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Fascinating Life Sciences Friederike Range Sarah Marshall-Pescini Wolves and Dogs between Myth and Science Fascinating Life Sciences Thisinterdisciplinaryseriesbringstogetherthemostessentialandcaptivatingtopics in the life sciences. They range from the plant sciences to zoology, from the microbiome to macrobiome, and from basic biology to biotechnology. The series not only highlights fascinating research; it also discusses major challenges associatedwiththelifesciencesandrelateddisciplinesandoutlinesfutureresearch directions. Individual volumes provide in-depth information, are richly illustrated withphotographs,illustrations,andmaps,andfeaturesuggestionsforfurtherreading orglossarieswhereappropriate. Interested researchers in all areas of the life sciences, as well as biology enthusiasts,willfindtheseries’interdisciplinaryfocusandhighlyreadablevolumes especiallyappealing. (cid:129) Friederike Range Sarah Marshall-Pescini Wolves and Dogs between Myth and Science FriederikeRange SarahMarshall-Pescini DomesticationLab,KonradLorenz DomesticationLab,KonradLorenz InstituteofEthology InstituteofEthology UniversityofVeterinaryMedicine UniversityofVeterinaryMedicine Vienna Vienna Vienna,Austria Vienna,Austria ISSN2509-6745 ISSN2509-6753 (electronic) FascinatingLifeSciences ISBN978-3-030-98410-6 ISBN978-3-030-98411-3 (eBook) https://doi.org/10.1007/978-3-030-98411-3 #SpringerNatureSwitzerlandAG2022 Thisworkissubjecttocopyright.AllrightsarereservedbythePublisher,whetherthewholeorpartofthe materialisconcerned,specificallytherightsoftranslation,reprinting,reuseofillustrations,recitation, broadcasting,reproductiononmicrofilmsorinanyotherphysicalway,andtransmissionorinformation storageandretrieval,electronicadaptation,computersoftware,orbysimilarordissimilarmethodology nowknownorhereafterdeveloped. Theuseofgeneraldescriptivenames,registerednames,trademarks,servicemarks,etc.inthispublication doesnotimply,evenintheabsenceofaspecificstatement,thatsuchnamesareexemptfromtherelevant protectivelawsandregulationsandthereforefreeforgeneraluse. The publisher, the authors and the editors are safe to assume that the advice and information in this bookarebelievedtobetrueandaccurateatthedateofpublication.Neitherthepublishernortheauthorsor theeditorsgiveawarranty,expressedorimplied,withrespecttothematerialcontainedhereinorforany errorsoromissionsthatmayhavebeenmade.Thepublisherremainsneutralwithregardtojurisdictional claimsinpublishedmapsandinstitutionalaffiliations. Frontcoverphoto:#RooobertBayer(www.rooobert.at) ThisSpringerimprintispublishedbytheregisteredcompanySpringerNatureSwitzerlandAG Theregisteredcompanyaddressis:Gewerbestrasse11,6330Cham,Switzerland Dedicated to our families (the two- and four- legged ones) Foreword Wolves and dogs—what a pair of species! Everybody knows them—at least most people have an opinion about them. And they are described and referred to in so manydifferentways.Oneisthewild,thebrave,thefearless,theclever,theselfish,or the greedy, while the other is rather tender, withdrawn, obedient, sharing, or kind, perhapsafriendofus.Sosomethinglikeyinandyangorblackandwhite? All this may sound familiar to people, especially those living in Europe, who grew uphearingthetalesbythe Grimm brothers, such asLittle Red Riding Hood. Butthereisalsoanotherstorytotellinwhichthesamewolvesareadmiredbypeople for their spirit and pure nature, and are regarded by many historic tribes as their ancestors.ThiscultureofwolveswasalsoresurrectedbytheRomansclaimingthata she-wolfraisedthetwofounders,RomulusandRemus,ofthecityRome. So,whatcouldbeagreaterchallengethantowriteabookaboutwolves,probably oneofthemostmythicalanimalsinhumanculture.Nottomentionthattherehave been only a handful of scientists who dared to step on this road. Friederike Range andSarahMarshall-Pesciniarejustattherightplaceattherighttimetofollowthis traditiontoupdatewhatonemaycallthe“ethologyandecologyofwolves”.Thereis no place here to list all the books on wolves that have dealt with their life history, behaviour,andevolution.Manyearlierworksarementionedinthisbook.However, thanks to the advancements of the last 20–30 years, and our growing knowledge about wolves, the content of this book is a very significant addition to the earlier publishedmonographs,forexample,bybiologistsErikZimen(1971),DavidMech (1981),andMechandBoitani(2010). Wolvesarealwaysatopicfordiscussion.Inanottoodistantpast,therewasthe worrythatwolveswouldbepushedtothebrinkofextinction,atleastinEuropeand