CEwolf consortium

CEwolf currently consists of 14 scientific institutions from eight countries. CEwolf members involve i) the laboratories that are in charge of genetic wolf monitoring in their respective countries, ii) institutions that are centrally involved in genetic sampling and closely cooperate with those central laboratories.

Please email to wildtiergenetik@senckenberg.de if you have general questions regarding the CEwolf consortium.

Austria

University of Veterinary Medicine Vienna, Department of Interdisciplinary Life Sciences

          Steve Smith: steve.smith@vetmeduni.ac.at

          Georg Rauer: Georg.Rauer@vetmeduni.ac.at

Belgium

Research Institute for Nature and Forest Belgium* ¹

          Joachim Mergeay: joachim.mergeay@inbo.be

          Weblink: https://pureportal.inbo.be/nl/persons/joachim-mergeay

University of Liège, Department of Life Sciences* ²

          Johan Michaux: johan.michaux@uliege.be

          Lise-Marie Pigneur: lmpigneur@gmail.com

          Weblink: https://www.uliege.be/cms/c_9054334/en/directory?uid=U016882

Czech Republic

Charles University in Prague, Department of Zoology

          Pavel Hulva: pavel.hulva@natur.cuni.cz

          Kamila Valentová: valentovakamila70@gmail.com

          Weblink: https://www.vertebrata.natur.cuni.cz/products/hulva-pavel/

Czech University of Life Sciences Prague, Department of Animal Sciences

          Barbora Černá Bolfíková: bolfikova@ftz.czu.cz

          Weblink: https://bit.ly/381vShy

Denmark

Aarhus University, Department of Biology*

          Philip Francis Thomsen: pfthomsen@bio.au.dk

          Weblink: https://pure.au.dk/portal/en/persons/philip-francis-thomsen(42e3d0c2-8389-45cd-aa6f-82004d696ccb).html

Aarhus University, Department of Ecoscience

          Peter Sunde: psu@bios.au.dk

          Weblink: https://pure.au.dk/portal/en/persons/peter-sunde(3a9b01f6-a55e-40f0-9f76-9f8bb4d5d466).html

Natural History Museum Aarhus

          Kent Olsen: kent@molslab.dk

          Weblink: https://www.naturhistoriskmuseum.dk/om-museet/medarbejdere/videnskab/kent-olsen

Germany

Senckenberg Research Institute and Natural History Museum Frankfurt, Center for Wildlife Genetics*

          Michelle Müller: wildtiergenetik@senckenberg.de (central contact person)

          Sebastian Collet: sebastian.collet@senckenberg.de

          Berardino Cocchiararo: berardino.cocchiararo@senckenberg.de

          Carsten Nowak: carsten.nowak@senckenberg.de

          Weblink: www.senckenberg.de/naturschutzgenetik

LUPUS – German Institute for Wolf Monitoring and Research

          Gesa Kluth: gesa.kluth@lupus-institut.de

          Ilka Reinhardt: ilka.reinhardt@lupus-institut.de

Luxembourg

Administration de la nature et des forêts

          Laurent Schley: Laurent.schley@anf.etat.lu

          Weblink: www.emwelt.lu

The Netherlands

Wageningen Environmental Research, Animal Ecology Team*

          Arjen deGroot: g.a.degroot@wur.nl

          Hugh Jansman: hugh.jansman@wur.nl

          Weblink: www.wageningenur.nl/wolven

Poland

University of Warsaw, Faculty of Biology, Department of Ecology*

          Robert Myslajek: robert.myslajek@gmaile.com

          Weblink: www.biol.uw.edu.pl

University of Gdańsk, Department of Vertebrate Ecology and Zoology

          Maciej Szewczyk: maciej.szewczyk@ug.edu.pl

          Weblink: https://en.biology.ug.edu.pl/

Association for Nature “Wolf”

          Sabina Nowak: sabina.nowak@polskiwilk.org.pl

          Weblink: www.polskiwilk.org.pl

 

* Laboratory in charge of wolf genetics in the respective country
*¹ Reference laboratory for the Flemish part of Belgium
*² Reference laboratory for the Walloon part of Belgium

All CEwolf laboratories use common marker systems and share protocols. This standardization of methods is the basis for a common genetic wolf monitoring across central Europe.

