 15 -Life of land are present only marginally in the current curriculum (below 1% from the total ECTS and number of hours). The lectures covered mostly general aspects relating to farming and non-agricultural activities (Figure 2, 3 and 4). There are some specific lectures dedicated to mountain areas that cover both SDGs. HNV farming is almost missing from all partner Member States.

Curriculum developments to promote specific skills adapted to the sustainable development of peripheral rural areas

The learning activities promoted by the current curriculum only partially sustain the development of the appropriate skills adapted to the peripheral rural area's needs. Important skills like organisation, cooperation, negotiation, teaching or multiculturality are missing in the curriculum of some partner Member States. In the others, the current curriculum develops some general practical skills. There are no particular developments targeting peripheral rural areas with the exception of mountain areas (Table 3).

The needs of professionals from peripheral rural areas

There were a total of 108 answers to the survey implemented at different scales (regional/local) in the partner Member States (Table 4). Professionals (advisory, farming, NGOs, industry and government bodies) represent around 80% from the number of participants.

Table 4. Type of participants in the study in partner Member States

Advisory Educatio n Farming/Primary production

Farming associations

Food & Beverage industry

Governmen t bodies

NGOs Research Other

Bulgaria 2 1 0 3 0 2 6 3 1 Croatia 1 5 3 2 1 5 0 0 1 France 1 5 6 1 0 4 0 1 2 Greece 3 1 2 1 0 6 3 0 4 Ireland 4 0 0 0 0 2 1 1 3 Finland 2 1 2 1 1 2 1 1 0 Romania 1 3 2 2 0 0 2 0 0 Total(%)
13.0 14.8 13.9 9.3 1.9 19.4 12.0 5.6 10.2

In terms of knowledge needs, results show that future graduates should master an in-depth understanding of interaction between land use, environment and economy, followed by the

CONCLUSIONS

The report helped finding answers to the following questions about the effective implementation of the RUR'UP project:

1 What kind of learning activities are needed in order to fulfil the needs of practitioners from PRA?

Significant differences were identified between the knowledge and capacities acquired by graduates and the needs that are regarded as important by local actors and authorities in PRA.

-First and foremost, employers need graduates that can master an in-depth understanding of interaction between land use, environment, and economy-specific for peripheral rural areas. Most of the educational programs specialise graduates in one of the directions. Usually some of them target more technical knowledge (land use and environment) while others more socio-economic areas; -In the area of farming techniques and management, employers look for graduates that know how to use production techniques with respect towards the environment and biodiversity. Such learning activities targeting the peripheral rural areas are mostly missing from the HEI curricula. Graduates learn general farming techniques. In some countries mountain farming is a specific topic, and to a less extent others, such as Natura 2000 and HNV farming;

-Product diversification and collective marketing actions represent the most important knowledge needed by a future graduate who will work in a peripheral rural area. Current lectures do not integrate the specific needs of peripheral areas;

-In addition, they acquire general knowledge of policies without always being able to specialize them based on the particular needs of the local actor (eg. develop a multifunctional exploitation; develop an agritourism business or handcraft), authority (eg. design and application of a territorial Plan) in a PRA.

The gaps and deficits identified in this report will be filled with the syllabus developed through RUR'UP project.

What skills and capacities are needed in PRA?

Practitioners coming from the PRA highlighted the skills and capacities needed for the sustainable development of PRAs. They are: self-direction; co-operation; negotiation, practical knowledge or project management. Important practical skills like organisation, cooperation, negotiation, teaching or multiculturality are missing in the HEI curricula of some partner member states.

The report clarifies the specific training needs of the PRA. Therefore, the development of RUR'UP training materials aims to fill in the identified training gaps.

HNV farmland = 1. Farmland with a high proportion of semi-natural vegetation; 2. Farmland with a mosaic of low intensity agriculture and natural and structural elements, such as field margins, hedgerows, stone walls, patches of woodland or scrub, small rivers, etc.; 3. Farmland supporting rare species or a high proportion of European or World populations [START_REF] Keenleyside | High Nature Value farming throughout EU-27 and its financial support under the CAP[END_REF].

