Wölfe In Schleswig Holstein 2016

Following massive persecution und eradication, strict legal protection assisted in a successful reestablishment von wolf packs in Germany, which has actually been ongoing because 2000. Here, we describe this recolonization process über mitochondrial DNA control-region sequencing, microsatellite genotyping und sex identification based on 1341 mostly non-invasively built up samples. We reconstructed the genealogy of German wolf packs betwee 2005 and 2015 kommen sie provide die info on trends bei genetic diversity, dispersal patterns und pack dynamics during the early development process. Our results show signs des a founder effect punkt the anfang of die recolonization. Hereditary diversity in German wolves is moderate compared zu other European wolf populations. Return dispersal among packs zu sein male-biased in the sense that females are an ext philopatric, dispersal ranges are similar betwee males und females when only dispersers room accounted for. Breeding with close relatives is regular und none of the six male wolves originating from die Italian/Alpine population reproduced. However, moderate genetic diversity und inbreeding levels of the recolonizing population are preserved von high sociality, dispersal amongst packs und several immigration events. Our results demonstrate in ongoing, rapid und natural wolf population expansion in an intensively used social landscape in Central Europe.

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Wilderness areas are rapidly decreasing across the planet, while easily accessible habitats and population numbers des large mammals shrink around the world (Di Marco et al. 2014; Watson et al. 2016). Big carnivores, weil das instance, are internationally threatened due to their lethal persecution von humans in reaction to livestock predation as well as die reduction des habitats und prey access (Ripple et al. 2014). Bei the present anthropogenic age, one necessary way zu protect huge animals, including big carnivores, may be to nähren human–wildlife coexistence within die same landscapes. Ideally, together a strategy requires common coadaptation (Carter und Linnell 2016), involving effective human–wildlife conflict management (van Eeden et al. 2018).

Interestingly, the global decline an large animal populations is not ubiquitous across all regions and species. In Europe, weil das instance, several large carnivore populaces (brown be afflicted with Ursus arctos, grey wolf Canis lupus, Eurasian lynx Lynx lynx and wolverine Gulo gulo) continuously thrive due to effective conservation measures und socioeconomic transforms (Chapron et al. 2014). Although die rewilding in Europe with huge mammal species has raised considerable public und scientific interest und may offer as a instance study on die potential for human-large carnivore coexistence in human-dominated landscapes, comprehensive knowledge about ns patterns of range und population growth into human-dominated landscapes is ausblüten limited. In this regard, ns grey ein wolf Canis lupus Linæus, 1758 to represent a specifically interesting research subject. Wolves are currently recolonizing their historic ranges within numerous human-dominated landscapes bei Western und Central Europe (Chapron et al. 2014).

In Germany, zum instance, the first wolf verpackt was confirmed bei 2000 an the eastern part des the country after much more than 150 year without resident wolf (Ansorge et al. 2006). Due to the fact that then, the population expanded, und is jetzt recognized as the Central European population (Chapron et al. 2014), spreading throughout mainly north Germany (Reinhardt et al. 2019) and Western Poland (Nowak and Mysłajek 2016; Szewczyk et al. 2019), with single individuals dispersing zu Denmark (Andersen et al. 2015). Hereditary comparisons indicate that ns Central European population likely derived from long-distance dispersers from ns Baltic wolf population bei North-Eastern Poland (Czarnomska et al. 2013). Distant areas are recolonized von wolves due zu their high dispersal capacity with jump-dispersal events von over 300 km (Kojola et al. 2006; Wabakken et al. 2007; Ciucci et al. 2009; Ražen et al. 2016). Despite their potential of long-distance dispersal, a huge proportion of dispersing individuals settle within 100 km from their natal package (Kojola et al. 2006; Caniglia et al. 2014).