the Americas, but their populations were also decreasing elsewhere. But thanks to conservationists,theirfatehaschangedduringthelast20–30years,andtheyarenow re-colonising many previously abandoned habitats both in Europe and in North America. Although the re-introduction of wolves to Yellowstone was heatedly debated, their showing up in many regions had a positive effect on the ecosystem. Wolfpopulationsrecoveredevenintheabsenceofactivehumaninvolvement;aban on hunting and providing an area with little disturbance appears to be largely sufficient. International students who visited our department were always asked to giveashortpresentationabouttheirwork.Manyyearsago,ayoungDutchstudent vviiii viii Foreword talkedaboutthemeasuresthatmaybeusefultointroducewolvesintheNetherlands. Wealllistenedtoherverypolitely,butafterthetalkwecouldnotresistcommenting: “Butthere arenowolvesintheNetherlands!”She wasprepared for sucharemark andrepliedveryfirmly:“Yes,buttheywillcome.”Andshewasright!In2019the mediareportedseveralsightingsofwolvesinthiscountry. So now we have achieved what we had wanted. Are we satisfied and happy? Well,therespectedandsacredwolvescanendupquicklyonthenegativeendofthe news,ifwestarttobelieveoractuallyfindoutthattheyarealsokillingourlivestock. Sonowweallhavetolearnhowtosharetheland.Weshouldallowthewolvesto establishstablepopulationswhichhelptobalancetheecosystem,butwealsohaveto ensure that humans and their livestock are protected from attacks from this top predator. Friederike Range and Sarah Marshall-Pescini are not afraid of describing and analysingatleast9differentdomestication hypotheses. Thereistherumourthatat the end of the nineteenth century the French academy “banned” the publication of newhypothesesontheemergenceofhumanlanguage.Itseemsthatmanypeoplein thecommunityofwolfanddogresearchenjoy comingupwith “THE”domestication hypothesis that explains “everything” about the history of our animal friends. However,ifoneappreciatesthecomplexityofevolutionaryprocesses,mechanisms ofselection,andtheinterwovennatureofgeneticandenvironmentalfactors,thenit turnsoutthatmostsuchproposalscanexplainonlyasmallpartofthechangesinthe variation of traits that gave raise to modern dogs. So, although in most of those hypotheses there is likely some “truth”, the problem is that we do not necessarily knowwhereitactuallyis.Themainissueisthatwedonothavethenecessarytools to provide experimental evidence for a specific and unique series of events that occurred in the distant past. This situation can be traced in the scientific adventure towards the understanding of where and when dogs were domesticated. Since the first paper using genetic data to establish the time frame and the location of dog domestication(Vilaetal1997),theevidenceforaspecifictimeintervalandlocation changedfromkeypapertokeypaper.Andoneimportantlimitationofthisapproach (probablyunlikelytobesolvedinthenearfuture)isthatitisimpossibletosavevery ancient DNA. Moreover, most of these hypotheses are rather complementary than exclusive; they look at different levels of biological organisation. For example, possibleselectionforgenesthatmaymakedogslooksoyoung(seeKinderschema, Lorenz) in the eyes of their human friends may not exclude earlier/parallel/later selection for more flexible agonistic behaviour at older age. Pleitropy and polyge- netic effects are also likely to make the unveiling of an evolutionary process very difficult. I,personally,totallyagreewiththeauthors’preferencetoregardwolvesanddogs astwospecies,buttalkingabout“wolves”and“dogs”inanevolutionaryscenario,as it often happens, is a major oversimplification. “Putting things into boxes” is a typical human way of thinking, probably based on our bias to think in linguistic terms. But in reality there are (were?) at least 10–14 subspecies of wolves ranging fromSaudiArabiatoAlaska,andtherearemorethan400breedsofdogsdispersed Foreword ix overtheentireglobe,nottomentionthedingoesandthemillionsofstrayandferal dogs. So when scientists seek to explain the emergence of differences in certain phenotypictraits,theyoftenimaginethetwospeciesasextremes,correspondingto the adjectives above that typically characterise “wolves” and “dogs”, but in reality therearemanywolf-likedogsandalsodog-likewolfpopulations.Ithinkweshould takebothwithinandbetweenphenotypicvariationinthesespeciesseriously. On the more positive side there is also some good news, when it comes to our much improved knowledge about the social life of wolves. Extensive field