Skill: the ability to carry out complex activities or job functions involving ideas (cognitive skills), things (technical skills) and/or people (interpersonal skills) (Korn Ferry, 2017).

Education for sustainable development (ESD): empowers graduates to take informed decisions and responsible actions for environmental integrity, economic viability and a just society for present and future generations.

Blended Learning: learning techniques that combine online educational materials and opportunities for interaction with traditional place-based classroom methods.

b. Identify the main education knowledge needs and gaps targeting peripheral rural area (30 minutes): After identifying the main educational gaps, the group will be asked to rank them (the most important gap from the three classes above (technical; regulation and market) needed to take informed decisions and responsible actions for environmental integrity, economic viability and a just society for present and future generations. Main skills:

1.

2.

3.

4.

5.

Main gaps:

1.

2.

3.

4.

5.

Examples of good practices developed in your area/region: 1.

2.

3.

4.

5.

Organization: graduates learn how people and organisations function in agricultural rural areas, focusing mostly on peripheral areas. They have multidisciplinary knowledge in farm and food technologies, socio-economy and environment.

Co-operation: build partnerships and work collaboratively with others to meet shared objectives. Graduates work in group case-studies or projects that involve multiple stakeholders (colleagues; educators; industry partners). There are transnational exchange initiatives that will be reported in the review process.

Negotiation: Handling conflict situations effectively, facilitating consensus between parties. Graduates learn to manage conflict by having conflict and negotiation management courses.

IT&C: graduates learn Information and Communication Technology (IT&C) knowledge adapted to the new farming technologies and innovations.

Project management: graduates learn project management and study entrepreneurial initiatives in order to be able to generate self-sufficient incomes, develop local high-value products or households' diversification in an environmental friendly approach.

Practical knowledge: graduates solve real-life problems and receive direct practical training from practitioners.

Teaching: graduates develop teaching and guidance skills appropriate for advisory and training activities in the peripheral rural areas. They use blended learning techniques which combine online educational materials and opportunities for interaction with traditional place-based classroom methods..

Legislation: ability to apply legislation changes in environmental requirements, farm management and market regulations, etc.

Acting in a multicultural environment: ability to interact effectively with people from different cultures and to successfully manage intercultural situations. Graduate courses include international exchange programs dedicated to students; they also feature lectures with teachers coming from abroad; it is a program taught in an international language that attracts students from different countries.

Reporting

Data collection was done in an on-line shared document using a common reporting guide.

Figure 1 Task 1 . 1 .

 111 Figure 1. IO1 interconnection

Figure 2 .Figure 3 .Figure 4 .

 234 Figure 2. Examples of learning activities targeting SDG8 -Decent work and economic growth

Figure 5 .

 5 Figure 5. Skills needed by future graduates to improve the sustainability of the peripheral rural area.

c.Table 2 .

 2 Identify the main skills needs and gaps targeting peripheral rural area (30 minutes): Reporting fiche concerning skills needs and gaps targeting peripheral rural area: Questions provided by the moderator Education gaps Q6. Discuss, based on the main skills listed below, what are the main important five skills that a graduate should have to respond to the problems of peripheral rural areas and that are currently missing. Provide us some examples of good practices developed in your region. Self-direction: graduates solve case-studies or projects where they manage goals and time and go beyond basic curriculum requirements such as to explore and to expand their own learning.