While there is considerable expertise about ein wolf dispersal an natural or semi-natural areas (reviewed bei Mech and Boitani 2003), few studies have documented the mechanisms of range extension and population establishment in more densely populated, anthropogenic landscapes. Couple of detailed multigenerational pedigrees documenting the expansion des wolf populaces have so far been generated and are mostly obtained from sparsely human-populated north regions such together Scandinavia and Yellowstone national Park (Liberg et al. 2005; vonHoldt et al. 2008; Granroth-Wilding et al. 2017), or in the Apennine Mountains in Northern Italy (Caniglia et al. 2014). An this study, us summarize ns results des intense genetic ein wolf monitoring during the anfangsverdacht 15 years von wolf recolonization an Germany zu reconstruct patterns des dispersal und population development into in intensively human-dominated landscape. Us hypothesized that grundlegend patterns of range expansion bei wolves would be similar bei a human-dominated landscape as bei more organic habitats. Considering ns large street (>400 km) to die source population in Eastern Poland as well as ns dispersal fads known zum wolves in other areas, consisting of Poland (Nowak und Mysłajek 2016; Szewczyk et al. 2019), us predicted that (i) we would find bei initial founder impact during ns early colonization phase and that (ii) recolonization would certainly follow a comparable process to that found in other areas (Mech and Boitani 2003). This procedure typically beginning with an initial verpackt from which die offspring disperse right into neighbouring areas. Ns local dispersers des the original verpackt then generally reproduce with newly immigrated individuals. We also expected the (iii) gene flow und dispersal amongst packs would certainly be primarily male-biased (vonHoldt et al. 2008; Caniglia et al. 2014). Moreover, we supposed that (iv) hereditary diversity would be lower und inbreeding levels would be higher des the recolonizing population compared zu larger, persistent European wolf populations (Hindrikson et al. 2017). Based on ns genealogical data, we survey trends in genetic diversity, inbreeding and population structure.


Study area and sample collection

Within die European union (EU), whereby Germany is a member state, ns wolf zu sein listed bei Annex II und IV under ns conservation legislation of the eu Habitats Directive (Council Directive 92/43/EEC), with the overall goal des reaching the ‘Favourable conservation Status, FCS’ (Article 2, board of directors Directive 92/43/EEC). Included in Article 2, ns conservation status von the wolf as priority types needs to be monitored von the member says (Article 11). Ns wolf population an Germany has been monitored because 2001. The taste monitoring methods used in Germany are existence sign surveys in combination v camera trapping and genetic analyses (Kaczensky et al. 2009; Reinhardt et al. 2015). The major objectives von the German wolf monitoring are ns annual assessment des the area von occurrence and the population size given as the minimales number von packs (including reproductions), scent-marking pairs und territorial single wolves.

In Germany, all wolf monitoring tasks are coordinated und conducted by the 16 commonwealth states, following the German security standards weil das large carnivores (Reinhardt et al. 2015), where genetic analyses constitute a crucial part. Searching weil das genetic samples such as scats or urine traces ist usually conducted on regional scales von a network of trained persons coordinated von the responsible State umwelt Agencies. early out to ns decentralized neighborhood responsibilities, surveillance intensity und strategies zum sample arsenal vary an space and time. However, genetic samples are routinely collected an all federal states with sometimes or regular wolf presence. As ns monitoring tasks during the initial phase von wolf recolonization were particularly intense, we assume that all packs to be identified hinweisen least until 2013.

In this study, scats made nach oben most of the genetic zutat used zum monitoring purposes und were generally accumulated all year-round during presence sign surveys. Various other frequently gathered sample species were hair (e.g. From job beds), urine and blood stains during ns pre-oestrus period collected while eye tracking. Saliva traces were gathered from livestock und wild ungulate kills. Tissue samples were built up from ein wolf carcasses und a this from one set des skeletal remains. Blood, hair und saliva samples were gathered from some wolf carcasses an addition kommen sie tissue samples. Blood samples were accumulated from injured wolves. Blood, hair or saliva samples were accumulated from wolves captured zum radio collaring.

Within the erste years of German wolf monitoring, anfangsverdacht genetic analyses to be performed at ns Institute of webocalendar.com Conservation, polish Academy von Sciences, in Krakow, Poland (Reinhardt and Kluth 2007). In 2009, all commonwealth states des Germany agreed zu use the Senckenberg research Institute as die central laboratory zum genetic wolf analyses zu guarantee ns generation von harmonized charme among the federal states. Bei this study, us used ein wolf samples the were gathered throughout Germany (47°16′–55°03′ N and 5°52′–15°02′ E) between January 2003 and April 2016 an the frame of the federal-state-based regional long-term monitoring tasks (Fig. 1). We began from 2005 with ns third und fourth breeding pair (GW006f and GW001m in N; GW012f und GW008m an NO, seen Tables S1 and S2 for details on individual genotype ID und wolf territories, respectively) zu describe ns trends of genetic diversity and inbreeding von the breeding pairs till 2015; bolzen 2000 und 2004 only two successive breeding pairs und a hybridization occasion occurred. The zuerst wolf reproduction was documented in the Muskauer Heide (MH), Saxony, in 2000 (Ansorge et al. 2006). Monitoring and genetic dünn suggest that die female GW023f and the male wolf ‘I’ with lacking genotype formed the zuerst breeding pair (Fig. 2), which reproduced in 2000 and 2001. From 2002 kommen sie 2004, GW023f reproduced with one more male GW064m in MH. Furthermore, a hybridization occasion occurred in 2003 (Reinhardt and Kluth 2007). Microsatellite evaluation revealed that die female GW006f mated with a domestic dog nearby to produziert natal territory. In winter 2003/2004, four mischung pups were ausblüten alive. Two von them were caught und brought into in enclosure, while the other 2 hybrids disappeared. Ns F1-hybridisation event was confirmed von further analyses based upon 93 single-nucleotide polymorphisms (Harmoinen et al. 2020).


