The Eurasian lynx Lynx lynx once colonised all Continental Europe but was ousted gradually with the decrease of forests and wildlife on the one hand and the growth of the human population, the expansion of cultivated land and the increase of livestock. The final eradication of the remnant populations happened in the course of the 19thcentury. The exceptions were the populations in the northeastern European lowland, the Carpathians and the southern Dinaric Range, which all reached a minimum in the late 1940s, but eventually survived (for a review of the historic downfall and the source literature see Breitenmoser & Breitenmoser-Würsten 2008). Reintroduction programmes started almost 50 years ago, mostly, butnot exclusively using funder animals from the Slovakian population. To date, Eurasian lynx were reintroduced in Continental Europe in France, Switzerland, Germany, Poland,the Czech Republic, Italy, Austria, and Slovenia (see contributions in this Special Issue). Some reintroduction projects failed at an early stage, and all emerging reintroduced populations are still relatively small, mostly isolated and show a rather high degree of inbreeding, inter alia because of the limited number of funder individuals (Breitenmoser & Breitenmoser-Würsten 2008).

At present, Continental Europe hosts 3 small to medium size autochthonous and 11 tiny to small reintroduced populations distributed over 23 countries (von Arx et al.2021). In recent years, new reintroduction projects were initiated (e.g. Idelberger et al. 2021, Tracz et al. 2021), efforts to connect distinct populations were made (Molinari et al. 2021) and the genetic remedy of earlier reintroduced populations was discussed and – in one case (Fležar et al.2021) – already launched. Such projects require access to adequate source animals and the transport of translocated lynx across international borders. According to the IUCN Guidelines for Reintroductions and other Conservation Translocations (IUCN/SSC 2013), but also to EU and/or national legislation, such conservation interventions require the identification of the adequate source population,an evaluation of the conservation status of the source population, and veterinary health precautions to prevent the transmission of pathogens. However, the “traditional”source population of the Carpathian Mountains and mainly from Slovakia has its own conservation issues (Kubala et al. 2021), which triggered (again) the discussion on the use of conservation breeding programmes as a source for reintroductions (Lengger et al. 2021). Eventually, all these now isolated populations should be merged into few large and viable metapopulations to mitigate the negative effects of habitat fragmentation (Premier et al. 2021). But to complicate the picture, continental Europe hosts three phylogenetically distinct lines of Eurasian lynx, recognised as valid subspecies (Kitchener et al. 2017). Hence, is any lynx welcome anywhere? How shall we delineate areas of subspecies in regions where we have no information on the original inhabitants?

Such questions and the complex situation call for a consensual strategy for the long-term goals for the recovery of the Eurasian lynx, for agreed standards and protocols facilitating the international and inter regional cooperation,and for common guidelines and a coordinated approach to lynx conservation in Continental Europe. Topical challenges are e.g. (1) the delineation of conservation units, (2)the genetic remedy of inbred populations, (3) the source populations for further translocations, (4) the connectivity of small populations, and (5) the management of lynx populations with regard to their coexistence with people.On 16–19 June 2019, some 50 lynx experts (Fig. 1, Appendix I) gathered in Bonn to discuss the conservation of the Eurasian lynx in Continental Europe. The participants of the workshop aimed (1) to review the conservation status of continental lynx populations and the implementation of conservation projects, (2) to discuss recommendations fora coordinated long-term approach to lynx reintroduction and conservation across Western and Central Europe, and(3) to agree on the development of standards and shared protocols for the practical conservation work. The proceedings of the symposium are compiled in this Special Issue.After the review, the participants developed Recommendations(Bonn Lynx Expert Group 2021) to help coordinating the conservation of the lynx in Continental Europe.The results from the Bonn Workshop were submitted to the Secretariat and the Standing Committee of the Convention on the Conservation of European Wildlife and Natural Habitats (Bern Convention) of the Council of Europe,which on 6 December 2019 adopted the Recommendation No. 204 (Standing Committee 2019), which are congruent to the Recommendations presented at the end of this Special Issue (Bonn Lynx Expert Group 2021).