A – Microsatellites (STR)

Like in most other regions of wolf distribution, microsatellite markers are the basis of wolf monitoring in the CE region. All CE wolf members apply a set of 13 nuclear microsatellite markers plus two sex-linked markers: CPH5 (Fredholm and Winterø 1995); FH2001, FH2010, FH2017, FH2054, FH2087, FH2088, FH2097, FH2137, FH2140 and FH2161 (Francisco et al. 1996); vWF (Shibuya et al. 1994); PEZ17 (Neff et al. 1999); DBX6 and DBY7 (Seddon et al. 2005). This panel of microsatellites allows to discriminate individuals, to assign samples to packs of origin and reference populations and to identify recent wolf-dog hybrids. Besides this core marker set, several CEwolf labs add other microsatellites in order to increase resolution. However, only the 13+2 marker core set is harmonized across all CEwolf labs.

B – Mitochondrial control region haplotype sequencing

Besides using microsatellites we sequence a hypervariable part of the mitochondrial control region (CR). This region provides important additional information regarding wolf dog discrimination and population origin. One main advantage is that mitochondrial DNA tends to be easier to analyze in samples with very low DNA quantity and quality. As genetic wolf monitoring heavily relies on the use of noninvasively collected samples, such as scats, urine, bones, or saliva traces from kills, wolf identification often relies on this mitochondrial marker.

All CEwolf members apply the identical markers to sequence a part of the control region, which allows for a standardized assignment of wolf haplotypes: Primers WdloopL and WdloopH (Caniglia et al. 2013) are specifically used for detecting the genus Canis on saliva samples from kills, while primers L15995 (Taberlet et al. 1994) / H16498 (Fumagalli et al. 1996) serve for general mammal species determination.

C – SNPs for hybrid detection and other methods

The two above-mentioned methods provide the backbone for genetic wolf monitoring in central Europe. Besides, other approaches are applied, including the use of additional microsatellite markers by some CEwolf labs. When aiming to test for signals of hybridization between wolves and dogs, however, microsatellites have certain drawbacks, including their high resolution and resulting sensitivity towards intraspecific genetic substructure, which may bias results when aiming for assessing wolf-dog-hybridization. We thus developed an alternative approach based on the use of SNPs selected for maximum discrimination between wolves and dogs, which provides accurate assignments to hybrid classes until the 3^rd backcross generation (Harmoinen et al. in review).  The method uses with noninvasively collected samples and allows for precise assessment of recent hybridization in the entire CEP.

Other methods are occasionally applied in the frame of research and monitoring, such as the determination of Amylase copy number variation (in cooperation with Steve Smith/VedMed University Vienna) or whole genome sequencing.

Caniglia R, Fabbri E, Mastrogiuseppe L, Randi E, 2012. Who is who? Identification of livestock predators using   forensic genetic approaches. Forensic Science International: Genetics 7, 397–404.

Francisco LV, Langsten AA, Mellersh CS, Neal CL, Ostrander EA,1996. A class of highly polymorphic tetranucleotide repeats for canine genetic mapping. Mammalian Genome 7, 359-362.

Fredholm M, Winterø AK, 1995. Variation of short tandem repeats within and between species belonging to the Canidae family. Mammalian Genome 6, 11-18.

Fumagalli L, Taberlet P, Favre L, Hausser J, 1996. Origin and evolution of homologous repeated sequences in the mitochondrial DNA control region of shrews. Molecular Biology and Evolution 13, 31-46.

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Pun KM, Albrecht C, Castella V, Fumagalli L, 2009. Species identification in mammals from mixed biological samples based on mitochondrial DNA control region length polymorphism. Electrophoresis 30, 1008-1014.

Seddon JM, 2005. Canid-specific primers for molecular sexing using tissue or non-invasive samples. Conservation Genetics 6, 147-149.

Shibuya H, Collins BK, Huang TH-M, Johnson and GS, 1994. A polymorphic (AGGAAT)n tandem repeat in an intron of the canine von Willebrand factor gene. Animal Genetics 25, 122.

Taberlet P, Bouvet J, 1994. Mitochondrial DNA polymorphism, phylogeography, and conservation genetics of the brown bear Ursus arctos in Europe. Proceedings of The Royal Society B – Biological Sciences 255, 195-200.