Fig. 1: ein wolf study area and sampling localities startseite the whole of Germany und are divided into die 16 federal states (black lines).

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*

Shown are die number des successfully genotyped samples collected between 2003 and April 2016 (blue circles) an 12 federal claims (labelled through grey initials, lakers Tables S1–S3). The smaller map shows ns wolf distribution across Europe bei 2011 through permanent event (dark pink) und sporadic incident (pale pink) according kommen sie Chapron et al. (2014) and the confirmed wolf occurrence in Germany weil das 2015 (red) follow to the Dokumentations- und Beratungsstelle ns Bundes zu Thema wolf (2017).


Fig. 2: Pedigree des reproducing German wolves an the monitoring period 2005–2015 reconstructed from a combination des microsatellite, mtDNA und field data.

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Bold black lines indicate successful reproductions bolzen female (circles) and male (squares) wolves, dual bold schwarze farbe lines highlight breeding des full-siblings, when thin schwarze farbe lines indicate parent–offspring relationships. Four individuals are depicted twice (arrow v dashed grey line). People with known source verpackt carrying haplotype HW01 (unfilled), individuals with unknown source verpackt carrying HW01 (filled through black) and individuals through unknown source verpacktem carrying haplotype HW02 (filled with dark grey). People found tot are crossed out. Individuals with missing genotype room indicated bei light grey with roman numerals. Reproduction pairs in which both breeders were notfall genotyped are not depicted. Framed through a grey dotted line space the erste (GW023f and I from 2000 zu 2001) and the 2nd breeding pair (GW023f and GW064m from 2002 kommen sie 2004) in the Muskauer Heide (MH), i beg your pardon are not included in the more analyses ~ above trends von genetic diversity and inbreeding von breeding pairs until 2015 (see ‘Materials und Methods’).


DNA extraction and genotyping

Data analyses

To determine individual genotypes, the ns package DNAtools (Tvedebrink et al. 2012; Curran and Tvedebrink 2013) an the r programming language (R Core team 2017) was used. Descriptive statistics von microsatellite loci und probabilities des identity to be performed with GenAlEx ver. 6.5 (Peakall and Smouse 2006, 2012). Ns mean number of different alleles über microsatellite locus (Na), number des effective alleles (Ne), it was observed heterozygosity (Ho) und unbiased supposed heterozygosity (He) were calculated, also as die probability des identity (PID) and probability des identity between siblings (PIDsib) (Waits et al. 2001). As ns number of distinct alleles rely on sample size, we additionally calculated allelic richness (Ar) using the rarefaction technique as implemented bei ADZE ver. 1.0 (Szpiech et al. 2008). Tests zum scoring errors caused über stutter peaks, big allelic dropout and the visibility of null alleles were performed in MICRO-CHECKER ver. 2.2.3 (van Oosterhout et al. 2004). CERVUS ver. 3.0.7 (Kalinowski et al. 2007) was used zu calculate the polymorphism information inhalt (PIC) und to produce input files zum the software program GENEPOP ver. 4.2 (Raymond und Rousset 1995; Rousset 2008) kommen sie conduct Hardy–Weinberg Equilibrium (HWE) testing; dememorization number = 5000; number von batches = 1000; number des iterations über batch = 5000. Zu detect sex-biased dispersal based upon microsatellite markers, the powerful mean des the repair assignment index (mAIc) experiment (Goudet et al. 2002) was conducted utilizing the r package hierfstat ver. 0.04–22 (Goudet 2005).

Pedigree reconstruction, genetic diversity und inbreeding

For pack und pedigree reconstruction, genetic säule were an unified with additional information recorded in the frame von the national wolf monitoring, including spatio-temporal charme on, weil das example, wolf occurrence, territories, society status des individuals or evidence of reproductions. As wolves live an families (Mech und Boitani 2003), the assignment von individuals to ns respective package allows zum reconstructing a consistent pedigree across several generations. A ‘breeding pair’ consists of two reproducing adult wolves, if a ‘pack’ zu sein the wolf family comprising the breeding animals und their offspring. The ‘territory’ ist the home variety inhabited and defended über the verpackt (Mech und Boitani 2003). Native all collected DNA samples, the scent-marking individuals and possible pups an the particular territories to be identified and separated native other breeding pairs and pups in adjacent territories. People that were figured out outside des the distribution range, at the range edge or in areas lacking energetic monitoring were checked against die known packs and pairs zum a possible assignment as offspring.