Continental Europe, in the context of the following Proceedings and Recommendations, refers to the historic and present distribution range of the Eurasian lynx south of the large autochthonous populations of Fennoscandia and Russia. The Bonn conference concentrated on the biological and ecological aspects with regard to the recovery of viable lynx metapopulations in this region. Although we are fully aware of the importance of the human dimension aspects of such an endeavour, we were, for practical reasons,not able to address also the social science aspects of lynx conservation. However, all participants agreed that the discussions in Bonn should be continued and that further topics of lynx conservation in Continental Europe need to be addressed in the future.

The Bonn lynx conference was jointly organised by the HIT Umwelt- und Naturschutz Stiftung, the Stiftung Naturund Umwelt (SNU) Rheinland-Pfalz, the IUCN SSC CatSpecialist Group, and the Foundation KORA. Financialsupport was generously provided by the HIT Umwelt- und Naturschutz Stiftung, the SNU, and the Council of Europe (Bern Convention).

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With a total of 17,000–18,000 individuals including European Russia, the Eurasian lynx Lynx lynx is listed Least Concern at the European level in the IUCN Red List. However,some of the larger autochthonous populations in Scandinavia and the Baltics have shown declining trends in the past decade, and the Balkan lynx subspecies was assessed as Critically Endangered. The reintroduced populations in West and CentralEurope remain small and are classified as Endangered (Alpine, Jura, Dinaric) or Critically Endangered (Vosges-Palatinian, Bohemian-Bavarian-Austrian, Harz). We present each of the populations regarding population numbers, range and threats and provide the justification for the respective Red List classifications.

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The north-eastern European lowland population of the lynx is commonly attributed to the “Baltic population” and believed to belong to the nominative subspecies Lynx lynx lynx. Geographically, its range extends from eastern Poland throughout the Baltic states north to Fennoscandia and east to central Asia. Within its range across north-eastern Europe the status of the species differs among the countries from fully protected year-round to a hunted species. The population is exposed to a varying degree of fragmentation with most severe habitat fragmentation in the south-western part of its range and well connected forest habitats in the north-east. The population is genetically structured, most likely as a result of habitat fragmentation. The genetically poorest subpopulation occurs in north-eastern Poland, which is most isolated from the remaining part, and the highest genetic diversity characterises the Latvian and Estonian lynx. The genomic data, however, confirm that lynx within the north-eastern European lowland population harbours a mitochondrial phylogeographic sublineage differentiated from the remaining central (Carpathian) and south-European populations, though its taxonomic value is still unclear. The lynx are facing various types of threats in different countries. Legal protection is not a sufficient measure to warrant the population’s demographic security, as it is exposed either to the lack of suitable and well connected habitat, forest logging, poaching or illegal amber mining. Hunted populations may be subject to excessive quotas. The challenges of lynx conservation include restoration of the suitability and connectivity of the habitat, reintrodutions, reconsideration of hunting quotas, establishing non-invasive robust population monitoring, and increasing public awareness about the lynx conservation needs.

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The population of Eurasian lynx in the Carpathian Mountains is one of the largest in Europe, with a total population size of ~2,100–2,400 individuals. However, the status of the species in the Carpathians was based solely on “expert opinions”, while relevant scientific data were restricted. Recent research indicated that these figures are not reliable and strongly overestimate the population size. Exaggerated data and misleading information on the status and trend of the lynx population have fostered conflicts between the lynx and human interests, and ultimately leading to illegal killings.Negative attitude of hunters towards lynx originates in a belief that the predator is responsible for the alleged decrease of roe deer populations in Slovakia.Moreover, illegal killing could have some synergetic effect with the development of traffic infrastructure, which increasingly disrupts the connectivity between suitable habitats and exacerbates human-induced mortality. Carpathian Mts. have been and still are a source for lynx reintroduction and reinforcement projects and are of great importance for the large-scale conservation of lynx in Europe. Authorities in charge,lynx experts and interested groups from the Carpathians should jointly establish a standardised robust population monitoring and seriously mitigate anthropogenic factors jeopardising lynx survival. A sound cooperation between all countries sharing the Carpathian population for the conservation and management of the lynx is required. We recommend in particular the adoption of a jointly developed Pan-Carpathian conservation and management strategy and related national action plans.