I. Origin of wolves in the CE region

Until the late 1990s wolves were extinct in western Central Europe. Comparisons of genetic similarity shows that the wolves occurring in Central Europe derive from Northwestern Poland (Szewczyk et al. 2019). Establishment of the new Central European wolf population (CEP) happened in the late 1990s with initial wolf detections in Saxony and lower Silesia, with the first wolf pack established on German military area near the German-Polish border (Reinhardt & Kluth 2007). Since then, the population has increased initially within the Lausitz and Lower Silesian Region, and showed an increased spread across large parts of the German and Western Polish flatlands during the past decade. In recent years the first territorial wolves have been as well detected in Czech Republic (2014), Denmark (2013), The Netherlands (2019) and Belgium (2018). The population has remained largely isolated since then, but occasional migrants from nearby populations (Alpine, Baltic, Dinaric, Carpathian) have been registered in the frame of CE-wide concerted genetic wolf monitoring.

II. Population separation

Likely due to a strong initial founder effect, CE wolves show a distinct genetic composition based on microsatellites and as well as mitochondrial haplotype frequencies that are distinct from the Baltic founder population. While there is occasional admixture with nearby wolf populations, the CEP has remained its genetic uniqueness until now (Szewczyk et al. 2019).

III. Hybridization

Domestic dogs are reproductively compatible with their wolf ancestors and hybrids are fully fertile. While genome-wide studies commonly find genetic traces of dogs within wolf genomes, there is relatively sparse evidence for recent hybridization events in most European regions. Although a comprehensive European-wide study on hybridization is still lacking, numerous genetic studies have revealed hybridization rates in different European regions. With the exception of few regions, such as the Tuscany (Bassi et al. 2017), hybrids within wild wolf populations occur at rather low rates. Within the CEP we have detected only very few incidences of hybridization, despite of >2500 individual wolf genotypes generated by our consortium. By October 2020, 8 hybrid litters have been detected within the CE region (Czech Republic: 1, Germany: 3, Poland: 5). In Germany, for instance, hybrid pups were removed from the wild and there is currently no evidence of hybrids which have successfully backcrossed into the wolf population. Despite of the low rate of hybridization between wolves and domestic dogs in the CE region, we urge to continue intense genetic monitoring to ensure the detection of hybrids at an early stage and prevent ongoing introgression of dog DNA into the wolf population.

IV. Information on wolf haplotypes in Central Europe

Naming of mitochondrial control region haplotypes follows Pilot et al. (2010). Currently, most CE wolves carry haplotype HW01, a haplotype commonly found across Eastern and Northern Europe, followed by HW02 and, more rarely, HW03, HW06, W17 (Montana et al. 2017) and HW22, the latter one being characteristic of wolves from the Alps or Italy.

Bassi et al. (2017) Trophic overlap between wolves and free-ranging wolf × dog hybrids in the Apennine Mountains, Italy. Global Ecology and Conservation 9, 39–49

Hulva et al. (2018) Wolves at the crossroad: fission-fusion range biogeography in the Western Carpathians and Central Europe. Diversity and Distributions 24, 179–192

Montana et al. (2017) Combining phylogenetic and demographic inferences to assess the origin of the genetic diversity in an isolated wolf population. PLoS ONE 12, e0176560

Pilot et al. (2010) Phylogeographic history of grey wolves in Europe. BMC Evolutionary Biology 10, 104.

Szewczyk et al. (2019) Dynamic range expansion leads to establishment of a new, genetically distinct wolf population in Central Europe. Scientific Reports 9, 19003

The ongoing return of wolves to central Europe raises mixed emotions and leads to often heated debates among involved stakeholders, including animal welfarists, conservationists, hunters, journalists, livestock farmers and politicians. Notably, a considerable amount of misinformation on wolves can be found in all kinds of media, including print, internet, TV, or social media. Some fake news and alternative facts appear to be placed intentionally to heat up the ongoing debate and raise refusal and fear concerning wolves. Similar to other environmental debates, such as climate change, scientific institutions aiming at producing solid data get discredited when they provide information which is disproving those claims. In the case of the wolf, several misinformation is circling around the genetic evidence provided by CEwolf labs and other European laboratories. Here we provide examples of common misinformation regarding wolf genetics in Central Europe as well as a brief fact check. Interestingly, similar or identical claims exist in most countries where wolves have recently spread, including Finland, France, Italy, Germany, Norway, Sweden and Switzerland.

