Topics in Biodiversity and Conservation F.P.G. Princée Exploring Studbooks for Wildlife Management and Conservation Topics in Biodiversity and Conservation Volume 17 Moreinformationaboutthisseriesathttp://www.springer.com/series/7488 F.P.G. Princée Exploring Studbooks for Wildlife Management and Conservation 123 F.P.G.Princée PopulationManagementandWildlifeConservation EastRudham,Norfolk,UK ISSN1875-1288 ISSN1875-1296 (electronic) TopicsinBiodiversityandConservation ISBN978-3-319-50031-7 ISBN978-3-319-50032-4 (eBook) DOI10.1007/978-3-319-50032-4 LibraryofCongressControlNumber:2016961290 ©SpringerInternationalPublishingAG2016 Thisworkissubjecttocopyright.AllrightsarereservedbythePublisher,whetherthewholeorpartof thematerialisconcerned,specificallytherightsoftranslation,reprinting,reuseofillustrations,recitation, broadcasting,reproductiononmicrofilmsorinanyotherphysicalway,andtransmissionorinformation storageandretrieval,electronicadaptation,computersoftware,orbysimilarordissimilarmethodology nowknownorhereafterdeveloped. Theuseofgeneraldescriptivenames,registerednames,trademarks,servicemarks,etc.inthispublication doesnotimply,evenintheabsenceofaspecificstatement,thatsuchnamesareexemptfromtherelevant protectivelawsandregulationsandthereforefreeforgeneraluse. Thepublisher,theauthorsandtheeditorsaresafetoassumethattheadviceandinformationinthisbook arebelievedtobetrueandaccurateatthedateofpublication.Neitherthepublishernortheauthorsor theeditorsgiveawarranty,expressorimplied,withrespecttothematerialcontainedhereinorforany errorsoromissionsthatmayhavebeenmade. Printedonacid-freepaper ThisSpringerimprintispublishedbySpringerNature TheregisteredcompanyisSpringerInternationalPublishingAG Theregisteredcompanyaddressis:Gewerbestrasse11,6330Cham,Switzerland Preface Some 35 years ago, I became involved in conservation of endangered animal species. It all started while I was a biology student at the Biological Department oftheRoyalRotterdamZoologicalandBotanicalGardens(“BlijdorpZoo”).Itwas around the time that the international zoo world introduced cooperative species management programmes, based on demographic and genetic science, in order to establish self-sustaining populations of endangered species. This created an environmentinwhichappliedandfundamentalresearchwereintertwinedandwhich shapedmyscientificbackgroundinpopulationgenetics. The introduction of personal computers opened a “new” creative platform that allowedmetodeveloptheskillsofcomputerprogramming.Thisresultedindevel- opingsimulationmodelstoexplorepopulationgeneticprocessesinrealpopulations with their complex pedigrees. The first computer program that I developed was followedbythenext,andthenext,etc.resultinginstudbooksoftwarethatincluded arangeofdemographicandgeneticanalyses. The catalyst for writing this book dates back to the years that I worked for the Dutch National Foundation for Research in Zoological Gardens/European Association of Zoos and Aquaria. Teaching, training and advising coordinators of European Endangered Species Programmes (EEPs) in population biology and studbook software meant exposure to a diversity of practical problems in species management. Solutions to some of these problems may be embedded in the studbookitself,andthatiswhereexploringstarts. I participated in various workshops of the IUCN/SSC/Conservation Breeding Specialist Group in which the knowledge of zoo and wildlife professionals was exchanged and combined. These workshops, the Population and Habitat Viability AssessmentoftheArubaislandrattlesnakeinparticular,exposedmetothereality offieldconservation. I became more directly involved in nature conservation projects in the 1990s through the conservation fund of the Dutch zoos. The induction was a population viability study on lions in the Queen Elizabeth National Park (Uganda) that was v vi Preface carriedoutbyMakerereUniversity(Kampala).Individualidentificationoflions,a smallpopulationandurbanisedareasactingasabarriertomovementdonotmake itdifficulttoforeseestudbook-stylemanagementinthefuture. A few years later, I started to write the first chapters while living next to Ranomafana National Park, a rainforest in the south-east of Madagascar. No zoo environment, but wild nature. However, the state of some endangered species in this42,000haprotectedareahighlightedthatnatureis(gradually)becomingabig zoo.Take,forexample,thecloselymonitoredgreaterbamboolemurpopulationin this park at that time. One day there were six animals, the next day one