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The Balkan lynx Lynx lynx balcanicus is a subspecies of the Eurasian lynx distributed in the south-west Balkans, with relict populations in the Mavrovo National Parkand surroundings in North Macedonia and the Munella Mountains in Albania, and with single individuals in Bjeshkët e Nemuna, western Kosovo. In 2015 the Balkan lynx was assessed as Critically Endangered in the IUCN Red List. Main threats involvesmall population size, limited prey base, habitat degradation, and poaching.The assessment was done on the basis of 10 years of lynx research and monitoring in the range countries through the Balkan Lynx Recovery Programme. This transboundary project strives to (1) create capacities for a long-term conservation project,(2) monitor and study the extant population, (3) understand local people's attitudes towards lynx and other large carnivores, and engage them in conservation efforts,and (4) establish a protected-area system for the benefit of the Balkan lynx and its prey. The ongoing project is focusing on diminishing the main threats to the Balkan lynx and engaging with stakeholders and local people with regard to awareness raising, knowledge gathering, improving conservation policies, and site protection.

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The development of transboundary cooperation in monitoring and conservation of the Bohemian-Bavarian-Austrian lynx Lynx lynx population is presented. From 2013–2020 two international lynx projects, co-financed by the EU, established monitoring standards and collected comprehensive demographic and genetic data on population level. Main threats for the population are illegal killing and inbreeding.

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Eurasian lynx Lynx lynx was reintroduced to the Dinaric Mountains in 1973 to bring back an extinct autochthonous keystone species, but also to enrich the hunting grounds. The institutions involved in the reintroduction were aware of the danger of genetic isolation and warned about the importance of connectivity between the lynx populations in the Dinaric Mountains and the Western Alps for their long-termstability. Unfortunately, these populations never connected and the once thriving Dinaric population declined dramatically. Today, lynx in the Dinaric Mountains and the neighbouring south-eastern Alps are again on the brink of extinction. To prevent this,reinforcement of the population is currently taking place within an EU-funded “LIFE Lynx” project. While the main threat is indisputable – the population is highly inbred– the relative importance of the other factors causing the decline is still under debate,especially because the reintroduced lynx in the Dinaric Mountains was legally hunted for more than two decades. Here, we describe the most important historical events which enabled the lynx to recolonise Slovenia, Croatia, Northeastern Italyand Bosnia and Herzegovina (BiH) and highlight the main management decisions that were made during the lynx population expansion. Since the hunters were the promotors of the reintroduction efforts but had also carried out all the hunting activities,we discuss also the relevance of hunting for the lynx population development in the region.

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Between 2000 and 2006, 24 zoo born lynx Lynx lynx have been released into the Harz Mountains HM in central Germany. In the monitoring year 2010/11, 25 cells of the EU reference grid were occupied by lynx belonging to the Harz Lynx Population HLP. In the same season, the first reproduction outside the HM was proven. Until the monitoring year 2018/19, the number of cells of the EU monitoring grid occupied by the HLP has increased to 84. This represents an average annual increase of 7.4 grid cells.Although the mountain range is surrounded by major roads and landscapes with low forest cover, reproducing females have established territories in five different areas outside the HM. Telemetry, genetic and photo data show that single male dispersers can be found in distances of up to 309 km from the source population in the HM,whereas reproducing females have not been proven further than 100 km away from the population centre and a female without cubs has been reported in a maximum distance of 143 km.

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The present report describes the situation of the Eurasian lynx Lynx lynx in the Jura Mountains Mts, shared by France and Switzerland. The species disappeared between the 17th and 20th century and recolonised its natural range from the 1970s following reintroduction in Switzerland. The current distribution in the region covers a total of 13,700 km², including 4,200 km² with “hard fact” evidence of reproduction(data of the biological year 2018/19). In France, the area of regular lynx presence is increasing and consolidates particularly in Franche-Comté region, while almost allsuitable habitat is colonised in the Swiss Jura. Between 2004 and 2020, we documented sixteen dispersal movements to adjoining mountain ranges includingthe Alps (9), the Black Forest and the Schwäbische Alb (6), the Vosges Mts (1) and two to the Swiss Plateau. The Jura Mts could probably form a larger functioning meta-population with the adjoining Vosges-Palatinate Forest, the Black Forest and Schwäbische Alb, and the Alps, although difficult to achieve without reinforcement. The lynx population of the Jura Mts is likely threatened to various extents by traffic accidents, conflicts with human activities and persecution. Furthermore, coordinated health surveillance and genetic variability monitoring are needed to secure population viability.