• Hybrids: “All or most European wolves are wolf-dog hybrids and should be removed for the sake of conserving the true wolf!”

Our data based on different up-to date methodologies proofs that hybridization between wolves and dogs occurs only rarely in central Europe. Several thousand individual genetic profiles analysed by CEwolf labs resulted in only a handful of hybrid occurrences. This finding is not surprising, given similar findings in other regions (e.g. Dufresnes at al. 2019 for Alpine region). In some regions of Southern and Eastern Europe, where large feral dog populations exist and effective wolf management is not always guaranteed, locally increased rates of admixed wolves can be found (e.g. Bassi et al. 2017). It should be noted that there is currently no scientific proof that wolf-dog-hybrids raised in the wild are more dangerous or have an increased propensity of showing bold behaviour compared than purebred wolves.

• Human transport: “Wolves did not actively recolonize Central Europe, they were brought back from captivity or Eastern Europe by secretly operating environmentalists.”

Wolves have started to recolonize western Poland and Germany in the late 1990s from Northeastern Poland through long-distance dispersal. This is confirmed by extensive genetic studies (Czarnomska et al. 2013; Szewczyk et al. 2019). Long-distance dispersal over distances of several hundreds to >>1000 km is a common phenomenon in wolves (Kojola et al. 2006).
Ongoing westwards range expansion can be observed for a range of mammal species, such as golden jackals and moose. This phenomenon is best explained, among other factors, by decreased hunting pressure and increasing population sizes within the source regions.

• Hide data: “There are many more wolves and wolf-caused damages than scientists and authorities admit.”

Scientists and environmental authorities work together to provide an effective wolf monitoring based on scientific data. All CEwolf labs are based in universities or large, public-funded research institutions and provide data generated by high scientific standards. Neither is there an interest to hide data on wolf occurrences nor are results of genetic analyses pre-selected for public information.

• Wrong methods: “Scientists use wrong, outdated methods in wolf genetics, thus their findings cannot be trusted.”

CEwolf labs use up-to-date methodologies which are commonly used among genetic laboratories worldwide. Our consortium is led by leading scientists in population and wildlife genetics and we conduct numerous scientific studies on wolf genetics, involving students and scientists from various academic institutions. Besides applying standard genetic tools, our research comprises sophisticated state-of-the-art approaches such as whole-genome-sequencing. For instance, CEwolf is involved in the sequencing >100 complete wolf genomes from central Europe in cooperation with Prof. Tom Gilbert, University of Copenhagen.

• Neutrality: “Scientific institutions are funded by pro wolf NGOs and thus not neutral.”

Analyses done by CEwolf labs come from two main sources: i) environmental ministries, ii) institutional or external research funds. All CEwolf labs are based in universities or large, public-funded research institutions and provide data generated by high scientific standards. Scientific neutrality and honesty are main principle of Good Scientific Practice. CEwolf labs are bound to strict institutional rules of scientific ethical standards.

Bassi et al. (2017) Trophic overlap between wolves and free-ranging wolf × dog hybrids in the Apennine Mountains, Italy. Global Ecology and Conservation 9, 39–49

Czarnomska et al. (2013) Concordant mitochondrial and microsatellite DNA structuring between Polish lowland and Carpathian Mountain wolves. Conservation Genetics 14, 573–588

Dufresnes et al. (2019) Two decades of non-invasive genetic monitoring of the grey wolves recolonizing the Alps support very limited dog introgression. Scientific Reports 9,148

Kojola et al. (2006) Dispersal in an Expanding Wolf Population in Finland. Journal of Mammalogy 87,281–286

Szewczyk et al. (2019) Dynamic range expansion leads to establishment of a new, genetically distinct wolf population in Central Europe. Scientific Reports 9, 19003

Andersen LW, Harms V, Caniglia R, Czarnomska SD, Fabbri E, Jędrzejewska B, Kluth G, Madsen AB, Nowak C, Pertoldi C, Randi E, Reinhardt I, Stronen AV (2015) Long-distance dispersal of a wolf, Canis lupus, in Northwestern Europe. Mammal Research 60: 163-168.

de Groot GA, Nowak C, Skrbinšek T, Andersen L, Aspi J, Fumagalli L, Godinho R, Harms, V, Jansman HAH, Liberg O, Marucco F, Mysłajek RW, Nowak S, Pilot M, Randi E, Reinhardt I, Śmietana W, Szewczyk M, Taberlet P, Vilà C, Muñoz-Fuentes V (2016) Decades of population genetic research call for harmonization of molecular markers: the grey wolf, Canis lupus, as a case study. Mammal Review 46: 44-59.