animal was “missing”, and one month later a young was born, bringing the group total backtosix.Thisdoesn’tsoundlikeawildpopulationbutmorecloselyresemblesa zoogroup.Fieldconservationisexploringtoo,asresearchersdiscoveredfournew groups, totalling more than 100 individuals, in fragmented bamboo forests within theperipheralzoneoftheparkin2009and2010. The idea of exploring studbook data for its relevance to conservation in the naturalhabitatwasreinforcedwhileIwaslivinginDzangaSanghaProtectedAreas (Central African Republic). The secretive lifestyle of many of the inhabitants of this tropical rainforest not only makes it difficult to assess population sizes but alsotoobtainnaturalhistorydatathatarerequiredforconservationmeasures.My backgroundpromptedmetoaskzoocolleaguesforstudbookdatainordertoextract basicbiologicalinformationonspeciesintheprotectedarea.IhaveincludedaPVA studyonlowlandbongoasanexampleinthisbook. Exploring studbook data “forced” me to explore statistics in more depth than I had done before. Studbooks are relatively small and the issue of reliability of estimated parameters cannot be ignored. Moreover, problem-solving involves comparative studies that simply require significance tests. However, this is not a bookonstudbookstatistics.Themainfocusisexploringthewealthof(biological) datainstudbooks,inwhichstatisticscanplayasupportingrole. Ihaveselectedtopicsindemographicsandgeneticsofstudbookpopulationsthat deservemoreattention,butwhichcanbestudiedinstandarddatasets.Thesetopics are interconnected, which means that, for example, life tables are used in tests for inbreeding depression and litter size is considered as a “trait” in quantitative genetics. Thisbookisintendedforprofessionalsinzooandwildlifeconservationwhoare involved in management of captive populations; reintroduction or management of small,isolated,fragmentedpopulationsinthewild;andforresearchersandstudents who are planning studies that are based on studbook populations or who want to explorewhatstudbookdatacanofferfortopicsinacademicresearch. EastRudham,UK F.P.G.Princée September,2016 Acknowledgements Amonographistheworkofasingleauthorbutnotofasinglemind.Thisbookisthe resultofexperiencesandideasthathavebeenaccumulatedoveryearsandhavebeen shapedbydiscussionswithcolleagues,fromzookeepertodirectorandfromranger togamewarden,aswellaswithcolleaguesingovernments,NGOsanduniversities. First of all, I would like to thank Anna Feistner for reading this book through severaltimeswithaneditorialeyeandforhercommentsandideasasaconservation biologist. I am indebted to Anna for her encouragement, support and trust in enabling me to realise a book which took considerably more time than I had once optimisticallyforeseen.IthankBertdeBoerforhisvaluablecommentsonanearlier draftandideasthathelpedinstreamliningthisbook. I could not have written this book without access to real-time computerised studbooksonendangeredspecies.Ithankthefollowingspeciesmanagers/studbook keepers and the institutions to whom they are or were affiliated: Richard Barnes (Port Lympne Reserve, UK), Rob Belterman (Rotterdam Zoo, The Netherlands), Leif Blomqvist (Nordic Ark, Sweden, and Helsinki Zoo, Finland), Lydia Frazier Bosley (Logsden, USA), Angela Glatston (Rotterdam Zoo, The Netherlands), GerardMeijer(OuwehandsZoo,TheNetherlands),HannyVerberkmoesandWim Verberkmoes (GaiaZOO, The Netherlands) for providing their studbook data to be used as examples in this book. I acknowledge Andrea Turkalo (the Wildlife ConservationSociety,USA)forfielddataonlowlandbongoinDzangaBaie. I would like to thank Jonathan Ballou, Laurie Bingaman-Lackey, Lisa Faust, Robert Lacy, Nathan Flesness and Andy Odum for discussions, some of which startedyearsago,onvarioustopicsindemography,geneticsandstudbooksoftware. Erik Postma is acknowledged for explaining assumptions that are made regarding missingparentsintheanimalmodel. Ludwig Siefert is acknowledged for introducing me to the reality of field conservationwhenIwasinvolvedinsupportingtheUgandanlion/carnivoreproject. I thank Jean-Claude Razafimahaimodison for all discussions when I exchanged the comfort of analysing zoo populations for interpreting transect data from RanomafanaNationalPark.AngeliqueToddandAndreaTurkaloareacknowledged forintroducingmetotheworldofgorillas,forestelephantsandlowlandbongo. vii Contents PartI Introduction 1 Introduction................................................................. 3 1.1 ABriefHistory...................................................... 