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The Eurasian lynx Lynx lynx became extinct in the Vosges Mountains in the early17th century. Between 1983 and 1993, 21 lynx were released during a reintroduction program. However, only 10 lynx survived notably because of illegal killings. Besides,tensions appeared between local stakeholders: the return of the lynx in the Vosges Mountains was not unanimously accepted. Almost 30 years later, the lynx is still critically endangered in the Vosges Mountains and acceptance remains still fragile.Thus, its conservation status needs to be improved urgently. This is a challenge for the species in the Vosges Mountains as well as for the Upper Rhine lynx metapopulation.In that context, a Regional Action Plan focusing on lynx in the Vosges Mountains has been written from 2016 to 2019 according a participative approach. A total of 18 actions answering to four topics (coexistence with hunters and breeders, ecological connectivity, monitoring, awareness) have been identified. The implementation phaseof this action plan will occur during 10 years, from 2020 to 2029, to restore durably the lynx conservation status in the Vosges Mountains.

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In the reintroduction programme in the Palatinate Forest – as part of the French/German biosphere reserve Palatinate Forest / Vosges du Nord – a total of 20 wild born lynx Lynx lynx from Swiss and Slovakian origin were released until 2020 to form a nucleus for the further expansion of the subpopulation. The reintroduction was accompanied by involvement of stakeholders, a management plan, environmental education, and public relations. Stakeholders, including hunters and livestock keepers,predominantly supported the reintroduction. The reintroduction has been indispensable to re-establish a subpopulation of lynx in its formerly natural range in the Palatinate Forest. The subpopulation will contribute to the conservation of the species, which is classified as critically endangered in the Red List of threatened species in Germany (Meinig et al. 2009) and is present in Germany and France only in few segregated, more or less isolated occurrences.

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Although the lynx Lynx lynx has been eradicated in north-western Poland several hundred years ago, the currently conducted analysis of habitat suitability shows that in this part of Poland there are habitats suitable for lynx and prey populations no smaller than in the eastern part of the country. The plan for the reintroduction of the Eurasian lynx to western Pomerania is implemented by the Western Pomeranian Natural Society and financed under the Infrastructure and Environment Program 2014-2020. The approach is to use a combination of soft release of captive-born lynx and possibly translocation of wild lynx from the north-east European lowland population.

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The discussion between different stakeholders about a reintroduction of the lynx Lynx lynx in the Black Forest has been going on since the 1980s. Since the installation of the Working Group Lynx Baden-Württemberg (WG lynx) in 2004, the knowledge about the lynx and the conflicts behind it has been improved and distributed for all stakeholders represented in the WG lynx. The joint development of a technical concept for dealing with the lynx helped to develop concrete solutions for identified concerns of the stakeholders. After the state elections in spring 2021, the governing parties stated in their coalition agreement that the chances for the return of the lynx should be improved through a reinforcement program to support the population in close cooperation with all stakeholders concerned.

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Assuming that large-scale reintroduction projects within the Alps or anywhere else in Western or Central Europe will be difficult to implement in the future, the stepping-stone approach to connect small and isolated populations seems an adequate strategy. Experiences indicate that the number of animals released in the frame of an adaptive release strategy (monitoring and further releases if needed) as well as distance to the nearest lynx population might be the crucial elements defining the success of these small-scale reintroductions.

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The project Status and Conservation of the Alpine Lynx Population SCALP is an ongoing programme aiming to coordinate the Eurasian lynx Lynx lynx monitoring, conservation and management activities in the Alps, but the monitoring approach has recently been expanded to the neighbouring Dinaric and Jura Mountains. The long-term goal of the SCALP is to help the still small and isolated reintroduced populations to expand and to recover in co-existence with people. The process is advanced and supervised by the SCALP Expert Group, which also prepares yearly distribution maps.