  Review the current programs and curricula developed in the EU tertiary education system for the sustainable development of the peripheral rural areas 3.1.1. Programs and curriculum developments to meet the United Nations Sustainable Development Goals 3.1.2. Curriculum developments to promote specific skills adapted to the sustainable development of peripheral rural areas

	1. INTRODUCTION
	2. METHODOLOGY
	2.1. Review the content of the tertiary education programs
	2.2. Methodology used for better understanding the needs of professionals
	from peripheral rural areas
	2.2.1. Focus -groups methodology
	2.2.2 Survey methodology
	3. RESULTS
	3.1. 10
	3.2. The needs of professionals from peripheral rural areas	13
	4. CONCLUSIONS	16
	5. REFERENCES	21
	APPENDIX 1 -RUR'UP FOCUS GROUPS RESEARCH PROTOCOL	22
	APPENDIX 2 -RUR'UP SURVEY RESEARCH PROTOCOL	25

Table 1 . Descriptive statistics

 1

		Bulgaria	Croatia	Greece	Finlan	France	Ireland Romania
					d
	Number of HEI	7	13	11	2	60	17	14
	Number of programs	14	24	34	6	150	67	79
	bachelor	12	16	7	2	78	39	43
	master	2	8	27	4	72	28	36
	Programs targeting peripheral	0	14	1	0	18	5	12
	areas
	bachelor	0	9	0	0	5	5	5
	master	0	5	1	0	13	0	7

Table 2 . Examples of learning activities targeting PRA

 2

			Bulgaria	Croatia	Greece	Finland
	Number	of	14	24	34	0
	programs
	Programs with	3	24	2	0
	topics targeting
	PAA (%)
	Topics
			Governance of	Hunting	Spatial planning and	No specific
			the territory and	development	development of mountain and	topic
			development	Endangered	disadvantaged areas
			Protected Areas	habitats of the	Monitoring of mountain
			Management	Republic of Croatia	environment
				Management of	Spatial economic, social, legal
				protected areas	dimensions of the development
				Responsible	and the environment of the
				tourism	mountain areas
					Introduction to the Environment
					and Society of Mountainous
					Areas
					Environmental Economics:
					Applications in Mountain
					Development Issues
					Environmental Management with
					ecosystemic approach and
					emphasis in mountainous areas
					Identity and development
					prospects of mountain areas

Table 2 . Examples of learning activities targeting PRA (continued)

 2

			France	Ireland	Romania
	Number	of	150	67	79
	programs
	Programs	with	26	5	12
	topics targeting PRA
	Topics		Mountains as challenging areas	Applied Ecology and	Protection of mountain
			Human-environment	Environmental	ecosystems
			interactions in mountains	Management; sections	Protected natural areas

Table 3 . Skills development

 3

		Bulgaria	Croatia	Greece
	Self-direction No specific learning	Direct sales and	Grassland and meadow
		activities	processing on the farm	management
				Fauna of natural ecosystems
				etc
	Organisation No specific learning	No specific learning	Organizational behaviour
		activities	activities	and human resource
				management
				Biodiversity Conservation
				and Exploitation
				etc
	Co-operation Economics of	No specific learning	Theory and methods of
		cooperatives	activities	participatory planning
				Environmental
				management with
				ecosystemic approach and
				emphasis in mountainous
				areas
	Negotiation	Ecological conflicts	No specific learning	No specific learning
		management	activities	activities
	IT&C	Electronic systems in	Biomimicry-solutions	E-entrepreneurship and E-
		agricultural	inspired by nature	marketing
		machinery		Geographic Information
		IT&C in agribusiness		Systems
	Project	Project management Natural pasture	Project management and
	management		sustainable	investment projects
			management	management
			Farm management and	Management plans
			agricultural products	etc
			marketing
			Management and
			entrepreneurship in
			agriculture
	Practical	Mechanisation	Farm management and	Grassland and meadow
	knowledge	Introduction to	agricultural products	management
		agriculture	marketing	Collective project and field
		Principles of	Nutrition of domestic	studies
		agriculture and	animals

Table 1 . Reporting fiche concerning education needs and gaps targeting peripheral rural area: Questions provided by the moderator Education gaps Q3. What do you expect from a future university graduate active in a peripheral rural area, in terms of farming techniques and farming management knowledge? What is currently missing?

 1

	Focus on:

• Sustainable use of resources and halt the biodiversity losses. • IT&C needs. • Existing knowledge about machines & technologies targeting peripheral rural areas. • Animal & plant breeds.