observations of wolf populations and individual groups are now complemented with observations on social structure, migration of individuals, and intra- and intergroup competition for resources. And the major discovery, attributed to Jane Packard,isalsoquiterecent:typicalwolvesliveinfamilies;thatisafather,amother, andtheiroffspringmakeupagroup.Sotheoldideaofthewolfpack,inwhichall individuals fight for being the leader, can be thrown into the garbage heap of outdatedscientificideas.Nevertheless,thisdiscoveryfindsitsplaceonlyslowlyin themindofpeopleasmanystillrefertowolfpacksinsteadoffamilies.Somebody mayask,whatisthedifference?Well,alot.Ifthefamilyisthesocialunitinwolves, theircooperationatvarioustasksandchallengesislessofasurprise.Theparentscan realisetheirfitnessonlyiftheyareabletoraiseasmanyyounganimalsaspossible, and the offspring can also gain a lot from cooperating with their parents. The collaboration among siblings can be explained on similar grounds. Ifcollaboration inwolvesisbasedonkinselection,thenactuallythetransitiontoacollaborativedog isactuallynotastraightforwardevolutionaryissuebecausethosekin-relatedgenetic factorsdonotoperatewhenhumansanddogsworktogether. This book is packed with much knowledge on wolves most of which was establishedinthelast20years.Ithinkweall,whoarefascinatedbytheevolution, ecology, and behaviour of wolves and dogs, have to thank those few enthusiastic scientistswhodedicatedapartoftheircareers tofollowthelivesoftheseamazing creatures.FriederikeRangeandSarahMarshall-Pescinideserveafineplaceamong these brave researchers together with Zsófia Virányi, Enikő Kubinyi, and Márta Gácsi,tonameonlyafewothers.Thesciencethatispresentedwithgreatcareinthis book, taking into account different views, perspectives, and lack of knowledge, should invite any readers interested in biology to get a closer look at this species withoutactuallygettingcoldordirty,andtheyalsodonotneedtostandthelookofa wolffromashortdistance. Congratulations,FriederikeandSarah! FamilyDogProject,Departmentof ÁdámMiklósi Ethology,EötvösLorándUniversity Budapest,Hungary 31October2021 Foreword Investigatingtherelationshipsbetweenhumans,wolves,anddogsprovidesamore important mirror to understand ourselves and our relationship with the world, than mostpeoplewouldappreciate.Homosapiensisthemostinvasivespeciesonearth, permanentlyinhabitingeventhemostunlikelyhabitatssuchasdesertsandthehigh Arctic. Dogs arguably contributed a lot to this roaring success. Unplanned and unintended, our ancestors bumped into wolves on their way into the Eurasian continent. It is still a miracle that this happened—and why it happened remains enigmatic. Even more astonishing is that some of these palaeolithic human–wolf associations turned into permanent companionship with domesticated wolves we calldogs.Thisamazingstoryraisesmanyquestions,forexample,howdidhumans andwolvesgetintocontactinthefirstplace?Howcomethatsuchagreatvarietyof dogs originated from a single wolf type? And in what way do modern dogs differ from wolves, i.e. what happened during adapting to live with, or close to, humans duringtheprocessofdomestication? Onewouldassumethatsciencehaslongprovidedmostoftherelevantanswers, butthisisnotthecase.Firstofall,thetopicissoclose(tooclose)tomanypeople, includingscientists,thatevenresearchwas—andstillis—anideologicalminefield. Second,andmostimportantly,onlythemostrecentdevelopmentsinpalaeogenetics, archaeology, experimental behavioural biology, and other disciplines provided the kind of firm ground needed to increasingly allow knowledge to compete with opinionation.Thisbookisamilestoneofknowledge,integratingmorethanadecade ofexperimentalresearchattheWolfScienceCenter(WSC)withtherelevantresults ofcolleaguesworldwide.Asananchoroffactsitwillprovidethegroundforfurther thinking,discussing,anddevelopingthefield. Thecontentsofthisbookevenradiateintoexperimentalarchaeology,enlighten- ing educated guesses as to why wolves and humans may have gotten into contact. The most promising candidate factor is the close match between the ecology, lifestyles, and social organisation of hunter-gatherer humans and wolves. Within- clan and within-pack cooperative orientation of humans and wolves was probably keyinboththeinitialmatchmakingandourcloser-than-evercompanionshipswith dogs. Our present knowledge even contributes arguments towards the early scenarios,makinghand-rearingandarespectfulnon-dominatinghumaninteraction stylemuchmorelikelythanthestillprominentideaof“self-domestication”,holding xxii