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For strictly protected species, the European Union’s Habitats Directive obliges member states to keep or head for a favourable conservation status FCS. Within the“3Lynx-Project”, 11 partners from 5 countries developed a population level based conservation strategy. To operationalize the FCS-concept, we used the criterion D(population size) of the IUCN Red List category Near Threatened and translated it into specific minimum population size. For the Bohemian-Bavarian-Austrian BBA lynx Lynx lynx population we came up with 250 mature animals, being implied that connectivity to other lynx populations is simultaneously assured. The key monitoring unit is defined as the number of reproducing females. To reach FCS, a number of 165verified females with kittens within the BBA-population is targeted.

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The reintroduction of lynx Lynx lynx to the Palatinate Forest with adjacent suitable habitats in France, Switzerland and Germany spurred the interest to augment the international cooperation to exchange information, develop harmonized approaches and to establish supporting organisational structures. Thus, the Expert Committee Lynx has been established under the auspices of the Upper Rhine Conference and functions as an administrative umbrella to the protruding lynx habitats.

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Following the reintroduction and natural expansion of various Eurasian lynx Lynx lynx populations, new challenges are being faced. The current lack of genetic exchange between small populations due to restricted dispersal caused by human activities(i.e. habitat fragmentation, persecution, vehicle collisions) puts them at risk of stochastic demographic events, genetic drift and inbreeding. Low genetic diversity has been reported for most reintroduced populations and evidence of ill-effects due to genetic impoverishment have been shown in two reintroduced lynx populations. We present the pertinent points discussed on the topic of fragmentation and connectivity of lynx habitat, with a special focus on the discussions surrounding genetics and the requirements for long-term management of a potential “Central European metapopulation”. The potential metapopulation management must be based on a good knowledgebase through genetic and demographic monitoring and targeted conservation research designed to deliver prognoses required by managers. The contemporary development of a spatially explicit individual-based demogenetic simulation model has the potential to confront the future metapopulation management questions.

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EUROLYNX (European Lynx Information System) is an open, collaborative project based on a spatial database that stores shared Eurasian lynx Lynx lynx data to investigate variation in behavioural ecology along environmental gradients or population responses to specific conditions, such as habitat changes, impact of human activities,prey densities, or livestock husbandry methods. EUROLYNX aims to promote comparative theoretical and applied research into Eurasian lynx behaviour and ecology at the European scale. The open, bottom-up and cooperative structure of EUROLYNX spurs proactive engagement of partners and assures that they are involved in all stages of research. Currently, 42 groups from 19 countries have joined the initiative and nine working groups have been established to address ecological questions, to prepare research protocols and to push methodological advances forward.

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Supporting Online Material

Table T1

Switzerland has become an important source of Eurasian lynx Lynx lynx for reintroduction projects in Europe. It is now widely accepted that translocations of animals are associated with a serious health risk. Therefore, the development of multidisciplinary expertise and the elaboration of veterinary protocols are needed, which require knowledge on the health status of the source population and information on potential health risks at the release site. Here, both disease cases and carriers of potentially threatening pathogens have to be taken into consideration. In Switzerland,a range of infectious agents circulate within lynx populations apparently without associated morbidity. However,genetic analyses combined with health investigations have pointed at a possible inbreeding depression. Furthermore, unexpected health issues arose in the framework of translocations. Overall, the Swiss experiences underline the necessity of long-term health surveillance of reintroduced and small isolated wildlife populations,the usefulness of well-established veterinary protocols in the framework of translocation projects, the value of multidisciplinary collaborations and of sample archives for further analyses, and the need for adaptive management based on scientific data. For a conservation programme of the Eurasian lynx on a pan-European level, procedure harmonisation should be sought.

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Supporting Online Material

Figures F1-F3

The use of captive-born lynx for reintroduction programmes has been controversially discussed in the past, but projects such as the reintroduction in the Harz Mountains have demonstrated that zoo-born lynx can adapt to living in the wild. However, phylogenetic considerations require that the zoo-based population is known and that subspecies are bred in separate lines. EAZA has established ESBs for Lynx lynx lynx and for Lynx lynx carpathicus. In the future, these ESBs could also serve as source populations for reintroduction, provided that sensible protocols for breeding, husbandry, and training of lynx to be released to the wild are developed and followed.

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Conclusions from the workshop of the “Bonn Lynx Expert Group” in Bonn, Germany, 16–19 June 2019

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