Q4 What do you expect from a future university graduate in terms of knowledge about public policy targeting peripheral rural areas? What is currently missing?

	Focus on:
	• Finding the most important type of knowledge that they need (cross-
	compliance rules; agri-environmental measures; assess specific rural
	development measures etc);
	• Other
	Q5

. What do you expect from a future university graduate active in a peripheral rural area, in terms of products and markets knowledge? What is currently missing?

	Focus on:
	• product diversification;
	• develop high-value added and labour-intensive products;
	• develop tourism services based on local culture and products;
	• marketing
	• Other

knowledge of specific rules and regulations that target the sustainability of peripheral rural areas (Table 5). As it concerns farming techniques and management, a future graduate should have better knowledge competences in areas of environmental protection. Also, knowledge related to farming techniques appropriate to halt the biodiversity losses from farming activities are a must (Table 6). Future graduates should master better knowledge about environmental regulations (crosscompliance rules; agri-environmental measures) and to know how to implement specific rural development intervention measures such as to valorise the rich biodiversity resources specific for the peripheral rural areas (Table 7).

Table 8. Products and markets knowledge competences expected from graduates to improve the sustainability of the rural peripheral area.

Product diversification (on farm activities)

Marketing of farm products

Collective marketing actions

Understandin g consumer needs APPENDIX 1 -RUR'UP FOCUS GROUPS RESEARCH PROTOCOL 1. Pre-select stakeholders

The group was composed of at least two participants from each stakeholder type: 1. Farmers and small industry; 2. Local action groups or AgroClusters and farmers' associations representatives; 3. Advisory services, educators, or training providers. 4. Employed and/or resident in rural areas; 5. NGO representatives; and, 6. Governmental bodies (at local/regional level). Such stakeholders should work or previously have worked in a peripheral rural area.

Select date, hour and place for the meetings

Meetings were organised on-line or face-to-face according to the COVID-19 pandemic situation. In the implementation phase, all partner countries were in lockdown due to the SARS-COV-2 virus thus the focus-groups meetings were organised on-line. Before the meetings email invitations have been sent to all identified participants explaining the main purpose of the meeting: to identify education and skills needs of the professionals active in the peripheral rural areas.

Prepare the meeting

Each local team designated a moderator for animating and conducting the meeting. Also, a supervisor checked that all topics from the interview guide were covered during the meeting. One team member recorded the main answers on a standardised fiche. Meetings were recorded such as to use the transcripts afterwards to check and to complete the in-situ notes.

Meeting

The moderator used open-ended questions arranged in a logical sequence in a common preestablished discussion guide. At the beginning of the meeting, the moderator presented the main goals of the meeting followed by a presentation of all participants (approx. 10 minutes). After collecting the answers to introduction questions (Q1&Q2), the moderator clarifies the meaning of the terms to be used later into discussion in order to avoid the possible insecurity of the participants as suggested by [START_REF] Fischer | Understanding mental constructs of biodiversity: implications for biodiversity management and conservation[END_REF].

Focus group discussion

For the purposes of the project we consider as:

Peripheral rural areas = regions affected by structural weakness due to the agri-environmental constraints created by mountains, HNV farming practices supporting biodiversity, natural park or Natura 2000 site.

Mountain area = regions characterized by a considerable limitation of the possibilities for using the land and by an appreciable increase in production costs due to altitude that shorten the growing season or the presence of slopes too steep for the use of machinery or requiring the use of very expensive special equipment; areas north of the 62nd parallel (EU Regulation 1305/2013).

Socio-economic profile:

Question 10. What is your highest degree in education?

• Vocational school in agricultural related areas;

• High school;

• University, not in agricultural related areas;

• University in agricultural related areas;

• Other?........... Question 11. How many years since your highest degree in education?

• Less than 5;

• 5 to 10; • 10 to 20; • More than 20; Question 12. Your age?

• 20 years to 30 years;

• 30 years to 50 years;

• More than 50 years;

II.3. Reporting:

Data collection was done in an on-line shared document using a common reporting guide.