Caniglia R, Fabbri E, Hulva P, Černá Bolfíková B, Jindřichová M, Stronen AV, Dykyy I, Camatta A, Carnier P, Randi E, Galaverni M (2018) Wolf outside, dog inside? The genomic make-up of the Czechoslovakian Wolfdog. BMC Genomics 19: 533.

Diserens TA, Borowik T, Nowak S, Szewczyk M, Niedźwiecka N, Mysłajek RW. (2017) Deficiencies in Natura 2000 for protecting recovering large carnivores: A spotlight on the wolf Canis lupus in Poland. PLoS ONE 12(9): e0184144.

Donfrancesco V, Ciucci P, Salvatori V, Benson D, Andersen LW, Bassi E, Blanco Juan C, Boitani L, Caniglia R, Canu A, Capitani C, Chapron G, Czarnomska SD, Fabbri E, Galaverni M, Galov A, Gimenez O, Godinho R, Greco C, Hindrikson M, Huber D, Hulva P et al. (2019) Unravelling the Scientific Debate on How to Address Wolf-Dog Hybridization in Europe. Frontiers in Ecology and Evolution 7: 175. 

Harms V, Nowak C, Carl S, Munoz-Fuentes V (2015) Experimental evaluation of genetic predator identification from saliva traces on wildlife kills. Journal of Mammalogy 96: 138-143.

Hatlauf, J., F. Böcker, L. Wirk, S. Collet, L. Schley, L. Szabó, K. Hackländer & M. Heltai (2021) Jackal in hide: detection dogs show first success in the quest for golden jackal (Canis aureus) scats. Mammal Research 66: 227-236.

Hindrikson M, Remm J, Pilot M, Godinho R, Stronen AV, Baltrūnaité L, Czarnomska SD, Leonard JA, Randi E, Nowak C, Åkesson M, López-Bao JV, Álvares F, Llaneza L, Echegaray J, Vilà C, Ozolins J, Rungis D, Aspi J, Paule L, Skrbinšek T, Saarma U (2017) Wolf population genetics in Europe: a systematic review, meta-analysis and suggestions for conservation and management. Biological Reviews 92: 1601-1629.

Hulva P, Černá Bolfíková B, Woznicová V, Jindřichová M, Benešová M, Mysłajek RW, Nowak S, Szewczyk M, Niedźwiecka N, Figura M, Hájková A, Sándor AD, Zyka V, Romportl D, Kutal M, Finďo S, Antal V. (2018) Wolves at the crossroad: fission-fusion range biogeography in the Western Carpathians and Central Europe. Diversity and Distributions 24: 179-192.

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Juránková J, Hulva P, Černá Bolfíková B, Hrazdilová K, Frgelecová L, Daněk O, Modrý D (2021) Identification of tapeworm species in genetically characterised grey wolves recolonising Central Europe. Acta Parasitologica: 1-5.

Karssene Y, Chammem M, Nowak C, Habib Yahyaoui M, de Smet K, Castro D, Lopes S, Muñoz-Fuentes V, Cocchiararo B, Kless D, Van Der Leer P, Khorchani T, Nouira S, Godinho R (2018) Noninvasive genetic assessment provides evidence of extensive gene flow and high movement ability in the African golden wolf. Mammalian Biology 92: 94-101.

Kraus RHS, vonHoldt B, Cocchiararo B, Harms V, Bayerl H, Kühn R, Förster DW, Fickel J, Roos C, Nowak C (2015) A single-nucleotide polymorphism-based approach for rapid and cost-effective genetic wolf monitoring in Europe based on non-invasively collected samples. Molecular Ecology Resources 15: 295-305.

Lesniak I, Heckmann I, Heitlinger E, Szentiks CA, Nowak C, Harms V, Jarausch A, Reinhardt I, Kluth G, Hofer H, Krone O (2017) Endoparasite richness and diversity increase with population size and change with individual age in a recolonising expanding large carnivore population. Scientific Reports 7: 41730.