Various computer system programmes space available zum inferring kinship and pedigree reconstruction (Jones et al. 2010; Walling et al. 2010). However, software-supported parentage assignments generally contain mismatches (Walling et al. 2010). Thus, zum accurate und robust pedigree reconstruction, we merged manual analyses of genetic und monitoring dünn additionally supported by parentage assignments des the program nest ver. 2.0.6.4 (Jones und Wang 2010). Male and female breeding systems were set kommen sie ‘polygamous’ and the inbreeding modell was selected. Verpacktem numbers and territories determined in the German security were acquired from ns public wolf database (DBBW 2017, https://dbb-wolf.de/). As wolf pups space born hinweisen the ende of April/beginning von May, a wolf monitoring year starts at the zuerst of May and ends at the ende of April an the adhering to year. Here, we mention ns year bei which ns monitoring year start (e.g. 2005 zum the security year 2005/2006).

Based on the pedigree data, the mean number of alleles (Na), allelic wealth (Ar), observed heterozygosity (Ho) und unbiased expected heterozygosity (He) were calculated for the various years based on ns genotypes von the inferred reproduction pairs to evaluate ns genetic diversity. Pedigree-based inbreeding coefficients (Fp) for the offspring from breeding pairs to be calculated with ns inbreeding function in the r package GeneticsPed ver. 1.40.0 (Gorjanc and Henderson 2007) using the method ‘meuwissen’ (Meuwissen and Luo 1992). For calculating pedigree-based inbreeding coefficients, people with an unknown source verpackt were thought about as unrelated founders or immigrants. We fitted an exponential growth modell to the yearly genotyped reproduction pairs and also to die yearly numbers of breeding bag inferred by the overall wolf monitoring activities and calculated the annual growth von reproductive systems from the modell using ns statistical programming language r (R Core team 2017). For the detection of trends in the hereditary diversity parameters explained above, we performed the Mann–Kendall test (Libiseller and Grimvall 2002) und calculated Sen’s steep (Sen 1968) using the ns package tendency (Pohlert 2018).

To better understand the recolonization process, we characterized three core locations (CORE1, CORE2, CORE3, seen Fig. S1) based on the spatio-temporal charme of all territories des the genotyped reproduction pairs identified betwee 2005 und 2015. We built spatio-temporal networks of territories, starting with the zuerst three territories of wolf packs, which developed further north–west indigenous a substantial distance zu already present territories: Neustadt (N) bei 2005 weil das core area 1, Altengrabow (AG) bei 2009 zum core area 2 und Munster (MU) in 2012 zum core area 3. The remaining regions were assigned to ns respective main point area according to ns smallest spatial und temporal distance kommen sie one of the three predefined main point areas. We created a minimal convex polygon zum each core area based on the constructed networks des our are data. The territory von the UEM verpackt was notfall included as die geographical position und time von pack bildete did notfall allow zum a clear assignment zu a distinctive core area.

In order zu gain insights into ns genetic diversity des the reproducing individuals an the 3 core areas, we calculated ns mean number des alleles (Na), number von effective alleles (Ne), allelic richness (Ar), it was observed heterozygosity (Ho) and unbiased expected heterozygosity (He) weil das the year 2015. Furthermore, we used the in der nähe des maximum-likelihood clustering an approach ‘snapclust’ (Beugin et al. 2018) bei the ns package ADEGENET ver. 2.1.1 (Jombart 2008) to reveal platz genetic patterns in the expanding ein wolf population. The snapclust.choose.k function was used zu identify die optimal number des clusters (K) based on the Bayesian info criterion (Schwarz 1978) weil das the reproducing individuals bei the year 2015.

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Overall comparison of dispersal distances von breeding females und males was conducted with ns Wilcoxon rank-sum test and observed kommen sie expected distribution of the geschlecht ratio was compared with the chi-square test in R. Direct distances bolzen the centres of the natal territory to die territory of the zuerst reproduction to be calculated through the r package GEOSPHERE ver. 1.5–7 (Hijmans et al. 2017) using the ‘Vincenty Ellipsoid’ method. For the analysis von dispersal distances, we tested two subsets, (i) including breeding individuals the stayed und reproduced in their natal territory, and (ii) including only breeding individuals the left their natal territory. Weil das breeders that reproduced an multiple years with die same or different breeding partners, however, only die distance between the zuerst reproduction and the natal territory was considered.