3 1.2 FromRegistertoManagement..................................... 4 1.3 PopulationManagementTriangle.................................. 5 1.4 ExploringStudbooks ............................................... 7 1.4.1 NaturalHistory........................................... 7 1.4.2 Census.................................................... 8 1.4.3 LifeTables ............................................... 8 1.4.4 GeneticAnalyses ........................................ 9 1.5 ConservationData .................................................. 10 1.6 MoreTopics......................................................... 11 References.................................................................... 11 2 StudbooksandSoftware................................................... 15 2.1 Introduction ......................................................... 15 2.2 StudbookExamples................................................. 16 2.2.1 RedCrownedCraneGrusjaponensis................... 16 2.2.2 AfricanWildDogLycaonpictus........................ 17 2.2.3 RedPandaAilurusfulgens .............................. 17 2.2.4 EuropeanWolverineGulogulogulo.................... 18 2.2.5 SnowLeopardUnciauncia ............................. 18 2.2.6 CalifornianSeaLionZalophuscalifornianus .......... 19 2.2.7 BongoTragelaphuseurycerus .......................... 19 2.2.8 BlesbokDamalisicuspygargusphillipsi................ 19 2.3 SoftwareTools ...................................................... 20 2.3.1 PopulationManagementLibrary........................ 21 References.................................................................... 22 ix x Contents PartII DemographicAnalyses 3 NaturalHistory............................................................. 27 3.1 Introduction ......................................................... 27 3.2 LifespanandLongevity............................................. 28 3.2.1 EstimatedDates.......................................... 30 3.2.2 Longevity................................................. 31 3.2.3 Pathologist’sView....................................... 34 3.2.4 HistoryandHusbandry.................................. 34 3.2.5 EarlyLifeStages......................................... 36 3.2.6 DifferencesBetweenSexes.............................. 37 3.3 ReproductiveLifespan.............................................. 40 3.3.1 AgeatFirstReproduction............................... 41 3.3.2 Artefacts.................................................. 42 3.4 LitterSize ........................................................... 42 3.4.1 AssessingLitterSize..................................... 43 3.4.2 DateRangeinLitter/ClutchSize........................ 44 3.5 Seasonality .......................................................... 45 3.5.1 MeanDate................................................ 45 3.5.2 SeasonalityinBirths..................................... 46 3.5.3 SeasonalityinDeaths.................................... 51 3.6 Inter–BirthInterval ................................................. 52 3.6.1 ManagementEffects..................................... 53 3.6.2 Egg–LayingSpecies..................................... 55 References.................................................................... 55 4 CensusAnalysis............................................................. 59 4.1 CountingAnimals................................................... 59 4.2 CensusDate......................................................... 61 4.3 Census............................................................... 61 4.4 PopulationGrowth.................................................. 63 4.5 Sex–Ratio ........................................................... 65 References.................................................................... 66 5 Births,DeathsandMigration ............................................. 69 5.1 CensusEvents....................................................... 69 5.2 Births ................................................................ 70 5.2.1 Sex–RatioatBirth ....................................... 73 5.3 Deaths ............................................................... 76 5.4 Migration............................................................ 78 5.4.1 MigrationPatterns ....................................... 79 5.5 BringingitTogether ................................................ 80 References.................................................................... 81