Montana L, Caniglia R, Galaverni M, Fabbri E, Bolfíková B, Czarnomska S, Galov A, Hindrikson M, Hulva P et al. (2017) Combining phylogenetic and demographic inferences to assess the extent and origin of the genetic diversity in an isolated wolf population. PLOS ONE 12: e0176560.

Mysłajek RW, Tracz M, Tracz M, Tomczak P, Szewczyk M, Niedźwiecka N, Nowak S. (2018) Spatial organization in wolves Canis lupus recolonizing north-west Poland: large territories at low population density. Mammalian Biology 92: 37-44.

Neradilova S, Connell L, Hulva P, Bolfikova BC (2019) Tracing genetic resurrection of pointing dog breeds: Cesky Fousek as both survivor and rescuer. PLOS ONE 14: e0221418.

Nowak S, Mysłajek RW, Szewczyk M, Tomczak P, Borowik T, Jędrzejewska B. (2017) Sedentary but not dispersing wolves Canis lupus recolonizing western Poland (2001–2016) conform to the predictions of a habitat suitability model. Diversity and Distributions 23: 1353–1364.

Reinhardt I, Kluth G, Nowak C, Szentiks C, Krone O, Ansorge H, Müller T (2019) Military training areas facilitate the re-colonization of wolves in Germany. Conservation Letters: e12635.

Salvatori, V., V. Donfrancesco, A. Trouwborst, L. Boitani, J.D.C. Linnell, F. Alvares, M. Åkesson, V. Balys, J.C. Blanco, S. Chiriac, D. Cirovic, C. Groff, M. Guinot-Ghestem, D. Huber, I. Kojola, J. Kusak, M. Kutal, Y. Iliopulos, O. Ionescu, A. Majic Skrbinsek, P. Mannil, F. Marucco, D. Melovski, R.W. Mysłajek, S. Nowak, J. Ozolins, G. Rauer, I. Reinhardt, R. Rigg, L. Schley, T. Skrbinsek, L. Svensson, A. Trajce, I. Trbojevic, E. Tzingarska, M. von Arx, & P. Ciucci (2020) European agreements for nature conservation need to explicitly address wolf-dog hybridization. Biological Conservation 248: 108525.

Schley, L, Jacobs M, Collet S, Kristiansen A, Herr J (2021) First wolves in Luxembourg since 1893, originating from the Alpine and Central European populations. Mammalia 85 (3): 193-197.

Smetanová M, Bolfíková BČ, Randi E, Caniglia R, Fabbri E, Galaverni M, Kutal M, Hulva P (2015) From wolves to dogs, and back: Genetic composition of the Czechoslovakian wolfdog. PLOS ONE 10: e0143807.

Stępniak KM, Szewczyk M, Niedźwiecka N, Mysłajek RW (2020) Scent marking in wolves Canis lupus inhabiting managed lowland forests in Poland. Mammal Research 65: 629-638.

Stronen AV, Iacolina L, Pertoldi C, Kusza S, Hulva P, Dykyy I, Kojola I, Faurby S (2019) The use of museum skins for genomic analyses of temporal genetic diversity in wild species. Conservation Genetics Resources 11: 499–503.

Sunde P, Collet S, Nowak C, Thomsen PF, Hansen MM, Schulz B, Matzen J, Michler FU, Vedel-Smith C & Olsen K (2021) Where have all the young wolves gone? Traffic and cryptic mortality create a wolf population sink in Denmark and northernmost Germany. Conservation Letters, published online.

Szewczyk M, Nowak S, Niedźwiecka N, Hulva P, Špinkytė-Bačkaitienė R, Demjanovičová K, Bolfíková BČ, Antal V, Fenchuk V, Figura M, Tomczak P, Stachyra P, Stępniak KM, Zwijacz-Kozica T, Mysłajek RW (2019) Dynamic range expansion leads to establishment of a new, genetically distinct wolf population in Central Europe. Scientific Reports 9 (1): 19003.

Von Thaden A, Cocchiararo B, Jarausch A, Jüngling H, Karamanlidis AA, Tiesmeyer A, Nowak C^*, Munoz-Fuentes V^* (2017) Assessing SNP genotyping of noninvasively collected wildlife samples using microfluidic arrays. Scientific Reports 7: 10768.